FAUNA NATURA

OPTIMA
MUNDI DUX

Volume 3

Alexander G. Radchenko
Graham W. Elmes

Myrmica ants
(Hymenoptera: Formicidae)

of the Old World

Warszawa 2010
 FAUNA NATURA
OPTIMA

MUNDI DUX

Museum and Institute of Zoology

Polish Academy oí Sciences

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 Myrmica ants (Hymenoptera, Formicidae)
of the Old World

ALEXANDER G. RADCHENKO1 and GRAHAM W. ELMES2

1
Museum and Institute of Zoology Polish Academy of Sciences, Wilcza 64,
00-679 Warszawa, Poland; e-mail: [email protected]
2
CEH Research Fellow, Centre for Ecology and Hydrology, Crowmarsh Gifford,
Wallingford, Oxfordshire, OX10 8BB, United Kingdom; e-mail: [email protected]

ABSTRACT

The "Myrmica ants of the Old World" by A. G. Radchenko and G. W. Elmes,

is the first full taxonomic revision of this important Holarctic ant genus that is
abundant in many biotopes throughout the Palaearctic and parts of the Oriental
Regions. Myrmica ants have important functions in the dynamics of the plant and
animal communities of many temperate habitats, e.g. myrmechory, commensalism
and social parasitism. They are especially renowned for their role in the life cycle
of the endangered species of Large Blue Butterflies (the genus Phengaris, now
senior synonym of Maculinea). Latest European research shows that during the
larval stage most Phengaris populations are obligate social parasites of particular
species, or even particular populations of Myrmica, while in East Asia there are
many more species of Phengaris butterflies whose biology are poorly known, but
probably interact with Myrmica ants. Therefore besides being a rigorous taxonomic
study, this book is also aimed to support the many groups of scientists and nature
conservationists who need to identify Myrmica ants and to place species in their
context both within the genus as currently known and within the evolutionary
history of the genus.
The book opens with a preface section that briefly looks at the mention of Red
Ants (Myrmica rubra) in early literature and outlines the collaboration between
the authors: Alexander Radchenko is one of the leading ant taxonomists of his
generation and has developed a special knowledge of the Myrmica species of the
former Soviet Union; Graham Elmes is- a'well known social insect ecologist who

3
has made special studies of the population dynamics of the genus Myrmica and
conservation problems of Phengaris butterflies. The preface is followed by 5 major
Chapters: it is not intended that these need be read sequentially, rather it is
expected that readers will dip in and out of the various chapters and sections as
required. Chapter 1 is an introduction to the genus, it starts by explaining the
scope and layout of the monograph and gives the reasons why the New World
fauna are not included, the main ones being that much primary taxonomy still
needs to be done on this fauna which shows no significant overlap with that of
the Old World. The main part of Chapter 1 comprises a well-referenced biological
review of the genus Myrmica sufficient to start a detailed literature search. It con-
cludes with photographs and biographies of the 19 myrmecologists who have
described two or more "good" Myrmica species.
The core of the book is Chapter 3 entitled "Review of Species". Here the 426
names that have been ascribed to the genus Myrmica are listed and discussed in
various ways and the 142 extant and five extinct taxa considered to be good Old
World species are highlighted. These are then reviewed taxonomically in alpha-
betical order and for each caste of each species a full set of accurate line drawings
are provided (using the type specimens where possible). The Keys to species are
provided in Chapter 5. Apart from the key to fossil species, the authors give
separate keys to the worker castes of extant species found within seven distinct
biogeographical regions. There is very little overlap between these regions and this
should make the keys less cumbersome for use by less specialist readers.
The remaining two chapters are more specialized and outline the authors'
latest views on the origins, evolution, spread and current distribution of the genus.
Chapter 2 starts by examining the taxonomic position and diagnosis of the genus
Myrmica in relation to other ants. The authors describe their concept of species
groups within the genus and list the main characteristics of the 17 groups that they
currently recognise, and the species therein. Recent molecular-genetic studies sug-
gest that most of these groups have some phylogenetic value. Chapter 4 addresses
the zoogeography and evolution among the species listed in Chapter 3; Radchenko
and Elmes recognise nine major Zoogeographie areas that generally are quite dis-
tinct from each other, the faunas of the two areas within the Oriental Region are
quite distinct from those of the Palaearctic Region. The paucity of fossil Myrmica
ants makes discussion of the timing of the evolution and spread of the genus Myr-
mica very subjective. However, the authors give reasons for supposing that the
genus evolved in ancient Eurasia more than 45 Ma and had spread throughout
Eurasia by the start of the Miocene Epoch, probably invading North America via
the Beringian land bridge several times in subsequent ages.

KEY WORDS

Systematics, ants, Myrmica, taxonomy, revision, identification-keys, biogeo-

graphy, evolution, biological-review, fossil, Palaearctic and Oriental Regions.

4
 CONTENTS

Abstract 3
Preface 7
Chapter 1. General background and biology of Myrmica 13
1.1. Scope of this monograph 13
1.2. The layout and use of this monograph 14
1.3. The biology of Myrmica 16
1.3.1. Myrmica in ecosystems 16
1.3.2. Anatomy and surface chemistry 17
1.3.3. Colony structure 18
1.3.4. Social interactions 20
1.3.5. Physiology 23
1.3.6. Social parasitism 25
1.4. A brief history of the taxonomic investigation of Myrmica in the Old
World ' 27
1.4.1. Pre 1950 27
1.4.2. Post 1950 30
1.5. Morphometries and Myrmica taxonomy 32
1.5.1. History of measurement of Myrmica 32
1.5.2. A brief critique of the value of morphometries 33
1.5.3. Measurements and indices used in this monograph 35
1.6. Biographies of the authors 37
1.6.1. Biographies of the authors of this monograph 37
1.6.2. Biographies of other authors (ordered chronologically) 38
Chapter 2. Taxonomic structure of the genus Myrmica 51
2.1. Taxonomic position and diagnosis of the genus Myrmica 51
2.2. Species group concept in the genus Myrmica 58
Chapter 3. Review of species 69
3.1. Material studied and its deposition 69
3.2. The 303 names definitely attributable to the genus Myrmica 71
3.3. Review of extant Myrmica species of the Old World 79
3.3.1. Review of species 79
3.3.2. Drawings of the details of structure of extant Myrmica species . . . 331
3.4. Fossil species, undoubtedly belonging to the genus Myrmica 653
3.5. Names incertae sedis in the genus Myrmica 657
3:6: Nomina nuda in the genus Myrmica 663
3.7. Unavailable names in the genus Myrmica 663
3.8. List of taxa originally described in the genus Myrmica, but afterward
transferred to other genera or excluded from Myrmica 664
Chapter 4. Zoogeography and evolution of the genus Myrmica 671
4.1. The centres of Myrmica species diversity 672
4.2. Comparison of the regional faunas of Myrmica 678
4.2.1. Diversity between the nine centres 678
4.2.2. Interpretation of the diversity between the centres 686
4.3. Origin and evolution of the genus Myrmica 689
4.3.1. Putative ancestor and estimated time of Myrmica origin 689
5
 4.3.2. Evolution within the genus Myrmica and origin of the species
groups 691
4.3.2.1. Ancestral Myrmica forms and place of their origin 691
4.3.2.2. Origin of the species groups and their probable relation-
ships 696
Chapter 5. Keys for the identification of the Myrmica species of the Old
World 711
5.1. A key for the identification of the Myrmica species of West Europe
and North Africa 713
5.2. A key for the identification of the Myrmica species of East Europe,
West Siberia, northern Kazakhstan, Caucasus, Asia Minor, Turkmeni-
stan and Iran 721
5.3. A key to the identification of the Myrmica species of East Siberia,
Russian Far East, Mongolia, Korean Peninsula, northern China, and
Japan 729
5.4. A key for the identification of the Myrmica species of the Middle
Asian Mountains 734
5.5. A key for the identification of the Myrmica species of the Tibetan
Plateau 735
5.6. A key to the identification of the Myrmica species of the Himalaya and
Afghanistan 736
5.7. A key for the identification of the Myrmica species of South and
South-East Asia 740
5.8. A key for the identification of the fossil Myrmica species 744
References 745
Index of the all Latin names 781
Myrmica names recognized here as good species 781
Other names 785

6
PREFACE

Red Ants - Myrmica

The name Red Ant as a description of reddish myrmicines, especially Myrmica ants,
does not appear in natural philosophical literature until the 17th century. However, it was
probably in common use much earlier because most modern European languages have some
version of the name. The ants that comprise the modern genus Myrmica must surely have
been recognized as a distinct form of animal life throughout most of the human cultural
history of the temperate Northern Hemisphere. They come to people's notice through their
readiness to administer irritating stings, especially in hot weather when it is easy to sit or
lie on an unnoticed nest. When one does so, the angry Myrmica workers have an uncanny
knack of getting under one's clothes where each can sting several times before exhausting
its supply of venom. For most people the result is a short-lived, localized stinging sensation
and, in some cases, small itchy bumps develop that can persist for several days. A few un­
fortunate people can develop severe allergic reactions to the stings. The average Roman sol­
dier, who served in northern Europe pitching camp in meadows and woodland clearings,
must have had some colourful names for Myrmica. During the European "Dark Age" the
tribal peoples who lived close to nature recognized many common insects, and although
we know of no name that refers directly to Myrmica they probably had one, the Saxon
peoples had a variety of words for ant, anthill, ant-nest, sting etc. Even today in England,
Myrmica ants are often called "Picnic Ants" or "Red Stinging Ants" while the name
"European Fire Ant" has been coined recently for the pest populations of Myrmica rubra
(L.) introduced into the USA.
Ants were well known to the classical Greek writers and although they are not discussed
at any length by Aristotle (384-322 ВС) in his "Historia Animalium" it is clear from his
knowledge and treatment of other insect groups that he recognized different sorts of ants.
The Greek understanding of natural history was barely advanced by European writers until
the 17th century. The Roman Pliny the Elder (20-79) re-hashed a lot of Aristotle's work
and compounded it with various myths and legends, while Claudius Aelianus (ca. 175-235)
simply reproduced much of Pliny's work with added errors. There is no description that
can be linked to the genus Myrmica which is hardly surprising given that classical study was
centred on the Mediterranean region where species of Myrmica are rare.
In northern Europe some of the early writers showed considerable knowledge of plants
and animals, including ants, but they were usually more interested in their medicinal
properties rather than their natural history, e.g. the 12th century Abbess Hildegard of
Bingham. The Dutchman, Albertus Magnus (1205-1282) was instrumental in reviving the
classical knowledge of natural philosophy by reiterating much of Aristotle's, Pliny's and
Aelian's works with some additions from Arabian studies. By the 16th century, there was
a sudden surge of interest in natural history with numerous works based on the classic
writers: e.g. the Swiss, Conrad Gesner (1516-1565), and the Bolognian, Ulysses Aldrovandi

7
(1527-1605). Aldrovandi devoted about 30 manuscript pages to ants, including a section
on types of ants, although it added little to the work of his predecessors it nicely
illustrates how by the mid 16th century natural historians were beginning to recognize the
great diversity of insects from different regions of the world. We have no information on the
extent of oriental knowledge of types of ants, but given the highly realistic pictorial depic-
tion of plants and animals that had developed in China by the 16th century it was probably
considerable.
So, it was not until the 17th century that the first formal descriptions of native Euro-
pean ants were made. The British natural historian, philosopher and theologian John Ray
(1628-1705), who some consider to have influenced Linnaeus' work, recognized five
species of ant in Britain (Wheeler 1926, p. 223). The posthumously published "Historia In-
sectorum" (Ray 1710) as Ray always acknowledged, owed a considerable amount to his
pre-deceased friend and colleague Francis Willughby; whichever of them contributed the
section on ants they recognized the following names: Formica maxima hippomyrmex,
Formica media ruja, Formica media nigro colore splendens, Formica minima rubra,
Formica minor ex fusca nigricans, that Wheeler equated to Camponotus herculeanus (L.),
Formica rufa L., Formica fusca L., Myrmica rubra (L.) and Lasius niger (L.) respectively
(though the first can not be C. herculeanus because Camponotus species are not present in
Britain). Some thirty years later (ca. 1743) this classification was well known in Europe.
For example, in an unpublished text the 18th century French natural philosopher Reaumur,
acknowledged great diversity among ants: "Ray n'en observe en angleterre que cinque
espèces; les descriptions qu'il en a faites m'ont mis en etat de juger que les mêmes sont
aussi en france, ou il est facile d'en trouver beaucoup plus d'espèces" (Wheeler 1926).
This then was the background knowledge available to 18th century writers such as the
Rev. William Gould (1747) who published the greatly under-rated volume "An Account of
English Ants", the first book on ants in the English language. Throughout, Gould compared
the biology of five species: the Hill Ant, the Jet Ant, the Red Ant, the common Yellow Ant
and the small Black Ant, that equate to F. rufa-like, F. fusca-like, M. rubra-like, Lasius
flavus-like and L. niger-like species. Gould used much the same morphological features
used earlier by Ray and later by Linnaeus, De Geer and others to define Red Ants. In his
first chapter on the various kinds of ants and their structure, Gould writes (ibid., p. 8)
"The ligament [petiole] that unites the breast [mesosoma] and body [gaster] in the Red
Ant consists of two lobes somewhat round and divided in the middle [petiole and
postpetiole]. A fine thread connects it with the breast, and another with the body. In other
ants their [there] appears but one lobe between the threads; which rises higher and is
broader than the lobes in Red Ants. This species has also at the end of its breast two sharp
prickles [propodeal spines]". This is followed (ibid., p. 9) by "The Red Species has also
a sting of the same contexture as a bee, only in miniature. With this it opens an orifice in
the flesh, and afterwards injects a venom which gives a smart but momentary pain". From
all the subsequent ecological description there is no doubt that in the main, Gould was
thinking of the genus Myrmica when discussing Red Ants.

The origin of our collaboration

Graham Elmes first studied Myrmica ants when he was recruited into the British Nature
Conservancy in 1961, to be an assistant to Michael V. Brian (1919-1990), a leading British

8
ant specialist at that time. Although employed as an ecologist, Brian was an "experimental
biologist" of the "Cambridge School" whose main interest was social biology, trying to un-
derstand how queens and workers interact to regulate factors such as colony growth, worker
oviposition and sexual production. Myrmica rubra was the main "research model" species,
used because its colonies frequently had many queens. This enabled one to make many
small replicated experimental cultures of ants to test the effect of single variables (e.g.
presence or absence of a queen, high and low temperatures or different types of food). While
such experiments have their limitations, they were extremely informative and form a bench-
mark against which many more recent studies fall short. The underlying rationale driving
Brian's team at that time could be summarized as "What we find for M. rubra almost cer-
tainly holds true for all Myrmica ants and probably all myrmicine ants, if not all ant species";
a view that in hindsight seems hopelessly naive.
The fact that most other common Myrmica species had fewer queens than M. rubra in-
trigued Elmes and he assumed that the difference might have an ecological basis. This re-
sulted in his 1972-1974 PhD study of the comparative ecology of Myrmica (southern
English species). He developed the idea that polygyny is an adaptive trait within different
local populations. Costs to individual queens being mitigated by shared benefits accrued at
the colony level, and so the interaction between queens evolved various checks and
balances that ensure an "ideal density" of queens per colony. A single mechanism would
thus explain the observed variation in queen-density between conspecific populations and
between species. He proposed an ¡dea that met with some scepticism, that colonies
periodically recruited new, often unrelated queens and that "surplus" queens (beyond the
minimum number required to maintain the worker population) could be thought of as tem-
porary social parasites (Elmes 1973a). Thus, while polygyny might be adaptive in some
circumstances, it always has the danger in some populations of degenerating into real
social parasitism; therefore Elmes regarded polygyny and microgyny as exhibited by
M. rubra as an "evolutionary precursor" to social parasitism (Elmes 1978a). The German
myrmecologist, Alfred Buschinger, explored similar ideas at about the same time to explain
social parasitism in Leptothorax Mayr species (Buschinger 1970).
Elmes' PhD study highlighted the fact that in many respects M. rubra does not provide
a model for all Myrmica species, rather it is the most unusual among the common British
species. Furthermore, as his post-doc studies developed, he found that population dynamics
and individual traits, e.g. basal physiology, larval ontogeny, varied between populations of the
same species. Consequently, he started retaining voucher specimens, working on the prin-
ciple that if one is going to compare different populations of the same species then one needs
to be sure that the populations are not different species. Elmes continued the practice of
voucher specimens when in 1980 he started a more applied study, collaborating with Jeremy
Thomas, on the biology and conservation of Large Blue Butterflies of Europe (the genus
Phengaris Doherty = Maculinea Van Eecke; for synonymy see Fric et al. 2007), which still
continues (Settele et al. 2005). This study chimed with Elmes' interest in social parasitism
in the genus Myrmica, because the 4th instar larvae of Maculinea butterflies can be consid-
ered as social parasites of Myrmica colonies. The details of the interaction vary between host
and parasite populations and may be related to variation between local races of Myrmica
throughout Europe. Examination of the voucher specimens led to Elmes' interest in Myrmica
taxonomy and since its modest beginnings of a few hundred British specimens it has grown
into probably the largest collection of a single ant-genus. It is now one of the major resources
that underpins this monograph.

9
 Alexander Radchenko's career followed a fairly conventional route within the scientific
community of Ukraine, then part of the former Soviet Union (FSU). In 1977, he graduated
from the State Pedagogical University in Kiev as a biogeographer. After some years
working as a teacher in a school, he was appointed as a PhD student to the Schmalhausen
Institute of Zoology of the Ukrainian Academy of Sciences (Kiev). The Institute adminis-
tration suggested that the taxonomy and biogeography of Ukrainian ants might be a useful
and appropriate topic for study, especially because the Institute holds the famous ant col-
lection of V. A. Karawajew. Radchenko successfully defended his PhD thesis in 1985 under
the tutelage of the Kiev hymenopterist Prof. Marina Zerova, and the famous Moscow
myrmecologist Prof. Gennady Dlussky. Subsequently, Radchenko concentrated on the in-
vestigation of taxonomy and zoogeography of various genera of Palaearctic ants. Many ant
species endemic to the FSU, were a mystery for western specialists : descriptions were often
brief and in archaic Russian, and many of the type specimens had been lost or misplaced
during the chaotic periods of the Russian Revolution and the Second World War. Further-
more, these problems were compounded by the general suspicions and lack of co-operation
by both sides during the Cold War period. Radchenko undertook a mission to resolve some
of the problems of the ant fauna of the FSU, especially for genus Myrmica, tracking down
type specimens and designating neotypes where specimens were lost. New horizons opened
for Elmes and others interested in the genus, when his revisions and keys for Myrmica
were translated into English (Radchenko 1994a-f). Radchenko was awarded the Doctor of
Science degree in 1999, for his work on the biogeography of ants.
Our collaboration, which culminates in the preparation of this monograph, started in
1982 when Elmes wrote to the Academies of Science of all 15 Soviet Republics, asking them
whether any entomologist would be interested in exchanging native ants (especially Myr-
mica ants) for taxonomic study. He had five positive results in a flurry of correspondence
with Soviet scientists that was quite unusual for that period of the Cold War. One of the
positive replies was from the President of the Ukrainian Academy of Sciences, who had
referred the request down the line until it dropped on the desk of Radchenko, who recalls
that by then it had become more of a "directive" rather than a request. There followed an
exchange of material and correspondence, particularly about the genus Myrmica, which
was already becoming a favourite group for Radchenko.
There was an immediate meeting of minds when we first met in person at the Second
International Colloquium of the Russian Language Section of the International Union for
the Study of Social Insects held in Rybnoe, the Ryazan Province of Russia, in September
1993 (see Kipyatkov 1993). The meeting was organized by Vladilen Kipyatkov of St. Pe-
tersburg, who with Elmes later obtained an INTAS grant (an EU initiative designed to help
FSU scientists survive the chaos of the Soviet Union break-up). Although the project was
primarily ecological, Radchenko headed-up an Ukrainian team because his taxonomic
expertise was essential for the reasons outlined above. We published our first paper together
in 1998 and thereafter have averaged two or three publications per annum. It almost goes
without saying that we could have achieved very little without the generous co-operation
of the other specialists and museums that are acknowledged in our various publications
and elsewhere in this monograph, and the grant funding and support provided by the
organizations listed below.
Firstly, we thank our respective Institutes: UK Natural Environment Research Council,
Centre for Ecology and Hydrology (Elmes), Ukrainian National Academy of Sciences,
Schmalhausen Institute of Zoology, Kiev, and Polish Academy of Sciences, Museum and

10
Institute of Zoology, Warsaw (Radchenko) for basic scientific support. Secondly, we are
grateful for the following awards that supported our collaboration EU INTAS grant (94-
2072), NATO Collaborative Linkage Grant (LST.CLG.976626), EU Framework VRTD
project "MacMan" (EVK2-CT-2001-00126) (Elmes and Radchenko); grant of the Ministry
of Science and Higher Education of Poland (2P04C 064 29) and UK Royal Society Collab­
orative Linkage Grant (Radchenko).
Finally, we thank our wives, Tamara Radchenko and Jenny Elmes, for their hospitality
and support, both moral and practical, during our fifteen years collaboration. We have spent
long periods in each other's homes and our wives have tolerated our visits and absences with
great patience; furthermore, even though we have often usurped our private holidays to
examine collections or to collect our own material, they freely provided invaluable scientific
assistance on these occasions.

и
CHAPTER 1. GENERAL BACKGROUND A N D BIOLOGY OF MYRMICA

1.1. Scope of this monograph

Neal A. Weber made the first and only attempt at a full worldwide revision of the genus
Myrmica and found that the plethora of names in the European literature, combined with
a paucity of material available to him from the Palaearctic, limited him to a revision of the
North American species with a synopsis of the Palaearctic taxa (Weber 1947,1948,1950).
He wrote "Since the North American Myrmicas are closely related to those of Europe and
Asia a study of the fauna of those regions was necessary. Unfortunately Bondroit and
others created numerous forms which cannot be evaluated from the descriptions alone.
Emery, Wheeler and others were highly skeptical of many. In the absence of much palaearc-
tic type material the original plan for a world revision has been abandoned" (Weber 1947:
438). We agree with Weber that many of the European varieties and sub-species of Myrmica
have no taxonomic value yet, strangely, he singled out Bondroit, who had a far better record
in naming "good species" of Myrmica compared to his contemporaries, probably because he
had been subject to criticism by his peers.
Early in our collaboration we also abandoned the attempt at a worldwide revision
constraining ourselves to the Palaearctic (or more correctly the Old World) for almost the
opposite cause to that of Weber. We decided to ignore the Nearctic Myrmica for several
reasons. Firstly, comparison of North American material in our own and other European
collections suggested that the American fauna of Myrmica is totally different from that of
the Old World, apart from a few species accidentally introduced from Europe. Secondly,
extrapolating from our preliminary studies of Europe, the Himalayan region and the Far
East of Russia we suspected that there are several biogeographically distinct faunas in North
America, and based on our more recent experience of southern Europe and South-East Asia
we inferred that there should be many locally endemic species, particularly in the southern
regions. This view was also expressed by our friend, the Canadian André Francoeur (pers.
comm.; Francoeur 1997), and is supported by our preliminary investigation of a very rich
collection of Myrmica recently taken from various places in North America for genetically
based taxonomic study by our colleagues Günther Jansen, Riitta Savolainen and Kari Vep-
säläinen. Thirdly, the keys for Myrmica available in North America (e.g. Weber 1947;
Creighton 1950; Gregg 1963 and other later regional keys based mainly on Creighton) now
appear to be quite outdated in their treatment of the species (Francoeur 1997 and personal
study), and might have led to regional inconsistencies in identification of the material
housed in various museums throughout the USA. Francoeur (2007) has started a revision,
but in the short term this might exacerbate the situation as some non-specialists use Fran-
coeur's revised names and other continue with the old keys. In our opinion, the time is
ready for a complete revision of North American Myrmica that should start with no pre-con-
ceptions and a fresh examination of the collections to erect a new taxonomy based on the

13
modem concept of species variation in the genus Myrmica. Only then, should the types be
examined and the existing names fitted to the modern taxonomy. We reluctantly concluded
that this major task could only be undertaken by someone based in America and we hope
that this revision of the Old World forms of Myrmica might stimulate a New World revision.
In Europe the situation was quite different. During the second half of the 19th century
and early part of the 20th century, European naturalists became obsessed with documenting
local forms and variation in plants and animals. The genus Myrmica also had this treatment
resulting in the plethora of names that Weber found so confusing, however the genus had
been subject to ongoing revision and major synthetic treatments with keys that more or less
agreed on the features for identification of the most common species (see section 1.4 below).
Consequently, our challenge was to decide on synonymies and sort out some outstanding
muddles of nomenclature in the literature. The task was similar in the case of Middle Asia,
Kazakhstan, Siberia, Far East, Mongolia and Korea, but was aided by the fact that Rad-
chenko already knew the Myrmica material available in FSU collections. Furthermore, Rad-
chenko (1994a-g, 2005) had made a first revision and key and established neotypes when
necessary due to some of Ruzsky's types being lost (or presumed lost, see Chapter 1.4 and
species catalogue, Chapter 3.3).
The Myrmica fauna of the Himalayan region, China, Japan and South-East Asia pre-
sented a different problem. The amount of material known to be in collections was relatively
small and the location of type specimens was not always known. Our first task was to revise
the ritae-group of Myrmica species (Radchenko and Elmes 1998) that at that time was be-
lieved to comprise a small number of morphologically archaic species (see Chapters 2 and 3).
Then we tackled the Himalayan Myrmica fauna (Radchenko and Elmes 1999, 2001b), where
many species had been confused and types and additional material was scattered in museum
collections worldwide. Once we had assembled the material it was relatively easy to order it;
for as Weber (1947, p. 437) surmised: "In a small collection from scattered localities it would
have been relatively easy to have assigned the ants various names, using new names which
differed more or less from previously recognised forms". However, once we published these
studies, various colleagues sent us unidentified material, that they had collected during the
last decades, which contained many more new species than we ever anticipated at the start
of our work. In fact, this new material has led us to reassess our views on the phylogeny,
biogeography and composition of the genus Myrmica (see Chapter 4).

1.2. The layout and use of this monograph

Four major chapters follow this introductory one, each is more or less self-contained
with some cross-referencing to other chapters. It is not intended that they need be read in
order therefore some ideas and information are repeated in the various chapters. Chapter
2 is an introduction to the basic taxonomy of the genus Myrmica. We discuss how Myrmica
fits with other related-myrmicines and outline the Myrmica species groups and complexes
that we currently recognize. We give the main diagnostic characteristics for the species
groups and discuss their use and relevance; a list of species assigned to each group and
a list of those that do not fit to any species group are also given.
Chapter 3 is a summary of the available data on Old World Myrmica. We first summarise
the collections that we have studied, and provide a list of all the names that definitely can
be assigned to the genus Myrmica. The core of this chapter (section 3.3) is an alphabetical
catalogue of the 142 names of extant Old World Myrmica that we consider to be "good"

14
species. For each of these species we provide full sets of detailed morphological drawings
for all known castes (usually based on type specimens). All figures (totally more than 1700)
are from originals drawn by Radchenko. Dimensions of the specimens drawn can be esti-
mated approximately from the mm scale lines. The catalogue can be used independently of
the rest of the book, being quite easy to find any particular name simply by flicking through
this section. Alternatively, any name and the page numbers relevant to it can be found in
the species tabular index given at the end of the monograph. For each species in the cata-
logue we indicate where type specimens are preserved, give a summary of its taxonomic
history, provide a re-description when the original description is inadequate by modern
standards, translate type-locality details when these were published in a non-mainstream
West European language (usually Russian), give a distribution map and outline of any
species-specific ecological information. This is followed by lists of fossil species and other
names that have been used in the genus Myrmica (nomina nuda, unavailable names etc.).
Chapter 4 is devoted to the zoogeography and evolution of the genus. We demarcate
nine centres of Myrmica species diversity and define and compare their faunas. Finally, we
outline the probable origin and evolution of the genus and discuss the possible origin of the
species groups. We drafted this section before we saw the fascinating molecular phylogeny
published by Jansen et al. (2010), subsequently we have tried to indicate to what extent
Jansen's results fit with our ideas on species groups (we thank Günther Jansen for making
his pre-publication results available to us).
Keys to the identification of the species form Chapter 5. Given the distinctiveness of
the fauna of the nine centres of Myrmica species diversity (see Chapter 4.1) we decided
that non-specialists would find it very hard to use a single key to the entire Myrmica fauna
of the Old World. Therefore we have produced separate identification keys for seven geo-
graphical regions that more or less coincide with the centres of diversity (see Chapter 5.1).
If users have an "unknown" specimen of Myrmica, they should first use the key for their
region, and if the species can not be identified therein, they should try the keys from the
neighbouring regions. Accurate identification of single male specimens of Myrmica is usu-
ally impossible without samples of workers from the same colony (or at least queens from
the same mating-swarm), therefore our keys are only for the workers. On the other hand,
many closely related (or at least morphologically similar) species are almost impossible to
separate without information on the males, therefore we give some male characteristics that
are useful in species separation. Users must accept that in several cases based only on
workers, without samples of all castes, one can only identify specimens to a small set of
cryptic species; any further determination is probabilistic, based on the location and
ecology of the site where the specimens were collected.
Most socially parasitic species (see section 1.3.6) are workerless, therefore we provide
a separate key to the social parasite queens and males; the few species that can have work-
ers or worker-like forms are also included in the appropriate regional keys to workers. When
parasites are captured in a host colony, they are readily distinguished from their hosts by
a combination of a suite of parasitic morphological traits (e.g. Kutter 1973; Radchenko and
Elmes 2003a). A problem arises when specimens are captured as solitary individuals because
some of the traits, such as increased hairiness and thickened waist segments, can be found
occasionally in mutated specimens, or "monsters" of normal species produced by disease
(e.g. Mermis Dujardin infestation; see Czechowski, Radchenko and Czechowska 2007; Cze-
chowski, Czechowska and Radchenko 2007; Csösz and Majoras 2008). We recommend that
caution should be observed when proposing "new".social parasites from single specimens.

15
1.3. The biology of Myrmica

Although there are about 1000 scientific publications that contribute to our under-
standing of the biology of Myrmica, most concern just a handful of species. The majority of
studies were made in Europe and the most studied species was M. rubra (L.) (= M. laevi-
nodis Nylander in many papers), which unfortunately is in many respects the least "typical"
biologically of the common European species. However, species from several other species
groups have been studied in detail and these results, supported by more anecdotal obser-
vation of some of the less known species, suggest that a number of behavioural and life-
history traits are probably common to all and might be considered basal within the genus
Myrmica. Below we simply provide an overview for the non-specialist reader to indicate
what sort of data might usefully be recorded for the vast majority of unstudied species; it is
not intended as an exhaustive review, the references given can be used as a starting point
for a more thorough literature search if required. Some extra information for the better
studied species, especially if they vary markedly from the general pattern, is given in the
catalogue (Chapter 3.3).

1.3.1. Myrmica in ecosystems

Myrmica colonies can be found in almost all habitats in the temperate zone
of the Northern Hemisphere, mainly north of approximately latitude 40-45°: meadows,
steppes, woodlands, forests and mountain tundra etc. Further south at sea-level, local
species are associated with meadows and woodlands that have some moisture in summer,
bordering rivers, lakes, marshes etc. or in habitats where there is high summer rainfall.
Otherwise, species live on southern mountains at a range of altitude where temperate con-
ditions prevail. Local isolation on such mountains appears to have led to a much greater
range of locally endemic species (see Chapter 4). In all habitats the average soil tempera-
tures governed by latitude, aspect and the shade provided by the vegetation, determines the
suite of species that can potentially utilize the habitat (Elmes et al. 1998; Berman et al.
2010). This is dynamic in natural habitats but often less so in woodlands and meadows
maintained by man's activities. In some places Myrmica colonies can be very abundant,
comprising only one or two species or in some cases up to eight species, in others colonies
are rare and well dispersed. In short-turf grasslands nests frequently are built in the soil,
often having an almost invisible single small (ca. 3-5 mm) entrance hole, sometimes these
nests are at densities of one per 4 m2 (Elmes and Wardlaw 1982c) and can only be found
by baiting (Elmes and Wardlaw 1982a). In longer vegetation colonies often build solaria,
small mounds of soil and dead vegetation, to enhance nest temperatures, or nests are built
in tussocks of grass and moss; in woodlands they often nest in and under pieces of rotten
wood, in rotten tree stumps or even in twigs and acorns (Banschbach and Herbers 1999);
in more exposed sites on hills and mountains Myrmica often nest in the soil under flat
stones that heat up quickly in the morning and retain their heat longer than the
surrounding soil. It is very hard to define nest-site choice for any particular species: they
seem to be able to use all when necessary. Similarly, widespread species vary over their
range, for example M. rubra is primarily considered a grassland species in Western Europe,
but is mainly associated with forests in Eastern Europe (Elmes et al. 1999).
Myrmica colony structure can be quite labile (see below) and nests can be fairly mobile.
In some circumstances colonies of various species migrate and it is possible to find a trail

16
of abandoned sites (Elmes 1978b), in others a colony changes location seasonally around
a tree or bush or grass tussock according to the shade afforded. On the other hand, some
nest-sites remain occupied for many years, for example we have observed M. rubra living
under the same stone for many tens of years (unpublished data). In areas of short grassland
there appears to be strong competition for nest-sites, i.e. the same system of tunnels and gal­
leries in the soil (Elmes and Wardlaw 1982a). If a nest site is abandoned it is soon re-colo­
nized by the same or a competing Myrmica species (Mouquet et al. 2005) and it has been
suggested that in such habitats populations are more limited by suitable nest-site availabil­
ity than by resource availability (Elmes 1973a; Hochberg et al. 1994; Zakharov and
Fedoseeva 2005). Joanna Pętal and her Polish co-workers showed that Myrmica colonies
retrieve large amounts of insect protein (Pętal 1967, 1981; Kajak et al. 1972) and can have
big impacts on the locai invertebrate community (Pętal and Breymeyer 1969), the detritus
and other waste products of a colony can have a large impact on the microorganisms and
nutritional content of the nest soil (Czerwiñski et al. 1969, 1971; Jakubczyk et al. 1972).
While colonies are affected by habitat management (Pętal 1974, 1976), local soil enrich­
ment might account for the different vegetation assemblage observed above soil nests
(Elmes and Wardlaw 1982b).
In recent years, understanding the population dynamics of Myrmica communities has
become important in the practical conservation of Phengaris (= Maculinea) butterflies (see
below) whose fourth instar larvae are obligate parasites in Myrmica nests (Elmes and
Thomas 1992). The UK, Centre for Ecology and Hydrology team have developed a series of
spatially explicit simulation models that take account of the population dynamics of the
host ants and the behaviour of the Phengaris parasites (Hochberg et al. 1994; Elmes et al.
1996). Not only have these models been rewarding in predicting and understanding the
factors that might affect the persistence of Phengaris populations (Clarke et al. 1997), they
have thrown light on the competitive interaction between Myrmica colonies. For example,
one study showed that in a community of two strongly competing Myrmica species, if one
of them is disproportionately affected by a Phengaris parasite (or for that matter any other
parasite or disease), the spatial distribution and proportion of the two species will alter
(Clarke et al. 1997; Thomas et al. 1997). This effect or "footprint" persists long after the
parasite goes extinct and takes tens of years before reverting to the original balance. The
models assumed colonization of newly vacated nest sites by division of neighbouring
colonies. Interestingly, it makes little difference to the outcome whether re-colonization by
solitary queens is permitted or not - the reason being that young incipient colonies are
usually absorbed by a bud of a neighbouring colony if the same species, or killed if
different species (unpublished data).

1.3.2. Anatomy and surface chemistry

There have been few publications on the internal anatomy of Myrmica since Charles
Janet used "Myrmica rubra" in many of his wonderful, detailed studies of ant and social
insect anatomy (see Billen and Wilson 2008 for an appraisal). Many of Janet's drawings
have been copied in various books on ants since he first published them (e.g. Wheeler 1910)
and there is no point in reproducing them here. Most subsequent publications have
been concerned with the structure of various glands and the sting apparatus of females
(e.g. Kugler 1978; Billen 1986). We have suggested that a functional study of the an­
tennae might prove useful in understanding why the shape of the female castes' scape and

17
length of the male scape is so important in Myrmica classification (Radchenko and Elmes
2001b).
It had long been known that some of the most important sense organs on the antenna
of ants detected odours and deprivation of antennae affected ants abilities to interact with
nestmates and aliens (e.g. Forel 1874) and by 1900 this was developed into a theory of nest-
mate recognition by "learning" odours (Fielde 1901). As analytical techniques for organic
chemistry were refined, efforts were made to identify the source and nature of the chemi-
cals used in social insect recognition. A short review of ant chemistry (Blum 1970) was
published in a wider review of insect chemical communication. Thereafter, just as Janet had
turned to Myrmica to represent a typical myrmicine ant, chemists used species from the
genus in studies of ant chemistry. The most productive in this respect was a team of Eng-
lish and Belgian scientists who frequently used Myrmica ants to study the secretions of
various glands, especially the Dufour's gland, and bioassay the chemicals' roles in trail-
making and alarm signalling (e.g. Cammaerts et al. 1982; Evershed et al. 1982; Attygalle et
al. 1983). Cuticular hydrocarbons were believed to be good candidates for the colony recog-
nition signals, though more recent writers believe this is too simplistic (reviewed by Van-
der Meer et al. 1998). The idea that social parasites might mimic their ant hosts by
synthesizing a similar suite of hydrocarbons has been studied in a few cases (Howard et al.
1990; Akino et al. 1999; Lenoir et al. 2001). Regardless of how these results might be in-
terpreted in terms of overall theory on "social odour" there is no doubt that the differences
in the cuticular hydrocarbon profiles of the common European Myrmica species are suffi-
cient to easily discriminate them, while within-species differences due to geographical lo-
cation, colony or age (including larvae) are comparatively much smaller (Elmes et al. 2002).

1.3.3. Colony structure

A colony of Myrmica ants is defined as those individuals living together in a discrete set
of nest chambers (Elmes 1973a). Colonies vary in size from a few tens of workers to many
thousands with an average size for the common European species between 200 and 500
workers depending on the species (Wardlaw and Elmes 1996). In some circumstances
colonies are quite labile, readily splitting or coalescing according to environmental condi-
tions (Elmes 1980, 1987; Banschbach and Herbers 1999). In other circumstances adjacent
nests interact and often exchange individuals so that they can be considered as a polydo-
mous colony (Pedersen and Boomsma 1999a). Sometimes the number of interacting nests
is sufficiently great that they more or less form a "super-colony". M. rubra is the best-known
species to do this in Europe and we have observed M. rubra super-colonies that comprise
many nests living at high density, completely dominating relatively small patches of habi-
tat, and totalling many hundreds of thousand individuals. Where M. rubra has been intro-
duced into north-eastern America, this occurs on a grand scale, super-colonies can comprise
millions of individuals dominating many hectares of habitat (Groden et al. 2005). However,
the size of the component colonies is about the same as that observed in Europe (ca. 1000
workers), but they are packed together at very much higher densities (Garnas et al. 2007).
In all the species where detailed population censuses have been made, at least some
colonies are polygynous, that is they have more than one functionally reproductive queen.
Average levels of polygyny vary between and within populations of the same species (Elmes
and Petal 1990). Among the European species the least polygynous is the normal form of
M. ruginodis, called M. ruginodis var. macrogyna by Brian and Brian (1955); the most

18
polygynous is the microgyne form, called M. ruginodis var. microgyna by Brian and Brian
(1955), that average about one and 50 queens per colony with about 400 and 20 workers
per queen respectively (Elmes 1978b; Elmes and Keller 1993). Polygynous and monogy-
nous forms of M. ruginodis were reported from Japan (Mizutani 1981), but it seems
probable that these refer to M. kotokui and M. ruginodis respectively. For further discus-
sion of microgyne forms see the section on social parasites (1.3.6) and the notes on these
species in the catalogue (Chapter 3.3). The frequency distribution of the numbers of
workers per colony in any population of colonies is more or less log-normal for all the
species studied to date, suggesting that providing there are sufficient queens to maintain the
worker population, the logistic growth model proposed by Elmes (1973a) might apply
generally. It was suggested that the actual values of the two parameters that determined
colony growth are functions of the particular nest-site that are determined by several inde-
pendent environmental factors. This has been used successfully in simulation models of
Myrmica populations (Hochberg et al. 1994).
The frequency distribution of queens is even more over-dispersed than a negative bino-
mial distribution, but examination of the 3 rd and 4th moments of the frequency distribu-
tions of queens for the common European species shows that they can all be generated by
the same underlying mathematical function known as a Pearson type III curve (Elmes and
Keller 1993), which suggests that the underlying process by which polygyny develops in in-
dividual colonies is the same for all Myrmica species. Hölldobler B. and Wilson (1977) sug-
gested that recruitment efficiency controlled by a simple random variable would produce the
same effect as the interacting host-parasite type growth model suggested by Elmes (1973a),
but in order to produce such overdispersed distributions more than one variable would be
required.
Elmes (1980) used demographic arguments to suggest that most Myrmica queens are
short-lived and that when a queen dies or becomes old, the workers "replace" her. In most
populations fertilized queens are foraging for much of the year and are "eager" to join
a colony when permitted to do so (Hölldobler K. 1938; Elmes 1982). Natural colonies vary
in the degree of intolerance and aggression shown towards new queens and this can vary
in populations from year to year (Elmes and Wardlaw 1983a). In the laboratory, if experi-
mental cultures of workers are deprived of their queen, they soon become very tolerant of
replacements (Brian 1988a). Beekeepers use much the same technique to introduce
replacement queens into hives. Elmes (1980) suggested that queen recruitment is under
the control of the workers: when the density of queens falls below a threshold level much
of the colony is effectively queenless, seldom encountering a queen due to the spatial
heterogeneity of natural nests. In queenless colonies some workers become reproductive
laying viable male-eggs (see below), but the majority become amenable to accepting re-
placement queen (s) even one(s) quite unrelated to the colony. If colonies are small then the
recruitment of one or maybe two replacements might be sufficient, but in larger, more spa-
tially dispersed nests recruitment might continue in a second year. Recruitment stops when
surviving queens are integrated into the society and become fully reproductive (this is
assessed by pheromonal communication, see below), and workers again become intolerant
of would-be joiner queens and oviposition by other workers. As queens die and age the
colony again enters a more tolerant phase and eventually recruits again, thus in larger
colonies (assuming that environmental conditions do not lead to a change in worker
numbers) one might expect a cycle in queen numbers. In the.case of M. sulcinodis such
a cycle was detected at the population level (Elmes 1987) and it had about a five year

19
periodicity, which would be expected if queens lived on average only two years (as pro-
posed by Elmes 1980; see also Seppä 2003). Elmes suggested that detection of the cycle
was fortuitous in the M. sulcinodis populations studied, because individual colony cycles
were put in phase with each other by an environmental disaster (major fire). Other major
environmental fluctuations might explain why different populations seem to some extent to
vary in polygyny levels synchronously (Elmes and Petal 1990).
Quite a number of North American species have some polygynous colonies (e.g. Talbot
1945; Banschbach and Herbers 1996; Evans 1996, and personal observations). M. kotokui
can be highly polygynous in Japan (Kasugai et al. 1983; Kikuchi et al. 1999) and Korea
(personal observations). The peculiar species M. arnoldii is usually quite polygynous
(Michal Woyciechowski, pers, comm.), and the occurrence of two or more queens in colony
samples of Himalayan and Siberian species is not unusual (personal observation; Berman
et al. 2010). Unfortunately, there is nothing comparable to the European data for queen
number in Myrmica from most of the other Zoogeographie regions. However, we suspect
that in each region there will be a few highly polygynous species, while most have relatively
small and isolated colonies that rarely have more than a single queen (rather like the
macrogyne form of M. ruginodis - see above). Little is known about the population struc-
ture of the ritae-group species, several queens have been found in a few colony samples
(our unpublished data), and we suspect that the group will comprise species exhibiting
much the same range of population structures as other groups. We conclude that all species
of Myrmica have the potential to develop polygynous colonies in some circumstances, and
a few are habitually highly polygynous; based on our own observations it seems that the
species that thrive in anthropogenic habitats, such as those of much of Europe, are more
polygynous than those associated with more natural habitats.

1.3.4. Social interactions

We include under this heading all aspects of the reproductive life cycle, individual be-
haviour and colony "management". The team based at the Furzebrook Research Station,
UK with principal investigators Michael Brian (from 1950 to 1980) and Graham Elmes
(from 1970 to 2000) were particularly concerned with developmental and physiological
aspects of European Myrmica species: these ants are particularly amenable to laboratory
experimental culture (for a review of methods see Wardlaw et al. 1998). The more recent
studies of the underlying genetic-influences and consequences of the social interactions,
using Myrmica ants as model systems, emanate from laboratories throughout the Northern
Hemisphere.
All fertilized queens lay eggs: M. rubra lay about 200-300 eggs per annum (Brian 1962),
and Brian (1951a) estimated about 300 eggs per queen for M. ruginodis in two wild
colonies, assuming eggs take about three weeks to hatch. Oviposition starts in early spring
when nest temperatures rise above ca. 14°C, depending upon the species (unpublished ob-
servation). Eggs are produced throughout the summer until temperatures drop in autumn
(see next section), individual queens oviposit in at least two pulses (Brian 1951a), mostly
producing a mixture of fertilized and a few unfertilized "male-eggs" and some trophic
eggs (Wardlaw and Elmes 1995). Although there have been no specific studies on the
workers' tolerance of unfertilized queens, called gynes, in colonies of Myrmica, it is generally
accepted that after the nuptial flight season they are either ejected or killed: no unfertilized
queens were ever found in polygynous colonies of M. sulcinodis, though occasionally

20
queens assumed to be old individuals, with degenerated ovaries, have been found (Elmes
1987). Some gynes fail to develop fully their spermatheca, but this is very rare and such
individuals usually have other intercaste features and adopt a worker role. Sub-fertile
old and young queens are relegated to the periphery of the nest and are not fed properly
(Brian 1988b) and we believe some have to forage for themselves outside the nest, and
may abandon the nest if they encounter another colony that is prepared to accept them
(see above).
Fertilized young queens must overwinter before their ovaries develop properly. Fully
functional M. rubra queens have between 10 and 30 ovarioles with the total length of
ovariole being proportional to the size (weight) of the queen (Elmes 1976). The average
number of ovarioles possessed by queens of different species probably accords to average
queen-size. Young queens vary in their reproductive potential (Brian 1988b). Even before
the ovaries fully develop, young queens are not equal, if placed in a pleometrotic group
some individuals fully adopt a worker role, tending eggs and foraging for a few "dominant"
individuals that fully adopt a queen role while others act as nurses (unpublished observa-
tion), and in some respects these temporary all-queen societies resemble those of primi-
tively eusocial wasps (Agrahari and Gadagkar 2006).
In all species eggs and young larvae are kept separately from the older larvae, more or
less in the centre of the nest in a region that has been termed the "core" (Brian 1986a).
The "key" to reproductive success for a queen is to gain entry to the "core" (Brian 1986a,
b, 1988b), where the egg-mass is protected and tended by a group of specialist workers that
could be considered "elite" (Elmes and Brian 1991). It was suggested that the elite
workers determine which queen reproduces because they control when the various queens
are permitted to enter the core: eggs laid in the core have a high chance of being added to
the egg-mass whereas those laid in the surrounding brood chambers are usually fed to the
developing 3 rd instar larvae by the nurse workers. In a large, dispersed natural colonies
there might be several "cores" that in some sense could be considered as competing. There
is a turn-over of queens in the core with some being expelled, and others gaining admission
as the season progresses. This process can easily explain the variation in effective repro-
duction between queens (e.g. Evans 1996; Pedersen and Boomsma 1999a, b; Seppä 2003)
and might lead to some occasional physical contest between queens, as reported by Evesham
(1984). For a queen to produce gynes, it is important that she should oviposit early in the
season as most eggs laid later become workers (see below).
All the Myrmica species studied to date have some workers capable of oviposition
(Wardlaw and Elmes 1998 and references therein). Individual Myrmica workers have zero
to six potentially active ovarioles, though one is the most common number (Wardlaw and
Elmes 1998 and unpublished data), the very few that have more than two ovarioles are usu-
ally larger individuals or "failed queens" (see below) and there are reasons to suspect that
these may dominate in the core. Workers lay two types of eggs: reproductive eggs that con-
tain an embryo that potentially can develop into males, and trophic eggs that have no em-
bryo and are used as larval food (Brian and Rigby 1978; Wardlaw and Elmes 1998).
Queenright workers adopt a different suite of behaviours compared to those of queenless
workers (Brian et. al. 1981) - the so called "queen-effect" - that is moderated by chemical
signalling (e.g. Brian 1976). When queenright, the ovaries of most workers are fairly
inactive and the vast majority of any eggs laid are trophic, but after a few days of queenless
conditions many workers start ovipositing (a few laying abundant eggs) and the majority
of the eggs are reproductive. Reintroduction of a queen reverses the process (see above).

21
However, if male-eggs are placed on the egg pile, they do not seem to be recognized as such
and appear to have similar survival chances to those of worker-larvae (Brian 1981). It has
been suggested that the majority of males in Myrmica are produced by workers (see Smee-
tonl981, 1982).
Thus, the egg pile comprises a mixture of fertilized and unfertilized eggs laid by the
queens and possibly some unfertilized eggs laid by the elite workers. Other than Weir
(1959b), relatively little is known about the interactions between larvae and workers on
the egg pile, because at this stage they are almost impossible to observe in a normal
experimental set-up. The following is our interpretation of anecdotes given in many publi-
cations supported by our own observations.
The eggs are stuck together by a "soup" of liquid regurgitations of the workers and
the fluid released by first instar larvae as they puncture their egg membrane. Larvae
cannibalize adjacent eggs, but we suspect they may find it easier to penetrate the outer
membranes of "flaccid" eggs, i.e. reproductive eggs with a dead embryo caused by
purposeful or accidental damage by the workers, and trophic eggs. Other than egg preda-
tion, it seems that first instar larvae get no feeding beyond the workers drinking the "soup"
and regurgitating it onto the pile, possibly with added antibiotic secretions. Both eggs and
larvae soon dehydrate if they are removed from the egg-pile. When larvae reach the second
instar, they get more attention from the workers and as they grow, they are removed from
the centre of the egg pile and are fed by regurgitations. Once they achieve the third instar,
they are taken from the core and placed in the surrounding brood chambers. The develop-
ment time from egg to third instar larvae can take only two to three weeks, depending on
incubation temperatures (Brian 1957).
The interactions between third instar larvae and nurses are much better known (e.g.
Brian 1956, 1965; Brian and Hibble 1964). They are fed on a mixture of regurgitations,
containing both sugars and proteins (Brian and Abbott 1977) and gain more than 90% of
their final weight (Elmes et al. 2001). The ontogeny of third instar larvae has been
examined in great detail (Weir 1959c; Brian 1954, 1955) and caste determination is con-
trolled by the balance of hormone production (e.g. Brian 1959; Brian and Hibble 1963),
which in part is intrinsic and in part, due to environmental conditions, such as a tempera-
ture, food availability, activity of nurse workers etc. (Brian 1962, 1963, 1973a). Female
larvae that are precipitated into premature metamorphosis become workers because they
have not fully developed the tissue-precursors of some adult characters (wings, ovaries,
spermatheca etc.). Thus, to a large extent the developmental path of larvae produced early
in the year is under the control of the nurse workers. When queenright, the workers place
the larvae at higher temperatures, feed them less and attack them mildly, usually by biting
(Brian 1973b) : all these treatments stimulate early metamorphosis into the worker caste and
this stage has been termed "the first switch". Queenless nurses adopt the reverse behaviours
and many fewer larvae metamorphose at this stage. These larvae grow quite large and
before winter they enter a full, or obligate, diapause (Brian 1975) that can only be broken
by some weeks at cold temperatures (ca. < 5°C). Larvae reared from later-laid eggs stop
growing and enter a facultative diapause when temperatures fall.
These studies showed that when a colony hibernates, most third instar diapausing lar-
vae survive winter, while eggs and younger larvae do not, being usually killed and eaten by
the workers before hibernation. The numbers, size and developmental potential of over-
wintered larvae vary between species (Elmes and Wardlaw 1981) and generally only the
largest of the overwintered third instar larvae have the potential to develop into new queens.

22
Those that achieve this, mate and must survive the following winter before producing their
first offspring, at which time they are already two years old (since egg). However, in spring
environmental manipulation, in much the same way as in the previous summer, can still pre­
cipitate the larvae into premature metamorphose (this has been termed "the second
switch"). Therefore, when social conditions change between summer and the following
spring (e.g. the recruitment of new queens - see above), workers still have control over the
sexual production. In small experimental cultures, any larvae that have passed "the-point-
of-no-return" towards developing as a gyne, are killed but those almost at that point meta­
morphose to become large workers, often having wing-buds or vestigial wings, and more
ovarioles than normal workers (Brian 1955). It may be that these individuals lay more eggs
and more probably become "elite" workers (see above).
We suspect that this basic developmental life-cycle will be shown to be present in all
Myrmica species. Cross-over experiments whereby experimental cultures were established
with workers of species A, containing a queen of species B, and rearing over-wintered lar­
vae of species С (Elmes and Wardlaw 1983a and unpublished observation), suggest that
the mechanisms and control signals for larval manipulation were the same. In other words,
the mechanism whereby workers detect the presence of a fertile queen (i.e. a chemical
signal), the change in behaviour engendered towards gyne-potential larvae and the response
of larvae towards the behaviours are probably old conserved traits within the genus.

1.3.5. Physiology

Details of the life cycle and individual social interactions (outlined above) obviously
interact with basal individual physiology and what we consider to be colony-level physio­
logical interactions with local environments: the team of Vladilen Kipyatkov (from 1980 to
the present) has contributed much to our understanding of the latter aspect. Kipyatkov and
co-workers have shown that queens of different species from different environments start
ovipositing at different times and have different brood production strategies in terms of
pre-winter and post-winter investment (Kipyatkov et al. 2002, 2005; Kipyatkov and
Lopatina 2002). Larvae, queens and workers use varying and different cues to determine
when to enter diapause (Kipyatkov and Lopatina 1989 and references therein), a rough
summary is that northern species and populations within the same species tend to use day-
length to determine when to diapause, whereas southern ones usually respond to falling
mean temperatures. Kipyatkov (1988) suggests that there may be a specific day-length-
mitigated pheromone released that co-ordinates diapause within a colony.
Nielsen et al. (1999) have shown that different species have different basal respiration
rates, the northern species respiring more actively, and that a similar trend can be observed
within-species sampled from different latitudes (Elmes et al. 1999). This effect is carried
over into the rate at which workers work and larvae grow, cross over experiments showing
that it is an attribute of both the nurses and the larvae. Populations from different latitudes
appear to have different fat-storage strategies (Elmes et al. 1999 and unpublished data).
Thus, each species is adapted to live and work at a different rate when reared at the same
average temperature, the response to temperature being the same for each of the species
studied (Elmes and Wardlaw 1983b), which can be thought-of as a series of species-specific
regressions each with the same slope, but a different intercept. In southern England, the dif­
ference between the intercepts for the four common species studied was the exact reverse
of the difference in average nest-temperatures observed in strong populations (Elmes 1982).

23
In other words, when living in their preferred habitat, each species has a basal physiology
that ensures larvae develop at about the same rate and sexuais are produced in early sum-
mer, ready for nuptial flights in August and September.
Thus, the overall life cycle strategy seems to be common to all Myrmica species, young
queens must over-winter before they start brood production because if they fly earlier there
is insufficient time to rear new workers. Kipyatkov (2001) reviewed seasonal life cycles and
dormancy in ants and used the term heterodynamic for species that have some adaptation
to survive cold periods (winter). He places Myrmica species in his endogenous heterody-
namic category, which can be found in all regions outside the tropics and subtropics. This
category includes species with good adaptations to survive winter, adults usually entering
an obligate diapause and overwintering larvae a facultative diapause (though this does not
account for the obligate diapause of sexual larvae, reported by Brian 1975). In our obser-
vations of low-altitude southern-living species of Myrmica it is possible that some might
verge on the exogenous heterodynamic strategy. One might raise the question, why Myr-
mica have not evolved physiologies that enable them to "live faster", rear brood immediately
after mating and so losing the extended brood development cycle, evolving life-cycles more
like that of the very successful temperate Lasius F. species. We think that the ancestral Myr-
mica life-cycle probably evolved in response to scavenging opportunistically on diverse tem-
porary food resources where low-energy, slow physiology and brood that could survive
periods of food shortage were advantageous. If the expansion of the genus into more tem-
perate environments occurred later than, for example, in Lasius and Formica L. species,
the basic life style might have been conserved because competitors were already much bet-
ter adapted to utilize seasonally abundant resources such as aphids and honeydew. Almost
all published observations on the timing of nuptial flights give dates in August and Sep-
tember, the only exceptions are anecdotal accounts of flights in May, June or July, from
populations living at very high latitudes or very high altitudes. However, in these environ-
ments the summer season is very short and there is anecdotal evidence that local popula-
tions have adapted to this, possibly having evolved a three-season brood cycle with gynes
hibernating twice as larvae and so would be three years old (since egg) before reproducing
(Bermanetal. 2010).
There has not been a major comparative study on the nuptial flights of Myrmica species,
probably because of the limited period when they occur, and of the difficulties of observing
exactly what happens within mating swarms. However, there are a great many anecdotal
accounts spread throughout the literature and the following is a sample (Beare 1913;
Donisthorpe 1927a; Boomsma and Leusink 1981; Collingwood 1958a; Ichinose 1990). In
our experience, Myrmica swarms start in late morning or early afternoon with males
leaving the nest, these start a local swarm usually over some prominent land mark:
a hilltop, a tree, telegraph pole, bush or large rock etc. The males appear to release a pun-
gent chemical (capable of burning small holes in cotton) that is noticeable to a human sense
of smell if one ventures into a swarm. This may be detectable by other males and gynes over
a large area because the swarm starts attracting more males and gynes. As the swarm in-
creases it often subsumes smaller male swarms until it gives the impression of smoke spi-
ralling above the landmark. M. rubra seems to be unusual in that individuals appear to fly
further and higher than is the case in many other species (Woyciechowski 1990a, b). Gynes
fly into the swarm, many copulate in flight and try to fly away from the swarm with the
male in copulation, but others fall to the ground where they are often besieged by several
competing males. Some gynes mate more than once (Pedersen and Boomsma 1999b) and

24
it is not known whether these are the ones that mate on the ground or whether those mated
aerially return to the swarm after copulation is finished (the duration of copulation appears
to be controlled by the gyne). In the case of M. ruginodis there is no indication of assortative
mating by size, but larger males are more likely to get a mate than smaller ones (Elmes 1991).
Finally, we and others have often observed mixed swarms of Myrmica species, occa-
sionally as many as six different species, but on the few occasions that we have checked
pairs taken in copulation, we have always found gynes mating with conspecific males. The
mating swarm stops when the day cools and the males cluster together and pass the night
on the ground or under the leaves of the tree or bush used as a mating post. If the weather
is suitable for swarming the next day, the surviving males reactivate the swarm and draw
in new males. Individual males probably only survive several days, but if weather condi-
tions permit the swarm can persist for much longer. We consider that the above observa-
tions indicates that the basic mating behaviour is very much the same between all groups
of Myrmica, and that the chemical signal males use for attracting.males and gynes to
a swarm must be very similar between all the species. Conversely, the failure to detect cross-
species pairings in mixed swarms suggests that there are strong barriers to hybridisation,
though what these might be are not known.

1.3.6. Social parasitism

Using a strict definition of social parasite to include only other ant species, the genus
Myrmica has long been known to have a considerable number of social parasites
associated with it (Kutter 1973) and using the modern more broad definition to include
species that in some way manipulate and exploit the ant society there are many more (Don-
isthorpe 1927b; Thomas et al. 2005). All the known ant social parasites of Old World Myr-
mica species are themselves members of the genus Myrmica (Bolton 1988; Radchenko and
Elmes 2003a), an exception being the New World Formicoxenus provancheri (Emery) (see
Kannowski 1957; Errard et al. 1990).
The presence of microgynes in some populations of ants was well known (Wheeler
1910), but none of the older authors found it particularly remarkable. Elmes (1978a) pro-
posed an evolutionary progression from microgyne (exemplified by microgyne form of
M. ruginodis - see above) through the "parasitic" microgynes of M. rubra, the species spe-
cific true parasites such as M. hirsuta Elmes, to the specialized parasites such as M. kara-
vajevi (Arnoldi) that can use several host species. This hypothesis envisaged some sort of
local sympatric speciation within populations, some genetical support for aspects of this
idea has published recently (Savolainen and Vepsäläinen 2003; Vepsäläinen et al. 2009;
Jansen et al. 2010). Experimental studies on the biology of true socially parasitic Myrmica
are few, mainly because the parasites can not be found with sufficient regularity in sufficient
numbers to ensure a coherent short-term research programme. Generally studies have been
opportunistic (Elmes 1994).
The microgyne forms of M. ruginodis and M. rubra have been better studied. Brian and
Brian (1955) showed that M. ruginodis microgynes are free living, produce normal
workers and they and their societies are more tolerant of foreign workers and queens. They
proposed that microgyny was an adaptation to the stable, persistent plagioclimax habitats
of the oceanic seaboard of Western Europe (see also Hölldobler B. and Wilson 1977).
Elmes (1978b) showed that while some nest of M. ruginodis contain only microgyne queens,
many also contain some normal queens; he classified these colonies as "microgyne" and

25
demonstrated that they are much more polygynous than colonies containing only normal
queens (see above). There is some evidence that the two forms have different strategies for
sexual production and larval hibernation (Elmes and Wardlaw 1981). Microgynes of M. ru-
ginodis are now known to be much more widely distributed than envisaged by Brian and
Brian (1955): we have found them in European populations ranging from Finland to Turkey
(unpublished data). The situation in Japan and Korea is not clear, oriental M. ruginodis are
slightly different morphologically from European populations and live sympatrically with
the morphologically and ecologically similar M. kotokui. Microgyny has been reported in
oriental M. ruginodis (Mizutani 1981; Kasugai et al. 1983) and polygynous and monogynous
populations of M. kotokui have been also described (Ichinose 1990; Kikuchi et al. 1999).
We are not absolutely clear from this as to whether both M. ruginodis and M. kotokui have
"normal" queens and more polygynous "microgyne" populations, or whether only one of the
species does, or whether M. kotokui has been confounded with M. ruginodis microgyne
(see species accounts in Chapter 3.3).
The fact that one could find microgynes in some populations of M. rubra was well known
(e.g. Donisthorpe 1927a; Collingwood 1958b; Elmes 1973b). For a short period the mi-
crogyne form was considered a separate species, M. microrubra Seifert, but recently this
name was synonymized with M. rubra (for details see Chapter 3.3). The first experimental
study was made by Elmes (1976) who showed that using the ratio of thorax-width (= maxi-
mum width of mesosoma in queens and pronotum in workers) to head-width as an index
of "queenness" most microgynes were simply an isometric reduction of normal queens, de-
spite the considerable range in their size. However, some winged individuals had the same
size and thorax-ratio as normal M. rubra workers, being intercastes with no spermatheca
and poorly developed ovaries. Elmes suggested that in one sense these could be considered
as "winged-workers", though there is no evidence that they did much work simply being
a drain on the colony's economy. Rearing tests showed that only the larger microgynes had
spermatheca and were fertile, and could exert the "queen-effect" associated with normal
queens (see above and Elmes and Brian 1991). Larvae reared from their eggs in summer
became intercastes (winged-workers), while most of the larvae that entered diapause and
overwintered became new microgynes (with a range of sizes and reproductive develop-
mental status), some became workers indistinguishable from those of normal queens and
a few became "winged-workers". For these reasons Elmes (1976) suggested that microgy-
nes arise within local populations due to some sort of mutation derived from normal
polygynous behaviour to take advantage of ecological conditions (see Brian and Brian 1955;
Hölldobler B. and Wilson 1977). However, in the case of M. rubra the microgyne strategy
has no ecological benefit and simply produces a proto-parasite: colonies of M. rubra con-
taining microgynes were shown to be disadvantaged (Cammaerts et. al. 1987). In support
of the sympatric speciation hypothesis, Savolainen and Vepsäläinen (2003) confirmed that
M. rubra microgynes are more related to their local host populations than to each other
(see also Vepsäläinen et al. 2009; Jansen et al. 2010). Nevertheless, despite many years of
breeding tests on M. ruginodis and M. rubra in Furzebrook Research Station and other labo-
ratories there has never been a report of normal queens giving rise to microgyne offspring
or vice-versa, so if microgyny is a result of a local mutation, this event must be quite rare.
Even less is known about the biology of M. hirsuta, but a study by Elmes (1983) indi-
cates that fundamentally its reproductive cycle is similar to that of M. rubra microgynes, but
even more adapted towards social parasitism. Small queens can not be fertilized, but the
production of intercastes is much rarer, only three worker-like specimens were found in

26
a population from Denmark (Elmes 1994; Radchenko and Elmes 2003a). M. hirsuta is a spe-
cific parasite of M. sabuleti Meinert and its close relative M. lonae Finzi; its queens are usu-
ally found in colonies containing an active host queen. Reproductively active M. hirsuta
queens can {via the host workers) suppress the sexual development of larvae of the host
queens and there is a suggestion that they can "delay" the oviposition by the host queen
until after they begin ovipositing themselves (Elmes 1983). There was less strong evidence
for a paraphyletic origin of different M. hirsuta populations, as was the case for M. rubra
microgynes (Savolainen and Vepsäläinen 2003), suggesting that if M. hirsuta has evolved
by a similar route to M. rubra microgynes, then the original event occurred longer ago so
that the parasitic form spread post-glacially throughout Europe together with its host. Even
so, the recent molecular phylogeny of Myrmica social parasites obtained by Jansen et al.
(2010) estimates that the M. hirsuta population studied diverged from its host < 1 Ma
and appears to be more related to its host population than to some other lineages of
M. sabuleti.
The Italian species M. laurae (Emery) uses as its host M. spinosior Santschi. The host
is probably closely related to M. sabuleti but can be confused with M. scabrinodis by its
relatively wide frons (see key in Chapter 5.1). We assume that M. spinosior represents
a different pre-glacial lineage of M. sabuleti-iike species that survived the glaciations in Italy
(see notes on M. spinosior in Chapter 3.3). Superficially, the queens of M. laurae appear
similar to those of M. hirsuta and we had thought that M. laurae probably evolved rela-
tively recently within the M. spinosior lineage in much the same way as M. hirsuta evolved
from its host. However, the molecular phylogeny of Jansen et al. (2010) indicates that
M. laurae is much older than M. hirsuta, diverging ca. 10 Ma, and appearing to be fairly
basal to the scabrinodis-group (see notes on M. laurae in Chapter 3.3). Thus Jansen and his
co-workers estimate the M. laurae lineage to be intermediate in age between that of M. hir-
suta and the highly specialized social parasite M. karavajevi, which appears to have evolved
about 20 Ma.

1.4. A brief history of the taxonomic investigation of Myrmica in the Old World

1.4.1. Pre 1950

Like all other areas of natural history, ant studies entered a new era with the works of
Carl Linnaeus (1707-1778) who gave the formal name Formica rubra to the "Red Ant" as
recognized by Ray, Gould and probably most common people (Linnaeus 1758). It was
another fifty years before Latreille (1804) established the modern generic name Myrmica,
placing there two species -Formica rubra Linnaeus and Formica Cephalotes Linnaeus (now
transferred to the genus Atta Fabricius). Until William Nylander published his first famous
paper on ants (Nylander 1846a), only nine further species had been attributed to Myrmica
(all described between 1834 and 1841), however, all these nine species were later trans-
ferred to other genera (see list of taxa, excluded from Myrmica, Chapter 3.8). Thus,
Nylander was the first naturalist after Linnaeus to describe "true" Myrmica species (1846a,
b, 1849, 1856, 1857). Six of his species (M. lobicornis, M. lobulicornis, M. ruginodis,
M. rugulosa, M. scabrinodis and M. sulcinodis) represent the most common forms found
in Europe, one (M. granulinodis) is considered in this monograph as incertae sedis in
the genus Myrmica, and two names (M. laevinodis and M. diluta are treated as junior

27
synonyms of M. rubra and M. ruginodis, respectively). In this respect, Nylander can be
credited with defining the basis of the modem taxonomy of Myrmica. However, while he
undoubtedly had a fine grasp of what represents a species, his overall concept of the genus
Myrmica was just as broad as that of his predecessors, for a further 18 of Nylander's
"Myrmica" species were later transferred to other genera.
Unfortunately, Nylander did not assign the Linnaean name rubra to any species, and
for the next sixty years or so the name Myrmica rubra meant different (or any) Myrmica
species, i.e. Red Ant in much the same sense as used by Gould. For example, the French
engineer and insect morphologist Charles Janet (1849-1932), based many of his anatomi-
cal studies made between 1893 and 1907 on M. rubra, though he more specifically referred
to the name M. laevinodis on some occasions. This convention was continued by Wheeler
(1910) in his ground-breaking book, and by Forel (1928) in his book "The Social World of
the Ants"; both seemed happy to consider all Nylander's species as subspecies of M. rubra
even though both authors had themselves described numerous species, subspecies and va-
rieties of the genus Myrmica. Santschi (1931a) corrected Nylander's omission by
synonymizing M. ruginodis with M. rubra. However, Yarrow (1955) inspected Linnaeus'
original material and concluded that M. laevinodis best matched Linnaeus' original
description of M. rubra (Linnaeus emphasized its aggressive nature and fierce sting, and as
Linnaeus' type series included specimens that conformed to Nylander's M. laevinodis, as
well as M. ruginodis and M. scabrinodis), he reversed Santschi's synonymy and
synonymized M. laevinodis with M. rubra.
This caused considerable confusion in the literature because the synonymy was accepted
slowly and reluctantly in most of Europe. On the other hand, M. V. Brian who probably
has published the most papers on M. rubra, created a different type of confusion (discussed
in Elmes 1973b). In short, in Brian's pre 1955 publications the name M. rubra = M. rugin-
odis sensu Santschi, and in his post 1956 publications M. rubra = M. laevinodis sensu
Yarrow. In papers published before 1956, based on studies of Myrmica made in Scotland,
Brian followed Santschi's synonymy using the name "M. rubra" for M. ruginodis. This is
most important to remember when reading his first papers on caste determination by
M. rubra and especially when considering his description (with A. D. Brian) of the forms
M. rubra (= M. ruginodis) macrogyna and microgyna, their study of M. rubra microgyna
must not be confused with later work by Elmes on the microgyne form of M. rubra
(= M. laevinodis, = M. microrubra, see above). When Brian moved to southern England in
1952, he concentrated his seminal studies of caste determination on the locally abundant
"M. laevinodis" populations and by strange coincidence prepared his first papers on these
studies shortly after Yarrow's (1955) revision. Trying keeping abreast of the taxonomic
changes, Brian immediately adopted the name M. rubra for these "M. laevinodis" popu-
lations.
After Nylander's work, relatively little was added to the understanding of Myrmica
taxonomy during the second half of the 19th century; new species appear to be described as
the result of occasional opportunistic study rather than methodical investigation of the
genus. Four more species were described by the greatest of the early myrmecologists,
Gustav Mayr, being two extant from Himalaya and Tibet, and two extinct species from the
late Eocene Baltic Amber. Carlo Emery described another three species: the parasitic
Sifolinia laurae (now transferred to Myrmica) was from Europe, but his two species from
Burma, M. ritae and M. margaritae, represent an important group within the genus (see
Chapter 2.2). Frederik Wilhelm August Meinert (1833-1912) described his one and only

28
Myrmica species, M. sabuleti, which is common throughout Europe, while the great Au-
guste Forel described only one Myrmica species before 1900, the parasiticM. myrmicoxena.
As a result, at the start of the 20th century, 130 taxa had been described in the genus Myr-
mica in the Old World; ten of these names are nomina nuda, 91 have been transferred to
other genera or excluded from Myrmica, 13 are considered as synonyms of other Myrmica
species, and only 16 of the names are still recognized as good species.
By the end of 19th and early in the 20th century some regional Keys that included Myr-
mica of Europe and British India, had been published (e.g. Mayr 1861; Forel 1874, 1903;
Bingham 1903; Ruzsky 1905), but the most comprehensive for that time was Emery's revi-
sion of the Palaearctic ants (the part, concerning Myrmica - Emery 1908a, e).
The first half of the 20 th century up to the time of the end of World War II (1945) was
a time of increased worldwide myrmecological study, especially taxonomical investigations
(see Bolton et al. 2006), and a considerable number of new forms and species were added
to the genus Myrmica (Fig. 1). By this time, the concept of the genus more or less con-
formed to our present understanding, so that most of the described taxa truly belonged to
the genus. Two authors stand out from this time: Forel, who between 1902 and 1909 de-
scribed twelve good Myrmica species from Himalaya, Middle Asian Mts. and Japan (plus
five other names now consider as synonyms), and the great Russian myrmecologist Michail
Dmitrievich Ruzsky, who documented the ant fauna of the former Russian Empire and
Tibet. Between 1895 and 1946, Ruzsky described 28 taxa of Myrmica, twelve of which are
still considered as good species. Significant contributions were also made by six other
authors (see biographies below): Karawajew (1916, 1926b, 1931b) described four Myrmica
species from the Middle Asian Mts., Ukraine and Siberia; Bondroit (1918, 1920a), added
four good species from Europe; Finzi (1923,1926) added four European species; Menozzi
(1925, 1930, 1939) described three species from Europe, Siberia and Himalaya; Santschi
(1931) added two species from Europe and Asia Minor; Wheeler (1928a, b, 1929,1930a) de-
scribed three species from Taiwan and China besides his work on New World ants. Several
regional Keys for the identification of ant species, including Myrmica, were published during
this period (Forel 1915-for Switzerland; Donisthorpe 1915a-for Great Britain; Emery 1916
- for Italy; Bondroit 1910, 1918 - for Belgium and France; Karawajew 1927b, 1934 - for
Ukraine; Stitz 1914,1939 - for Germany; Novak and Sadil 1941 - for Central Europe).
There were three significant publications before World War II that were especially
devoted to the genus Myrmica. The first was provided by Finzi (1926), who summarized the
known data for the European species, provided an identification Key to the species and
infraspecific forms, redescribed many taxa, and gave drawings of most species. Five years
later, Santschi (1931b) published a paper that discussed several unresolved problems and
made some important improvements to the taxonomy of the genus. The third significant
contribution to Myrmica taxonomy was made by Arnoldi (1934), who considered the fauna
of the FSU using morphometries in a way that has become increasingly important in
Myrmica taxonomy (see section 1.5). In addition, Weber made a first revision of the genus
Myrmica in a 1935 PhD thesis, which was not published until after World War II (Weber
1947, 1948, 1950). In these papers he revised the Nearctic Myrmica and provided a com-
prehensive taxonomic review of the Old World species with a description of two new species
from Himalaya. By 1950, the number of good Old World Myrmica species had increased to
65 (Fig. 1), a further 94 names, described and attributed to Myrmica in the period 1900
to 1949, are now considered as synonyms of other Myrmica species, or have been trans-
ferred to other genera, or are unavailable names, nomina nuda, incertae sedis etc.

29
1.4.2. Post 1950

. The first important paper on Myrmica taxonomy published in the second part of the
20th century, was a revision of the Central European fauna made by Sadil (1952), in which
he described one good species (and four more taxa now considered as synonyms). It is in-
teresting to note that between 1952 and 1969 only four more species were described
(M. arnoldii Dlussky, 1963, M. bibikoffi Kutter, 1963, Aphaenogaster gigantea Colling-
wood, 1962, and Sifolinia lemasnei Bernard, 1967; two latter species now transferred to the
genus Myrmica). The well-known monograph on French ants, that included a comprehen-
sive section on the genus Myrmica, was published by Bernard (1967).
In the 1970's and 1980's a few more species were added to the Old World Myrmica
fauna, so that by 1990 a total of 81 good species were known. However, this was a period
of considerable synthetic activity and several important revisions of the genus Myrmica
were published. First, there was those of Arnoldi (1970, 1976), dealing with Myrmica of the
European part of the former USSR and Middle Asia, where five good species were described
and identification Keys were provided. In 1973 the Swiss entomologist Heinrich Kutter
(1896-1990) reviewed the parasitic species of Myrmica and its "satellite" genera (Sifolinia
Emery, Symbiomyrma Arnoldi and Sommimyrma Menozzi) and subsequently the great
English ant taxonomist Barry Bolton (born in 1945) described a new socially parasitic
species from Himalaya, and more importantly synonymized with Myrmica the socially-
parasitic genera Sifolinia, Symbiomyrma, Sommimyrma and Paramyrmica Cole, and the
subgenus Dodecamyrmica Arnoldi (Bolton 1988). Henri Cagniant (1970) described a new
socially-parasitic species from Algeria, and Elmes (1978) described another social parasite
from England. Finally, by the end of the period Kupyanskaya (1986a, b, 1990) had described
six new species and had written a useful monograph on the ants of the Far East of Russia,
and Seifert (1987) had described one new species from Caucasus, and had revised
the Myrmica of Europe and Caucasus, making several important synonymies (Seifert 1988).
Other significant papers, monographs, reviews and regional Keys relevant to Myrmica
taxonomy include those of Kutter (1970, 1977), Bolton and Collingwood (1975), Tarbin-
sky (1976), van Boven (1977), Collingwood (1978, 1979), Arnoldi and Dlussky (1978),
and Agosti and Collingwood (1987a, b).
In the 1990's some data concerning European and Far Eastern Myrmica were published
in various monographs and papers (e.g. Atanassov and Dlussky 1992; Seifert 1996; Rad-
chenko et al. 1997), but the most important was the revisions of Myrmica, mainly of the
FS U, made by Radchenko (1994a-f). Although the data and ideas in that revision have been
added to, modified and refined as our current collaboration progressed, it was our starting
point for the work that has culminated in this monograph (see Preface). We discovered in
collections a very rich Myrmica fauna from the Himalaya and South-East Asia and described
it in papers on the ritae species group and the Myrmica fauna of the Himalaya and Taiwan
(Radchenko and Elmes 1998, 1999a, b; Elmes and Radchenko 1998). In total, we described
20 new Myrmica species in these papers, including one very peculiar species from Taiwan
(Elmes and Radchenko 1998). In the 1990's, the Spanish myrmecologist Xavier Espadaler
described a new species from Morocco. Therefore, in the second half of the 20 th century
the number of good Old World Myrmica species had increased by about 60% to a total of 106.
After 2000 we were fortunate to be given access to considerable amounts of unidenti-
fied material (see Preface), which allowed us to expand our investigation of the Old World
Myrmica. We were able to revise several other species groups and describe the Myrmica

30
fauna of many previously poorly studied regions: China, Korea, Vietnam, Turkey, Sicily etc.
(Radchenko and Elmes 2001a, b, 2002, 2003a, b, 2004, 2009a, b, e; Radchenko et al. 2001,
2008; Elmes et al. 2001, 2002; Radchenko, Elmes and Alicata 2006; Radchenko, Elmes and
Viet 2006; Elmes and Radchenko 2009; Elmes et al. 2009). We were also able to revise the
fossil Myrmica from the late Eocene European ambers (Radchenko et al. 2007). At the same
time several important additions to the taxonomy of European Myrmica were provided by
Seifert (2000, 2002, 2003, 2005; Seifert et al. 2009), and several new identification Keys
for Europe and East Asia were published (Czechowski et al. 2002; Imai et al. 2003; Rad-
chenko et al. 2004; Radchenko 2005; Seifert 2007). In this first decade of the 2 1 s t century
41 new Myrmica species have been described from the different regions of Eurasia (39 of
them by us).
Consequently, 147 species oìMyrmica are recognized in this monograph: 142 are extant
species distributed in the Old World (one of them, M. kozakorum, is described in this mono-
graph, see Chapter 3.3), and five are fossil species from the European late Eocene ambers
(see Chapter 3.4.) Between us, we have added 60 species (or more than 40%) to the
known fauna of the Old World, the majority of our new species living in southern
mountain regions, especially in the subtropical and tropical mountain areas of Eurasia.
One (M. emeryi) is the first Myrmica to be described from the Southern Hemisphere (see
Chapter 3.3). These species emphasize the previously unrecognized richness and diversity
within the genus Myrmica and we are certain that many more species await discovery,
especially in the Himalayan region, southwest China and South-East Asia.
Extrapolating to the New World we expect that after a cardinal revision (see Chapter
1.1) the number of good species in North America should be at least double the 30 or so
currently recognized species. Finally, we readily admit that apart from taxa still to be dis-
covered, there are many unresolved problems in the details of the taxonomy of Myrmica in
the Old World that are not addressed by this monograph. We hope that our work will be of
some help to future generations of myrmecologists when they tackle these questions.

Figure 1. The total number of "good" Old World Myrmica species, described from 1758 until 2010.

31
1.5. Morphometries and Myrmica taxonomy

1.5.1. History of measurement of Myrmica

Early taxonomists often gave an indication of an insect's size when they first described
it. Usually this was the whole body length, but sometimes they measured the length of
various sections of the body. The first use of morphometries (measurements of size of
various body parts) of the genus Myrmica (perhaps the first use in ant taxonomy) was made
by the Dutchman August Stärcke (1880-1954) who made six measurements to compare
various forms of Myrmica lobicornis (Stärcke 1927). He measured head-width (HW,
including the eyes), the minimum distance between the frontal carinae (frons-width in our
modern terminology - FW; see Chapter 1.5.3 for measurements and indices used by us), the
maximum extent of the frontal laminae (our frontal lobe-width - FLW), the length and
height of the petiole, and the length of the propodeal spines (measured in two ways, the
second being the same as our ESL). Stärcke also introduced the concept of comparing
ratios, specifically the Frontal index, which he defined as head-width / frons-width (we
prefer the reciprocal of this).
Konstantin Vladimirovich Arnoldi was also one of the first myrmecologists to use mor-
phometries: first in a paper on Stenamma Westwood (drafted in 1927) and later as an aid to
identification of Myrmica ants (Arnoldi 1934). He used four measurements: FW and FLW,
head-length (HL) and HW measured in front of the eyes (i.e. excluding the eyes). In the case
of Myrmica (but not all ants) it makes little difference whether HW is measured immedi-
ately in front or immediately behind the eyes. Stärcke maintained that including the eyes re-
duced errors due to orientation of the head, a view strongly supported by Bernhard Seifert,
on the other hand including the eyes arguably confounds two separate characters (head-
width and eye size) and other authors (including Arnoldi, Sadil, Brian and Brian, and us -
see Chapter 1.5.3) measured HW excluding the eyes. This minor rivalry in methodology is
unlikely to be resolved in the near future (see also Espadaleri: 1981b).
Arnoldi (1934) placed much greater emphasis on the value of ratios of the measure-
ments compared with Stärcke, using Head-index = (HW/ HL) x 100, Frons-index = (FW
/HW) x 100 and Frontal-lobe-index = (FW/FLW) x 100. Later Arnoldi (1970, 1976) used
the reciprocal of these (i.e. Head-index = HL/HW, Frons-index = HW/FW and Frontal-
lobe-index = FLW/FW). These indices (or their reciprocals, though usually not scaled by
100) were adopted generally in Myrmica taxonomic studies and identification Keys (e.g.
Radchenko 1994c) and they are still used today. Sadil (1952) was the next who used
morphometries in a major revision of Myrmica, however, despite being a mathematician,
he used the same measurements as Arnoldi and did not develop the tool significantly.
A few years earlier, Brian and Brian (1949) used morphometries more imaginatively to
separate M. ruginodis from M. laevinodis {- M. rubra sensu Yarrow).
Elmes (1978) introduced a new more statistical approach when he described the social
parasite M. hirsuta, using multivariate Canonical Variate Analysis (CVA) of a priori de-
fined groups; this technique had long been known in theory but became a practical possi-
bility at that time due to the development of modern computers. Elmes assembled three
groups of specimens based on separation by traditional subjective morphological features,
being M. hirsuta, the host M. sabuleti and M.. scabrinodis (queens of M. hirsuta superficially
resemble those of M. scabrinodis and males superficially resemble M. sabuleti). He
measured 10 body parts and count of the number of hairs on the petiole. CVA showed that

32
on the basis of the features chosen, M. hirsuta queens and males were statistically very dif-
ferent from the other two groups and could easily be discriminated by morphometries alone;
Elmes illustrated his results graphically. He considered that this technique greatly rein-
forced the case for the separation and description of M. hirsuta as a separate species. Later
Elmes showed how this technique could be applied to other questions of Myrmica
taxonomy (e.g. Elmes and Thomas 1985). Analysis of morphometries could also be used to
apportion variation geographically (Elmes and Clarke 1981), a Hierarchal Analysis of
Variance illustrating the effects of different reproduction strategies between the microgyne
and normal forms of M. ruginodis populations.
Since the 1980's Bernhard Seifert has become more or less "The King of Morphomet-
ries" using them in many of his numerous publications on ants including his major revision
of European Myrmica (Seifert 1988) and papers on various Myrmica species (e.g. Seifert
1984,1989,1993,2000, 2005; Seifert et al. 2009). In general, Seifert prefers to illustrate his
results by mathematical discrimination functions rather than graphically.

1.5.2. A brief critique of the value of morphometries

The use of morphometries and associated statistical analysis can be a useful addition to
the taxonomist's tool kit, but it is not the only tool, furthermore considerable caution needs
to be taken when using measurements and indices. Shortly before his death Stärcke (1951)
wrote a critique of Brian and Brian's (1949) use of morphometries; although some of his
criticisms appear rather pedantic, he did make several important general points.
The first concerned errors: Stärcke was concerned about the accuracy of Brian and
Brian's measurements. If X is a measured parameter and T is the true value then: X = T +
Er + Es where Er = a random error and Es = a systematic error. In the case of an ant mor-
phometries the most frequent random errors are firstly due to rounding up or down to the
nearest measuring-graticule unit, secondly errors that result from parallax due to orienta-
tion of the specimen, then misreading the graticule and finally chance transcription errors
when recording the observation. Random errors if truly random should not affect the over-
all mean value of X but slightly increase the variance about the mean, slightly reducing
the value of X as a discriminator (alone or in combination with other variables). Related to
this is pseudo-accuracy resulting from scaling-up graticule calibration, Stärcke objected to
Brian and Brian giving the value of one graticule unit to 100th of a micron (although he did
the same thing in his 1927 paper) because there must be some small discrepancies between
the engraved graticule scale units and certainly those generally available in non-specialist
laboratories will not be accurate to 100th of a micron. However, if one is measuring to the
nearest graticule unit the average error must be ± V2 a unit (in Stärcke's case half of
0.01815 « 0.009 mm) so there is no benefit in giving any individual measurement to an
accuracy > 1 scale unit (0.01 mm at magnification x 100 or 0.02 mm at x 50, using the
most common graticule scale of 1 mm divided into 100 units).
Systematic errors are a much greater problem because by definition they will affect the
value of the mean of variable X. The greatest systematic errors occur between individual
observers. For example, one observer might consistently orientate his specimen differently
to another observer which might induce say a consistent 5% difference in the two's values.
Stärcke (1951) disagreed with the way Brian and Brian measured ESL (even though it
seems to more or less accord with his method - figure 3, loc. cit.), the problem being exactly
how they determined the basal ends of the spines. Stärcke also did not approve of the way

33
Brian and Brian measured head-width (see below). Systematic errors can occur also when
the same observer uses different equipment or makes measurements at a later date. How-
ever in general such errors by an experienced observer are relatively unimportant compared
to differences between observers.
About 30 years ago we made a simple test of observer errors (unpublished) : we asked
four observers to measure 11 characters (10 of them were measurements plus a count of the
bristles on the petiole) on the same set of 40 workers of M. scabrinodis, each observer using
his favourite microscope set to the same magnification (x 50 for four parameters and x 100
for the other six). Two of the observers measured the same ants again some days later,
the maximum difference between the two sets expected by rounding graticule readings was
± 1 unit or ± 0.02 mm and 0.01 mm for the two levels of magnification (or ± 0.014 mm
averaged over all ten morphometries). The second set of measurements made by
observer 1 (experienced in measuring ants) deviated from his first set of measurements by
an overall absolute average of 0.011 mm (i.e. within the random rounding errors expected
from graticule readings) and it seemed probable that this level of error represented a com-
bination of random errors (e.g. due to specimen orientation, transcription etc. as well as
graticule error). In subsequent comparisons we took the best estimate of the true mean for
any morphometric, to be the average of observer l's two sets of measurements.
In her first set of measurements observer 2 (a myrmecologist not very experienced in
measuring) deviated from the estimated true means (based on observer 1) by an absolute
average of 0.023 mm and one hair on the petiole bristle count indicating some systematic
error. However, in her second set, after reading the instructions again and consulting
observer 1, she deviated by only 0.006 mm and 0.5 hairs respectively, within the random
error of observer 1 and indicating that a little experience soon reduces systematic error.
The other two observers both had similar errors to observer 2 on her first try: observer 3
(another myrmecologist not very experienced in measuring) deviated 0.019 mm and
1.3 hairs, while observer 4 (an experienced entomologist but not an ant specialist) deviated
by 0.018 mm and 1.1 hairs. Generally, observers 2 and 3 underestimated measurements
and petiole hair number compared to observer 1, whereas observer 4 overestimated values
(i.e. the systematic errors associated with the observers being negative and positive
respectively), which probably represented a combination of errors due to equipment dif-
ferences and systematic orientation error. As predicted by Stärcke, the systematic errors
were greatest and most directional for features that were somewhat difficult to define
and-or orientate consistently, mainly ESL, scape-length and postpetiole height. In general,
one can conclude from this simple test that when measurements are made by the same
experienced observer, systematic errors are likely to be small and not greatly affect any
resulting discriminations. In other words, morphometrical analysis can be a useful a re-
search tool for an individual scientist comparing features of species.
Much greater care must be taken when mixing results recorded by two or more
observers, with some features being much more prone to generate systematic observer error
than others. The practical consequence of the latter is that when morphometries (or indices
calculated from them) are used in Keys etc., the authors should try to avoid using those fea-
tures whose definition is more ambiguous and should not rely entirely on indices (or other
discriminants) without support from more traditional morphological features. One of Stär-
cke's (1951) criticisms of Brian and Brian (1949) was that they had not paid sufficient at-
tention to more subjective morphological criteria, shape of spines, degree of rugosity of the
petiole and thorax etc. In the particular case Stärcke seemed to miss the fact that they sent

34
samples to the leading English ant taxonomist of the time, Horace Donisthorpe (1870-1951),
who identified the specimens using these "traditional" features. Nevertheless, Stärcke makes
an important general point that can be summarized as: just because one can find significant
morphometric differences (or not) between two populations it does not necessarily prove
(or disprove) the hypothesis that these belong to different (or the same) species.
Finally, Stärcke (1951) was worried by how the statistical differences Brian and Brian
observed should be shown and interpreted. Brian and Brian (1949) plotted HW against
ESL and calculated linear regressions and they could have compared the regressions sta-
tistically but did not do so because they were so obviously different (figure 2, loc. cit.). In
theory, if one plots one of the measurements that we use (see Chapter 1.5.3) against another
the best descriptor of the relationship between the variables is the geometric mean slope (i.e.
the geometric mean of slopes X Y and Y X) which usually hardly differs from the ratio of
the two variables, especially when the regression intercept is not statistically different from
zero (usually the case with our data). The indices used by us, like those of the older authors,
are simple ratios between two morphometries, although not strictly statistically valid one
can get a good idea of the value of any ratio for separating two species by comparing the
means and their standard deviations. A robust index is one that clearly separates a pair of
species and is comprised of two morphometries that can easily be measured by a non-
specialist using a fairly simple microscope (i.e. usually maximum magnification x 50);
the index must be able to discriminate most individuals even when there is moderate sys-
tematic observer error. In general, the original indices favoured by the old authors fit this
criterion (e.g. Frontal index, Frontal lobe width index and Propodeal spine length index).
So, what is the best way to compare morphometries? Should it be done graphically, or
should one compare mean values (including indices, see above)? Should one use the
powerful statistical tools now available or is simply "eyeballing" the data sufficient? To
what extent should one worry about allometry? (see the various publications of Bernhard
Seifert loc. cit. for discussion of this problem). The answers vary according to whether the
authors are using morphometries as a research tool or as an identification aid. In the first
case the author might be investigating whether material assigned to a particular species in
fact includes specimens from two or more different but morphologically very similar species,
in which case graphs, indices and discriminant functions and other statistical techniques
might all aid the author's presentation of the results (e.g. see Elmes and Radchenko 2009).
On the other hand, statistics and discriminant functions can be quite daunting especially
when non-specialists encounter them in identification Keys, often users discard such Keys
in favour of simpler ones, especially when faced with the prospect of measuring perhaps
10 or more variables and making a mathematical calculation. We consider that at most a few
simple robust indices should be used in Keys.

1.5.3. Measurements and indices used in this monograph

In our previous papers concerning Myrmica taxonomy that are cited in this monograph,
we used all or some of the following measurements and indices (Fig. 333 in Chapter 5 in-
dicates how we make the measurements) :
HL - maximum length of head in dorsal view, measured in a straight line from the ante-
rior point of clypeus (including any carina or ruga, if they protrude beyond the an-
terior margin) to the mid-point of occipital margin;
HW - maximum width of head in dorsal view behind (above) the eyes;
FW - minimum width of frons between the frontal carinae;

35
FLW - maximum distance between the outer borders of the frontal lobes;
SL - maximum straight-line length of scape from its apex to the articulation with condy-
lar bulb;
AL - diagonal length of the alitrunk (seen in profile) from anterior end of the neck shield
to the posterior margin of propodeal lobes (workers) and from the most antero-
dorsal point of alitrunk to posterior margin of propodeal lobes (queens and males);
HTL - maximum length of hind tibia;
PNW - maximum width of pronotum in dorsal view (workers) ;
PL - maximum length of petiole in dorsal view, measured from the posterodorsal margin
of petiole to the articulation with propodeum; the petiole should be positioned so
that measured points lay on the same plane;
PW - maximum width of petiole in dorsal view;
PH - maximum height of petiole in profile, measured from the uppermost point of the
petiolar node perpendicularly to the imaginary line between the anteroventral (just
behind the subpetiolar process) and posteroventral points of petiole;
PPL - maximum length of postpetiole in dorsal view between its visible anterior and pos-
terior margins;
PPW - maximum width of postpetiole in dorsal view;
PPH - maximum height of postpetiole in profile from the uppermost to lowermost point,
measured perpendicularly to the tergo-sternal suture;
ESL - maximum length of propodeal spine in profile, measured along the spine from its tip
to the deepest point of the propodeal constriction at the base of the spine (note that
in some our earlier papers we followed earlier authors by calling the propodeal
spines "Epinotal Spines" - subsequently we retained the abbreviation ESL etc. to
avoid confusion) ;
ESD - distance between the tips of propodeal spine in dorsal view;
SCW - maximum width of scutum in dorsal view (queens and males);
SCL - length of scutum+scutellum in dorsal view (queens and males);
AH - height of alitrunk, measured from upper level of mesonotum perpendicularly to the
level of lower margin of mesopleuron (queens and males).
We calculate the following indices from these morphometries (simple ratios - see Chap-
ter 1.5.2.) but in the Keys (Chapter 5) we used a reduced number, usually CI, FI, FLI, SI1,
SI2, PIj, PPI4, ESLI and ESDI.
Cefalic (CI) = HL/HW;
Frontal (FI) = FW/HW;
Frontal-lobe (FLI) = FLW/FW;
Scape-1 (SIj) = SL/HL;
Scape-2 (SI2) = SL/HW;
Petiolar-1 (PlJ = PL/PH;
Petiolar-2 (PI2) = PL/HW;
Petiolar-3 (PI3) = PW/HW;
Postpetiolar-1 (PPIJ = PPL/PPH;
Postpetiolar-2 (PPI2) = PPH/PPW;
Postpetiolar-3 (PPI3) = PPW/PW;
Postpetiolar-4 (PPI4) = PPW/HW;
Propodel spine-length (ESLI) = ESL/HW;
Propodel spine-distance (ESDI) = ESD/ESL;
Alintrunk (AI) = AL/AH;
Scutum (SCI) = SCL/SCW.

36
1.6. Biographies of the authors

We thought that readers might find it interesting to have some biographical details of the
main authors of the Old World Myrmica species. We therefore give details of persons who
have described two or more good species (see Table 1.1), or in the case of Sadil has a major
review of Myrmica. For the purposes of this analysis we credit species to the author who first
described a good species even though they unintentionally used an unavailable name for the
species, rather than "crediting" the species to the author of the replacement name (see e.g.
Kupyanskaya below).

1.6.1. Biographies of the authors of this monograph

A. G. Radchenko
(born in 1955)

Alexander Grigorievich Radchenko was born on 28 th

January 1955 in Kiev, Ukraine, the son of a school-
teacher, he is married to Tamara. He trained as a school-
teacher specializing in geography and biology, gradu-
ating from the Kiev State Pedagogical Institute in 1977.
After a short spell as a teacher, he worked in the Insti-
tute of Molecular biology and genetics of the Academy
of Sciences of Ukraine, where he started his research ca-
reer by studying viruses of insects. In 1980 he was ap-
pointed as a PhD student in the Schmalhausen Institute
of Zoology of the Ukrainian Academy of Sciences
(Kiev), successfully defending his PhD thesis in 1985.
In 1999 Radchenko was awarded a Doctor of Science
degree for a thesis entitled "The ants of Palaearctic", and in 2002 he was awarded a full
professorship in the Kherson University (Ukraine).
The main theme of Radchenko's professional career has been the study of systematics,
taxonomy, faunistics and ecology of ants together with the associated problems of zoo-
geography, origin and evolution of the Palaearctic myrmecofauna. He has described more
than 120 new species and five new genera of the extant and extinct ants, and revised many
Palaearctic ant genera. His published papers and monographic revisions currently total
about 200; these have resolved several complicated questions concerning the taxonomy,
systematics and biogeography of ants of southern and eastern regions of the Eurasian con-
tinent. He has taken part in many scientific expeditions, collecting ants throughout the ter-
ritories of the former USSR and many other countries, e.g. Vietnam, Korea, China, India,
Turkey and most European countries. His collection of ants now comprises many tens of
thousand specimens and is deposited in the Schmalhausen Institute of Zoology, Kiev, which
also houses the famous memorial ant collection of V. A. Karawajew. Taken together, these
collections form one of the richest ant collections in Europe.
Radchenko started an especial investigation of the genus Myrmica about 20 years ago,
and this has been a significant part of his work since the late 1990's,'cooperating with
G. W. Elmes (see Preface), culminating with the publication of this monograph. He has
described 59 (40%) of the Old World Myrmica species (55 of them jointly with Elmes and
other co-authors).

37
 G. W. Elmes
(born in 1943)

Graham Wakely Elmes was born on 7th November 1943

in the small town of Wareham, Dorset, England, the son
of a Post Office clerk. He is married to Jenny and they
have two married daughters, Elizabeth and Rosanna.
From 1961 to 2003 he worked as a British government
research scientist first as an assistant to Dr. M. V. Brian
who was head of the British Nature Conservancy's
Furzebrook Research Station. After completing a first
degree, he started his independent research career in
1970 as a scientist in the Natural Environment Research
Council. He was awarded a University of London
PhD in 1975 for a thesis entitled "Population studies on
the genus Myrmica", and a Doctor of Science degree
from the London University in 1993 for his contribution to ant ecology. Currently he is an
Honorary Research Fellow with NERC's Centre for Ecology and Hydrology, Wallingford
laboratories. He has published more than 130 papers on three related areas of research:
1) the population ecology of Myrmica (especially the behaviour and evolution of their
social parasites); 2) the ecology, population dynamics and behaviour of Phengaris (= Ma-
culinea) butterflies, which have larvae that parasitize Myrmica ant nests; 3) the taxonomy
of Myrmica species mostly in collaboration with A. G. Radchenko. He is also especially
interested in the investigation of cryptic species and varieties the latter being important in
the interpretation of field studies arising from his primary ecological interests.

1.6.2. Biograhies of other authors (ordered chronologically)

W. Nylander
(1822-1899)

William Nylander was born in Oulu, Finland on 3rd Janu-

ary 1822. His father came from a prosperous merchant
family and was a city alderman. In 1839 he graduated
from the gymnasium in Âbo (now Turku) and entered
the University of Helsinki where he is said to have been
a brilliant student, passing examinations as a candidate
in philosophy in 1843 and becoming a Doctor of Medi-
cine and Surgery in 1847 at the young age of 25. How-
ever, his boyhood interest in natural history soon came
to the fore and he neglected his medical career in favour
of botanical and entomological studies. His first interest
was in Hymenoptera and he published several papers in this field; the most notable, in the
context of this monograph, being his papers in 1846, 1849 and 1856, in which he estab-
lished the modern treatment of the genus Myrmica. However, he is best remembered as

38
a lichenologist. He became Helsinki's first Professor of Botany and played an important
role in establishing botanical research in Finland. Nylander never married. In 1850 he
moved to Paris and continued his lichenological researches there. Sadly, he became
something of a recluse being estranged from most of his contemporary lichenologists as
a result of his "vitriolic abuse" and consequently finding it hard to get his work published.
He withdrew from the academic world and died at his desk in Paris, on 29 th March 1899,
aged 77 (Vitikainen 2001; Lawrence 2008).

G. L. Mayr
(1830-1908)

The Austrian, Gustav L. Mayr was born on 12th October

1830 in Vienna. He held chairs in both Pest, Hungary
and in the University of Vienna. Not much seems to be
known about his private life. He published widely on
ants from 1853 to 1907 and some consider his works to
be among the first written in "proper scientific style".
Forel in 1874 wrote about Mayr (paraphrased): "Un-
doubtedly Mayr is an important author. He has raised
the study of ants to the highest level of entomological
knowledge by his remarkable perception of genera, the
recognition of zoological characters and the precision of
his work". Mayr was a great man in the context of ant
taxonomy generally although he did not contribute much to the taxonomy of the genus Myr-
mica. He can be credited with providing the basis for modern ant taxonomy especially at
the generic level describing about 60 genera which are mostly still valid names, and more
than 400 species. Mayr died in Vienna on the 14th July 1908, aged 77 (Anonymous 1908).

A. Forel
(1848-1931)

Auguste-Henri Forel was born on 1st September 1848

in La Gracieuse, to the west of Lausanne, Switzerland.
The son of a middle class family, he studied medicine
and was a psychiatrist by profession. He married Emma
Steinheil in 1883 and they produced a large family. In
1879 he was a professor of psychiatry at the University
of Zurich Medical School and was running the Burghöl-
zli asylum. He published papers on insanity, prison re-
form, social morality and the brain structure of humans.
More controversially he was a supporter of eugenics,
though in later life he abandoned these views and joined
the Bahá'í Faith. He became interested in ants as á boy,

39
when he read Huber's book on ants and he maintained this interest throughout his life
becoming one of the World's most notable myrmecologists as evinced by the use of his image
on a 1000 Franc Swiss banknote. By the time he died, at the age of 83, on July 27lh 1931,
he had published a five-volume book and more than 500 papers on ants, having described
many hundreds of new species of ants in many genera, including 13 good species
of Old World Myrmica and many more varieties. His fame in both medicine and
myrmecology has resulted in many published tributes and obituaries (e.g. Donisthorpe
1931; Steck 1948; Cherix 1980) and most recently his life and contribution to myr-
mecology has been reviewed by Sleigh (2006). Forel had strong views on morality
and influenced by his wife, he was life-long abstainer from alcohol. These attitudes
probably contributed to the somewhat tempestuous relationships he had with most of his
myrmecological peers, though he seemed to have maintained a long friendship with
Carlo Emery.

C. Emery
(1848-1925)

Carlo Emery was born in Naples, Italy on 25 th October

1848. He was sufficiently well-liked and important to
have many obituaries, including ones authored by Au-
guste Forel and William Wheeler who were the world's
preeminent myrmecologists at that time (Forel 1925;
Wheeler 1925), the following personal information is
taken from these. Other Italian obituaries and tributes
include those of Ghigi (1925), Grandi (1925), Gestro
(1926), Conci and Poggi (1996). The affection with
which he was regarded is indicated by the fact that 56
ant species and infraspecific forms were dedicated to
Emery, more than to any other myrmecologist in history.
His parents were Swiss émigrés. He married an
Italian wife (Rita) and had two children (Lucia and Luigi). Like many early zoologists, he
first trained in medicine, specializing as an ophthalmologist before turning to biological
studies. From 1878 to 1881 he was Professor of Zoology at the University of Cagliari,
Sardinia before moving to the Chair of Zoology in the University of Bologna, where he re-
mained until about 1920. His early research interests were broad, he published on fish and
reptiles, but he soon specialized in entomology, publishing about 300 papers on ants, de-
scribing about 130 new genera, three of which became type genera of subfamilies (Aenic-
togitinae, Aneuretinae, Leptanillinae), and more than 1100 species and infraspecific forms
of ants. His important collection, which contains many type specimens from other spe-
cialists is housed in the Museo Civico di Storia Naturale "Giacomo Doria" in Genoa, Italy.
He is perhaps most famous among his generation of myrmecologists for his contribution on
Formicidae in Wytsman's "Genera Insectorum". Emery was apparently a polyglot, fluent in
most European languages, besides being a skilled draftsman. He suffered a severe stroke in
1906 but trained himself to draw with his left hand. He was a long time friend of Forel who
tended him during his illness; Forel himself suffered a stroke six years later. Emery died in
Bologna on 11 th May 1925, aged 76.

40
 M. D. Ruzsky
(1864-1948)

Mikhail Dmitrievich Ruzsky was born in 1864 in

Gdovsky Uezd (District) of St.-Petersburg Gubernia
(Province) and died on 29 th April 1948, aged 84, in
Tomsk. His family moved to Simbirsk, where he gradu­
ated from the gymnasium in 1884. He enrolled in the
Kazan' University graduating in 1888, successfully de­
fending his diploma work on ichthyology. He began his
distinguished academic career in the same university
first as an assistant in the zoological laboratory, and
then progressing to Custodian of the Zoological Mu­
seum, Assistant-professor and finally Head of the Cathe­
dra of Zoology. In 1913 Ruzsky was simultaneously
elected as Professor of both the Kiev and Tomsk (West Siberia) Universities, electing to go
to Siberia rather than to Ukraine. For the next 35 years he was Head of the Cathedra of
Zoology of the Tomsk University (Berezhkov 1937; Garanin 2002; Krasilnikov 2003).
Ruzsky was a very able general zoologist with an encyclopedic knowledge of many groups:
he studied fishes, birds, mammals, amphibians and reptiles, publishing papers and several
monographs on these taxa during the first half of the 1890's and again later in the 1920's and
1930's. However, outside of Russia Ruzsky is best known as one of the great myrmecologists
of the first half of the 20th century. His first paper on ants was published in 1895, and only 10
years later he published his renowned monograph "The ants of Russia" ("Муравьи России"),
which was first published in two volumes (1905, 1907). For its time it was one of the most
comprehensive books on ants and furthermore, in many respects it is still relevant today, de­
scribing a new ant genus and more than a hundred species and infraspecific names (many now
treated as good species) from throughout the Russian Empire. There is no doubt that Ruzsky
provided the foundation for future myrmecological studies in Russia and states of the former
Soviet Union. Publication of this book won Ruzsky the highest award of the Russian
Academy of Sciences - the K. Ber's Prize; it is interesting to notice that he was only the third
scientist to be awarded this prize, following the great evolutionary biologists 1.1. Mechnikov
and A. O. Kovalevsky.

V. A. Karawajew
(1864-1939)

Vladimir Afanasievich Karawajew (also spelled as Ka­

ravaiev or Karavajev) was born on 9 th March 1864 in
Kiev, Ukraine, the son of a doctor. He graduated from
the Kiev University in 1890 and started his scientific ca­
reer as an assistant in the Cathedra of Zoology in that
University. His early scientific interests included the em­
bryology of hemipteran insects and studies of several
groups of marine invertebrates before he specialized in
myrmecology. In 1898 and again in 1912-1913 he made
expeditions to Java and other Indonesian Islands, where

41
he collected a very rich zoological material on various groups of animals, including ants.
In 1919 Karawajew was one of the founders of the Zoological Museum of the Ukrainian
Academy of Sciences in Kiev, he became its first Custodian, donating his huge private zo-
ological collections. He became Director of the Museum in 1927 and remained so until his
retirement in 1934, thereafter until his death he remained a senior scientist attached to the
Institute of Zoology (Paramonov 1941). Karawajew was married but we know little else
about his personal life, he was spared the traumas of World War II, dying in Kiev on 7th
January 1939, aged 74.
Apart from Indonesia Karawajew collected ants in various European countries, North
Africa and the former Soviet Union, especially Ukraine, Crimea, Middle Asia and Caucasus.
As a result, he left the next generations of entomologists a large collection, in which all
specimens are splendidly mounted, labelled, determined and arranged. He published more
than 50 papers on taxonomy of ants from the different parts of the Palaearctic and Orien-
tal Regions. His two-volume monographs "The ants of Ukraine" (1934, 1936) formed the
basis for later Ukrainian myrmecological studies.

W. M. Wheeler
(1865-1937)

William Morton Wheeler was born in Milwaukee, USA

on 19th March 1865. We have found several obituaries
written shortly after his death (Carpenter 1937; Ice-
lander and Carpenter 1937; Parker 1937; Bruch 1937)
from which the following information is drawn. See also
the account of his career in Sleigh (2006). By his own
account he was rather a naughty child so his parents
sent him to a strict German school where he gained an
interest in nature. He graduated from the German-
American Normal College in 1884 and worked a year as
an assistant in the Ward's Natural Science Establish-
ment, which was a sort of the museum factory. He
taught German and physiology in the Milwaukee High School (1885-1887), and then be-
came Custodian of the Milwaukee Public Museum (1887-1890). He published on insect
embryology and continued to do so when he became Fellow and Assistant in Mor-
phology at the Clark University, being awarded a PhD in 1892. Wheeler transferred to the
University of Chicago in 1892 remaining there until 1899, when he became Professor
of Zoology at the University of Austin, Texas. During this period he spent some time
in Europe, where he made many contacts, and he married Miss Dora Bay Emerson of
Rockford, Illinois, who was in the words of Howard Parker "a woman of great personal
charm and delightful presence, who in the years that followed made his household
a hospitable centre for friends and for distinguished visitors from all quarters of the globe".
In 1903 Wheeler became Curator of Invertebrate Zoology in the American Museum of
Natural History in New York City, but returned to teaching at the Harvard University in
1908 and remained associated with that University until his death on 19th of April 1937 at
the age of 72.

42
 Wheeler is most remembered for his book "Ants: their structure, development and
behavior" published in 1910, which opened a new chapter in the history of myrmecology, but
arguably he was the greatest ant taxonomist of the first half of the 20th century. His approach
to taxonomic work was much more "modern" than those of all earlier authors, including
Emery and Forel, and he provided the model on which modern myrmecologists, such as
W. L. Brown and E. O. Wilson, built during the 1950s. He published > 320 works about
ants and described > 600 new ant species. His monograph "Ants of the American Museum
Congo expedition. A contribution to the myrmecology of Africa" (1922) was the first and still
the most comprehensive monograph about Afro-tropical ants, while his paper "The ants of
the Baltic Amber" (1915) became the basis for subsequent ant palaeontological studies.

F. Santschi
(1872-1940)

Felix Santschi was born in 1872 and is considered to be

a native of Lausanne, Switzerland. He studied medicine
and practised as a doctor in Tunisia, mainly in Kairo-
uan, now known as Al Qayrawân, or the "City of 100
Mosques". We know nothing more of his private life.
His main interest seems to have been the study of desert
ants. Santschi appears to have been a good experimen-
talist, pioneering studies on the navigational abilities of
ants. He showed that harvester ants used the sky to
navigate and as long as a small patch of sky was visible,
the ants could return to the nest. On the other hand, the
American taxonomist William Brown, considered his taxonomy as erroneous and outdated
due to his profligate naming of an abundance of new species, subspecies (or "stirpes" in his
terminology) and varieties, based mainly on slight differences in colour and pubescence of
a few individual specimens (Brown 1955). The practical effect of this is illustrated by his
1929 monograph on Cataglyphis (Wehner et al. 1994). Nevertheless, Santschi is still con-
sidered to be an important collector and describer of ants. He died in 1940 aged 68. A short
obituary was published by Donisthorpe (1941).

J. Bondroit
(1882-1952)

Jean Bondroit was an enigmatic character who appeared

on the zoological scene in Belgium after World War I,
when he made some important contributions and then
fell-out with his contemporaries before returning to
relative obscurity. Much of the following is summarized
from the interesting biography of Belgian entomologists
published by Pauly (2001). Apparently Bondroit was in
Paris at the start of World War I, he volunteered for
military service in 1914 and was severely wounded in
October 1916, being evacuated from the front and
put on indefinite leave. Judging from his subsequent

43
behaviour one wonders to what extent his mental balance was also affected. He is said
to have been proud of being an active soldier and despised those who avoided front-line
service.
He probably had artistic leanings before the war because he appears to have been
a biological illustrator by profession being trained in microscopic techniques in the
Laboratoire d'Evolution des êtres organisés, at the Sorbonne, Paris. He seems to have
fallen-out with his employers in Paris and moved to Liege in 1920, where he worked
for the Faculty of Medicine. It seems he was not treated properly by the Professor
and believed he was being taken advantage-of: so, one day he locked the door and
told the Professor exactly what he thought of him and his work, culminating in shouting
that "if he [the Professor] had not been an old man he would have thrown him
down the stairs". He remained in Liege until 1925 eking an existence in various employ-
ments, before being recruited as a "preparator" by Prof. Lameere of the University of
Brussels. Lameere wrote that Bondroit was "a draftsman of talent and an excellent
naturalist".
He often dressed like a workman and took little sustenance. It is said "He had contempt
for the middle-class and even of affectations that facilitate a life in [polite] society" and
while his contemporaries appear to have had some sympathy with him on this matter most
agree that Bondroit had a very difficult personality. On one hand, he was always ready to
dismiss anyone who disagreed with him, having been said to once wave a revolver at the au-
dience in a scientific meeting. On the other hand, he is said to have been affectionate and
friendly to those few people with whom he agreed and his few friends are said to have felt
sorry that he chose such a difficult life for himself. He died in impoverished and lonely cir-
cumstances in 1952 at the age of 70.
As a naturalist Bondroit published and illustrated several works on beetles and is
said to have had a great eye for detail, perhaps attributable to his artistic skills.
Nowadays he is best remembered for his work on ants, his ant collection was "rescued"
after his death and is preserved in the Institut Royal des Sciences Naturelles de Belgique,
Bruxelles. His monograph "Les fourmis de France et de Belgique" (Bondroit 1918),
caused great controversy among contemporary myrmecoiogists because he more or
less disregarded Forel's concept of hybrids (especially among the genus Formica) resur-
recting a number of names considered to be synonyms, using female caste characters,
such as leg pilosity and gastric sculpture. However, general acceptance of this novel
approach was not helped by his freely expressed contempt for the older generation
of taxonomists, he is supposed to have written "I could understand nothing about
ants while I took Forel and Emery seriously ... Once I examined almost all the litera-
ture since Systema Naturae, and I decided to give Emery and Forel a first class
burial, with the cross, the compass and the square, so that everyone is content". Such
"heresies" probably accounted for his academic isolation and dismissal by authors
such as Weber (see Section 1.1) and Bernard (1967). Nevertheless, many of Bon-
droiti ideas on ant species have stood the test of time; we consider that his apprecia-
tion of the genus Myrmica was far superior to that of many of his better known contem-
poraries.

44
 С. Menozzi
(1892-1943)

Carlo Menozzi (also spelt as Minozzi in his early papers)

was born on 7th April 1892 in Spilamberto, Province
Modena, Italy. As a child he was interested in Natural
History, particularly of entomology. His professional
work mostly involved applied agricultural entomology;
after his degree studies he worked for a range of organi­
zations, e.g. the Geological Institute in Padua, the
Laboratory of the Agricultural Entomology in Portici, the
Agricultural Department in Modena and the Phyto-
pathological Laboratory in Chiavari. From 1929 until his
death Menozzi worked as an entomologist in the Na­
tional Consortium for Sugar Production (Grandi 1943a,
b; Anonymous 1943a, b, 1946; Conci and Poggi 1996).
Menozzi's scientific interests were very wide: he published many paper on Dermaptera,
Solifugae and other groups of arthropods. However after he met Carlo Emery his principle
research interest was the taxonomy of ants on which he published ca. 60 papers; these in­
cluded several important contributions to Myrmica taxonomy (including a first key to the
Himalayan Myrmica), Also he carried out valuable work by partly sorting Emery's collec­
tion after his death. Menozzi had a large collection of Dermaptera, Homoptera, Orthoptera,
Diptera and other arthropods as well as his important ant collection. After his premature
death in Spilamberto on 4th March 1943 at the age of 51, his collections and his personal
library were housed in the Istituto di Entomologica, Bologna.

В. Finzi
(1897-1941)

Bruno Finzi was born on 25 th August 1897 in Trieste,

Italy. The following data are extracted from an obituary
by Menozzi (1941) and a biography of Italian entomo­
logists (Conci and Poggi 1996). Finzi was a school­
teacher having graduated from the Istituto Magistrale
in 1920. From 1929 onwards he was Director of the
Scuola Commune of Trieste besides acting as an assis­
tant to Curator of the City Museum of Natural History.
He was married but we know little else of his personal
life. He joined Mussolini's Italian Fascist Party and in
1936 he became Comandante di Coorte della 175a Le­
gione. He died on 30th July 1941 in Trieste at the early
age of 44 years. It is not clear whether this was from
natural causes or a result of wartime actions, most probably it was the former because
Menozzi made no mention of how he died.
Besides his pedagogical and military interests, Finzi was a talented entomologist having
joined the Italian Entomological Society in 1921. Like many of his contemporaries he had
a wide range of interests in entomology having, for example, made studies of Odonata and
Libellulidae. Nevertheless, his primary interest was ants. He amassed a large and important
ant collection, which after World War II his widow sold to the United States and it is pre­
served in the Museum of Comparative Zoology, Harvard.

45
 К. V. Arnoldi
(1901-1982)

Konstantin Vladimirovich Arnoldi was born on 5 th Jan­

uary 1901 in Moscow and died there 12th December
1982, aged 81. He came from a family of scientists, his
father was V. M. Arnoldi, the well-known algologist
and hydrobiologist, a professor of the Kharkov and
later of Moscow Universities; his grandfather (from
the mother's side) was the botanist, Professor I. N.
Gorozhankin. Arnoldi attended a school in Kharkov
(Ukraine), graduating in 1921, before entering the
Natural History faculty of the Moscow University, from
which he graduated in 1926. After graduation Arnoldi
worked in the Moscow University. In 1934 he moved to
Leningrad (now St.-Petersburg), to the Zoological
Institute of the Academy of Sciences of the USSR. From 1942 until 1969 when he retired,
he was employed as a senior scientist at the Institute of Evolutionary Morphology and
Ecology of the Academy of Sciences of the USSR in Moscow (Anonymous 1982; Gilarov
et al. 2003; Dlussky and Zakharov 2001).
At first it looked as if Arnoldi would follow his father, his first scientific paper was de­
voted to hydrobiology, but later he started to work as an entomologist with a particular in­
terest in myrmecology. Besides his work on ant taxonomy he was a recognized expert on
many groups of beetles and bugs, and soil arthropods generally. Among his papers there are
several important publications concerning theoretical problems of systematics, zoogeogra­
phy and biocoenology that unfortunately are not known to most modern Western biolo­
gists. As a myrmecologist, he not only contributed to basic taxonomy, describing several
tens of species and two ant genera, but also made important revisions of many genera, in­
cluding Myrmica (1934,1970,1976). Until the 1980's he was the author or co-author of all
Keys for ants of the USSR. He was a keen collector, taking part in many scientific expedi­
tions throughout the former USSR. He amassed a very rich and important collection of ants
that is mostly preserved in the Zoological Museum of the Moscow State University, and in
part, in the Zoological Institute of the Russian Academy of Sciences, St. Petersburg.

46
 N. A. Weber
(born 1908)

Neal Albert Weber was born in 1908, we believe him to

be dead but can not find a date. He completed his Mas-
ters degree at the Harvard University in 1933 and was
awarded his PhD in 1935 for his thesis entitled "A revi-
sion of the ants of the genus Myrmica Latreille" made
under the supervision of William Morton Wheeler. He
published descriptions of five new forms of North Amer-
ican Myrmica in 1939. While at Harvard he participated
in scientific expeditions to the West Indies (1933-
1936), where he probably developed his interest in Leaf-
cutting ants and Army / Driver ants for which he is
better known. He took up a post in the University of North Dakota and made several field
trips to South America and Africa. By 1947 he had obtained a post at the Swarthmore Col-
lege, Pennsylvania where he remained until 1974. This move seems to have stimulated him
to publish his revision of Myrmica (in three parts, 1947, 1948 and 1950) in which he de-
scribed two good Old World Myrmica species. During his time at the Swarthmore College
he participated in numerous American Museum of Natural History expeditions: e.g. to Cen-
tral Africa in 1948, to the Middle-East in 1950 and 1952, and to Tropical America in 1954.
Weber's interest went wider than ants: he was a visiting professor at the University of Bagh-
dad, Iraq (1950-1952), U.S. Dept. of State, Scientific Attaché, Buenos Aires (1960-1962),
a member of the National Academy of Sciences Committee on Polar Research (1958-1960)
and deeply involved with the Entomological Society of America and local ecological groups.

J. Sadil
(1919-1971)

Josef Sadil was born 19th March 1919 in Prague and

died there on 19th January 1971 at the early age of 51.
He was an astronomer by profession and an enthusias-
tic amateur entomologist with a particular interest in
ants. He started studying astronomy at the Charles Uni-
versity in Prague but his studies were interrupted by
World War II, were not resumed until after 1945. Sadil
was well known in Czechoslovakia as a populariser of
astronomy and he published nine books on the Earth,
planets and aspects of astronomy (in the Czech lan-
guage). As a myrmecologist he was best known in
Czechoslovakia for his book published in 1955, "Our ants" (written in Czech), which more
or less marked the end of his work on ants. In the context of this monograph he is remem-
bered for his detailed revision of Central European Myrmica species, in which he provided
comprehensive tables of morphometries (Sadil 1952). Sadil left a small collection of ants
(ca. three thousand pins, mainly from the Czech Republic), which is preserved in the
general entomological collections of the National Museum, Prague.

47
 A. N. Kupyanskaya
(born 1934)

Alina Nilovna Kupyanskaya was born into a teacher's

family on 16th April 1934 in Kalach, the Voronezh
Province of Russia. She graduated from the Natural His-
tory faculty of the Voronezh State University in 1959
and worked as a school teacher in Vladivostok until
1964. From 1965 until 1968 she was a post-graduate
student in the Soil and Biology Institute of the Far
Eastern Scientific Centre of the Academy of Sciences of
the USSR in Vladivostok, and in 1970 she successfully
defended her PhD thesis, entitled "Pest insects of the
green plantations of towns of Primorsky Krai". Kupyan-
skaya still works at the Soil and Biology Institute in
Vladivostok. Her main scientific interest is now the
ecology and taxonomy of ants, mainly of the Far East of Russia, a huge territory comprising
several million square kilometres of forest and mountains. Kupyanskaya has published
about 70 scientific works, including two monographs. Among these she described six Myr-
mica species; three of them were given replacement names by Bolton (1995) because,
unfortunately, she used homonymous names (M. aspersa, M. carinata and M. bicolor).

G. M. Dlussky
(born 1937)

Gennady Mikhailovich Dlussky was born 4 th October

1937, in Moscow, the son of a railway-engineer. He is
married to Irina and they have two married sons and
three grandchildren. From an early age Dlussky had a
strong interest in natural history, especially entomology
in general and ants in particular. After finishing school
in 1954 he became a student in the Faculty of Biology of
the Moscow State University, graduating in 1959. In
1957 he first met Prof. K. V. Arnoldi, who inspired his
future work on ants; after graduation he started work
as Arnoldi's assistant in the Institute of Evolutionary
Morphology and Ecology of the Academy of Sciences of
USSR in Moscow. At that time, his main work was on the biology and taxonomy of the
genus Formica, especially the practical use of red wood ants in pest control. He successfully
defended his PhD in 1967, entitled: "The ants of the genus Formica of the USSR" and in
the same year published the monograph "The ants of the genus Formica", that still is a stan-
dard reference on this genus.
In 1972 Dlussky transferred back to the Moscow State University, Cathedra of Evolu-
tionary theory, where he still works as a Professor, being awarded a Professor's degree by
the Moscow State University in 1988. He successfully defended a Doctor of Science thesis
in 1981 entitled "The ants of deserts"; this excellent monograph has no analogues in the
world literature, being devoted to the biology and synecology of desert ants and addressing

48
the problems of the origin, function and evolution of desert ant communities. Dlussky has
published > 150 scientific papers and several faunistic monographs on ants, including "Ants
of Turkmenistan", "Ants of Bulgaria", "Ants of Fiji and Samoa". His scientific interests re-
main very wide: taxonomy and ecology of ants, problems of the theory of evolution, history
of biology, general ecology, etc.; however over the last 25 years the palaeontology of ants
has become his main interest to the extent that he is now probably the leading expert on the
fossil ants of the Old World.

B. Seifert
(born 1955)

Bernhard Seifert, the son of a surgeon, was born on 28 th

September 1955 in Zwickau, East Germany (then part
of the DDR). He is married with one son. From 1974-
1979 he studied biology at the University of Leipzig spe-
cializing in animal physiology. For a short while he
researched in developmental biology at the Central In-
stitute of Genetics and Crop Research at the Academy
of Sciences in Gatersleben before being appointed to the
Staatliches Museum für Naturkunde Görlitz.
Since then his main professional interest has been
the taxonomy and ecology of Palaearctic ants. He
published his first paper on ants in 1982 and was
awarded a doctorate in 1985 for a thesis entitled "Comparative studies on habitat selection
of ants in central and southern part of the GDR" (in German). This work on the ecology
and habitat use by common ants in the southern DDR showed his considerable originality
as an ecologist (Seifert 1986). In 1990 Seifert was made a Senior Curator in the Görlitz
museum, with special responsibility for pterygote insects. Since then he has concentrated
on taxonomy publishing about 100 papers, including a revision of Myrmica of Europe and
the Caucasus (Seifert 1988) and several books, the most recent being an account (in Ger-
man) of the ants of Middle and North Europe (Seifert 2007). Seifert has participated in
several expeditions to the Caucasus, Middle Asia and elsewhere in Europe. As a result, he
has amassed an important ant collection that is housed in the Görlitz Museum. His work
on Myrmica has mainly concerned infra-specific variation for which he uses very accurately
measured morphometries and various mathematical discriminant techniques.

49
Table 1.1. The number of good Myrmica species (both extant and extinct) of the Old World,
described by different authors (author listed according number of species described; the list
includes only those authors that have described at least one species as a senior author; for this
purpose species having replacement names for preoccupied names are credited to the authors of
the original description).

2Author names: Living authors Number of described species

Radchenko X 59 (55 together with Elmes and other co-authors)*
Elmes X 52 (51 together with Radchenko and other co-authors)*
Forel 13
Ruzsky 12
Arnoldi 7
Nylander 6
Kupyanskaya X 5
Mayr 4
Wheeler 4
Karawajew 4
Bondroit 4
Finzi 4
Emery 3
Santschi 3
Menozzi 3
Weber 2
Dlussky X 3 (2 together with Radchenko and Elmes)
Seifert X 2
Linnaeus
Meinert
Sadil
Kutter
Bernard
Collingwood X

Bolton X

Cagniant X

Espadaler X

Wei X

* The following scientists are junior co-authors of good Myrmica species described mostly in
collaboration with Radchenko or Radchenko and Elmes: S. Zhou, China (6 species), N. Aktac, Turkey
(4 species), F. Rigato, Italy (3 species), B.-J. Kim, South Korea (1 species), A. Alicata, Italy (1 species),
M. Liu, China (1 species), B.-T. Viet, Vietnam (1 species), and R. Savolainen, Finland (1 species).

50
CHAPTER 2. TAXONOMIC STRUCTURE OF THE GENUS MYRMICA

2.1. Taxonomic position and diagnosis of the genus Myrmica

Myrmica Latreille is a type-genus of the subfamily Myrmicinae of ants (Formicidae) and,

accordingly, of the tribe Myrmicini. Below we summarize the taxonomic history of this tribe
(for full details see Bolton 2003, Bolton et al. 2006 and literature cited therein).
By the time Emery produced his catalogue (Emery 1921) tribe Myrmicini comprised
three genera: Myrmica with its subgenus Manica Jurine, Pogonomyrmex Mayr with its sub-
genus Hylomyrma Forel, and Cratomyrmex Emery. Subsequently, the name Cratomyrmex
was considered to be a junior synonym of Messor Forel (tribe Pheidolini), and Manica and
Hylomyrma were raised to genus level. Although since Emery several genera (e.g.
Ephebomyrmex Wheeler, Sifolinia Emery and Huberia Forel, see Dlussky and Fedoseeva
1988, Hölldobler and Wilson 1990) have been included in the Myrmicini, by the time the
most recent additions were made (Bolton 1994, 1995, 2003) the composition of the tribe
had hardly changed. Bolton transferred to Myrmicini from Tetramoriini the genus
Eutetramorium Emery, containing two species from Madagascar (Bolton 1976), and added
Secostruma Bolton that comprises a single species from Borneo (Bolton 2003). The com-
position of the tribe was confirmed by Bolton et al. (2006), who also included to Myrmicini
Nothomyrmica Wheeler, an extinct genus from the Baltic Amber. At more or less the same
time we synonymized Nothomyrmica with Myrmica, transferring also some of its species to
Temnothorax Mayr (Radchenko et al. 2007). Most recently, two new extinct primitive
genera Protomyrmica and Plesiomyrmex, described from European Late Eocene amber,
were added to the tribe (Dlussky and Radchenko 2009).
Hence, currently the tribe Myrmicini contains nine genera: thè extant Myrmica,
Manica, Pogonomyrmex, Hylomyrma, Eutetramorium, Huberia and Secostruma and
the extinct Protomyrmica and Plesiomyrmex. With the advent of modem molecular-
phylogenetic analysis it is unlikely that the tribe will remain intact in its present state
(Dlussky and Radchenko 2009). Furthermore, Bolton noted that "Secostruma is only very
dubiously associated with Myrmicini core-genera" (pers. comm. 2008), and very.recently
Jansen and Savolainen (2010), using molecular techniques, concluded that Eutetra-
morium and Huberia do not form a monophyletic group with the other genera in
Myrmicini.

Subfamily MYRMICINAE Lepeletier, 1835

Myrmicites Lepeletier, 1835: 169 (as a group name). Type genus -Myrmica Latreille, 1804.
Myrmicidae: Mayr, 1855: 290 (as subfamily Formicidae) (emendation of spelling).
Myrmicinae: Forel, 1893: 163 (as subfamily Formicidae) (emendation of spelling).

51
 Tribe MYRMICINI Lepeletier, 1835
Myrmicites Lepeletier, 1835: 169 (as a group name). Type genus -Myrmica Latreille, 1804.
Myrmicini: Forel, 1891: 143 (as tribe Myrmicinae).

Genus Myrmica Latreille, 1804

Myrmica Latreille, 1804: 179. Type species -Formica rubra Linnaeus, 1758: 580, by subsequent
designation: Latreille 1810: 437.
Sifolinia Emery, 1907: 49. Type species - Sifolinia laurae Emery, 1907: 49, by monotypy. Synonymy
by Brown 1973: 184 (provisional); Bolton 1988: 3; Czechowski et al. 2002: 16; Radchenko and
Elmes 2003a: 224; Bolton 2003: 222 (see also Notes below).
Sommimyrma Menozzi, 1925: 25. Type species - Sommimyrma symbiotica Menozzi, 1925: 25, by
monotypy. Synonymy by Brown 1973: 184 (provisionai); Bolton 1988: 3; Radchenko and Elmes
2003a: 224; Bolton 2003: 222.
Symbiomyrma Arnoldi, 1930: 267. Type species - Symbiomyrma karavajevi Arnoldi, 1930: 269, by
monotypy. Synonymy by Samsinak 1964:156 (as synonym of Sifolinia); Bolton 1988: 3 (as synonym
of Myrmica), 1994: 106; Radchenko and Elmes 2003a: 231; Bolton 2003: 222 (see also Notes below).
Paramyrmica Cole, 1957: 37. Type species - Paramyrmica colax Cole, 1957: 37, by monotypy.
Synonymy by Brown 1973: 183 (provisionail); Bolton 1988: 3; Radchenko and Elmes 2003a: 219;
Bolton 2003: 222.
Dodecamyrmica Arnoldi, 1968b: 1803 (as subgenus of Myrmica). Type species -Myrmica (Dode-
camyrmica) arnoldii Dlussky, 1963: 191, by original designation. Synonymy by Brown 1973: 180
(provisional); Francoeur, 1981: 759; Bolton, 1988: 3; Radchenko and Elmes 2003a: 222; Bolton
2003: 222.

Workers (Fig. 2)

Diagnosis. Myrmica workers are monomorphic and intermediate in size compared to

ants generally. The length of body of most species is 4.5-6.0 mm, though some Oriental
species can be much larger, up to 8.5 mm long. Typically the overall colour is reddish-brown,
but species can be found in all shades varying from light brownish yellow, through chest-
nut brown, dusky reddish-brown to almost black. As in all ants, young, callow workers have
almost no body pigment and take some months to acquire their final adult colour. When ma-
ture, the head often appears distinctly darker than the alitrunk and the appendages appear
somewhat paler, so that the workers of some species appear distinctly bicoloured. Colour
and the degree of bicolour can vary between local populations of the same species.
Seen from above, the shape of the head is sub-oval, the sides usually being slightly con-
vex and the occipital margin straight or somewhat convex, as a rule with well-marked
occipital corners. The anterior clypeal margin is convex and, in many species, it has a dis-
tinct notch medially. The eyes are well developed being situated on the sides of the head,
approximately half way along its length, and they always project beyond the lateral mar-
gins. Ocelli are absent in normal workers, but intercastes can have reduced versions. The
mandibles are elongate-triangular in shape; their masticatory margins rarely have more than
10 teeth, usually comprising two large apical teeth and a series (5-7) of smaller ones. The
maxillary palps are 6-segmented and labial palps 4-segmented. The antennal sockets are
distinct, being situated somewhat anterior to the eyes and approximately mid way between

52
the sides and the longitudinal axis of the head, though the precise location varies between
species. Seen from above, the frontal carinae can appear almost straight, but always to some
extent curve outwards anteriorly and posteriorly so that the position of the minimum
distance between the frontal lobes is easily recognized. Anteriorly, the carinae extend out-
wards to form frontal lobes, flattened plates that partly project over the antennal sockets.
The size of the frontal lobes relative to the minimum distance between the frontal carinae
can vary greatly between species. Antennal scrobes are not present. The antennae are
12-segmented, comprising a scape that usually surpasses the occipital margin, and
11 funiculus segments that are enlarged apically to form a moderately defined
3-4-segmented club. The scape is always bent at its base; the degree of bending can vary
from a smooth natural curve to a distinct, almost right-angled bend. The bend of the scape
can be twisted to some extent, which has the effect of rotating the scape around the plane
of its longitudinal axis. In many species, the effect of bending and twisting has led to the
development of various lobes, dents or carinae on the bend of the scape that have become
important in the taxonomy of the genus.
The alitrunk is usually distinctly narrower than the head seen from above. The widest
part being the pronotum, near its broadly rounded anterior margin, generally, it is
obviously wider than the mesonotum and propodeum. The promesonotum (seen in profile)
appears somewhat convex, while the promesonotal suture (seen from above) is indistinct
or absent in most species, but can be distinctly obvious in some. On the other hand, the
metanotal groove is always present and often appears deep and abruptly marked although
in some species it appears very shallow. The propodeum possesses a pair of spines that can
be useful diagnostically: according to the species these vary in length and can range in shape
from stout broad-based thorns to fine needle-like structures.
The waist is 2-segmented and variation in its structure is very useful in the taxonomy of
the genus. The anterior segment is the petiole that extends anteriorly to form a well-
developed peduncle that articulates with the alitrunk, and posteriorly (when seen in profile)
the petiole is enlarged dorsally to form a distinct node that, depending on species, can vary
in shape ranging from rounded to triangular or sub-rectangular. The posterior segment, the
postpetiole, is distinctly more rounded, from above its appearance ranges from sub-
globular to fig-shaped. The ventral surface of the petiole has a small rounded tooth that is
usually absent from the postpetiole, although in some socially-parasitic species both
petiole and postpetiole have well-developed ventral lobes.
The structure of the legs is not much used in the taxonomy of Myrmica with the excep-
tion of the tibial spur. The middle and hind tibiae usually have well-developed pectinate
spurs that are usually longer than maximum width of the tibia, but in some species (espe-
cially socially-parasitic ones) the development of the spurs is reduced, spurs might be sim-
ple with no pectination or even absent altogether. The gaster has no obvious external
morphology apart from punctures and striations on the surface of the first tergite in a few
unusual species; consequently it is not much used in Myrmica taxonomy.
In contrast, the sculpture and pilosity of the head, alitrunk and waist is quite variable
between species and differences are often considered to be diagnostic for species identifi-
cation. Almost all species have quite coarse sculpture that ranges from fine to coarse
reticulation and from fine striations to coarse rugosity, usually rugae are longitudinally
orientated but in some species (mainly Oriental ones) they can be transverse. In addition,
the surface between the rugae and reticulations can be coarsely or finely punctated,
making the surface appear less shiny or even matt. Only a few species have sculpture that

53
is so reduced that the body appears wholly or at least partly smooth and shiny. Often only
decumbent pubescence is present on the appendages but the number, location and length
of erect hairs on the head and body can be quite variable and diagnostic between species,
the hairs on the body are usually rather long and more abundant than those on the head.
A few species, mainly parasitic ones, can be extremely hairy.

Figure 2. Morphology of Myrmica workers: A, head, frontal view: as - antennal socket, ci - clypeus,
ey - eye, fc - frontal carina, fl - frontal lobe, fr - frons, ft - frontal triangle, gn - gena, md - mandible,
oc - occiput, om - occipital margin, tl - temple; B-E, differently shaped scapes, lateral view:
B, gradually curved; C, angulate; D, with horizontal lobe; E, with vertical lobe; F, alitrunk
(= mesosoma) and waist (petiole + postpetiole), lateral view: al - alitrunk, mg - metapleural gland,
mn - mesonotum, mtg - metanotal groove, pd - petiolar peduncle, pl - propodeal lobe; pn - pronotum,
pnd - petiolar node, ppt - postpetiole, pr - propodeum, psp - propodeal spine, ps - propodeal spiracle,
pt - petiole; G, alitrunk and waist, dorsal view: mn - mesonotum, ns - neck shield, pn - pronotum,
ppt - postpetiole, pr - propodeum, psp - propodeal spine, pt - petiole; F, hind leg: bts - basitarsus;
ex - coxa, fm - femur, tb - tibia, tr - trochanter, trs - tarsus, ts - tibial spur.

54
 Queens (Fig. 3)

In almost all cases queens have similar external morphology to their workers, apart from
the "fully developed" alitrunk structure and the presence of ocelli. With a few very rare
exceptions they are distinctly larger (ca. 25%) than their workers (body length ranging
5.5-8.0 mm), generally they have about double the mass. This causes natural allometry in
the relative size of various structures, for example the width of the frons relative to the head
is usually slightly greater in queens compared to the conspecific workers. Queens of the
very large species, found in South-East Asia, are still unknown, but we suspect some could
have head-widths in excess of 2 mm. On the other hand, queens of many of the workerless

Figure 3. Morphology of Myrmica queens: A, head, frontal view: ocl - ocelli (other parts of head
- as in workers); B, scape, lateral view: C, alitrunk (= mesosoma) and waist (petiole + postpetiole),
lateral view: mpl - mesopleuron, pn - pronotum, pr - propodeum, ppt - postpetiole, psp - propodeal
spine, pt - petiole, sct - scutum, scti - scutellum; D, alitrunk and waist, dorsal view: pn - pronotum,
pr - propodeum, ppt - postpetiole, psp - propodeal spine, pt - petiole, sct - scutum,sctl- scutellum.

55
socially-parasitic species are small (body length 4.0-5.0 mm), much smaller than their host-
workers. A few species produce two sizes of queen, large normal queens or macrogynes, and
small queens or microgynes, which are not much larger than their workers (see Chapter
1.3). Wings are present only in young unfertilized queens (gynes) (for details of the wing
venation see males).

Males (Fig. 4)

Myrmica males are normally a brownish black colour, varying from dark brown to wholly
black. The size of individual males from the same colony can vary greatly; on average their
body length is slightly smaller than that of conspecific queens although the two sexes are
often about the same size, it is very rare for males to be larger than queens. However, com-
pared to queens, males appear distinctly more "fly-like" being more slender with longer legs
and a distinctly much smaller head. The head is relatively short compared with its width,
generally being only very slightly longer and sometimes shorter than the head-width so that
it appears spherical. The eyes and ocelli are often much larger than those of conspecific
queens and appear very large in proportion to the head. The mandibles are less broadened
at their apex compared with workers, nevertheless males have a distinct masticatory mar-
gin with 4-6 (very rarely with 8-10) teeth. The antennae are normally 13-segmented, but
many socially-parasitic species and a very few free-living species have only 12 segments.
The length of the male scape is an important taxonomic diagnostic feature in Myrmica;
according to species, it can vary in length from < 30% of head-width to longer than the
head-width. Also, the scape shape can range from almost straight, to a smooth curve or being
sharply bent at the base, however it never has any lobes or projections at the bend, as do the
workers and queens. The funiculus has a moderately defined 4-5-segmented apical club and
the joints distinctly increase in length from the second to the apical. The scutum has
Mayrian furrows and the propodeum has no spines, only small rounded tubercles or at most,
short denticles. The forewing has closed cell mcu, an open cell 3r and vein 2+3RS reduced
proximally so that cells l+2r and rm are partly separated (Fig. 3, E). However, in many
socially-parasitic species vein 2 +3RS is completely reduced and cells 1 +2r and rm are fused;
furthermore, cross-vein m-cu sometimes is reduced, so that cell mcu is open (Fig. 4, F, G;
for some more details see also Bolton 1988). The tibial spurs are as in female castes.
Distribution. Myrmica is predominately a Holarctic genus, but many species are also
found in the mountains of the Oriental Region (Himalaya, Taiwan, southern China, Viet-
nam, Burma, Thailand and Indonesia) and in the Neotropics (Mexico) (see Map 1).
Notes. Several socially-parasitic genera, closely related to the genus Myrmica, were de-
scribed during last 100 years (see above). Despite several attempts to clarify the relation-
ship between genus Myrmica and its parasitic "satellite genera" (e.g. Kutter 1973), nobody
could discriminate them consistently and satisfactorily. Many species had characters inter-
mediate between those of, for example, Sifolinia and Myrmica, and it became somewhat ar-
bitrary as to which genus authors ascribed new species (e.g. Elmes 1976). Eventually,
Bolton (1988) showed that there are no diagnostic characters for any of the "satellite"
genera that were not also present in Myrmica, and he formally synonymized Sifolinia,
Sommimyrma, Symbiomyrma and Paramyrmica with Myrmica. Subsequently, Seifert
(1994,1996) did not entirely agree with Bolton's treatment, and revived from synonymy the
name Symbiomyrma, including in this genus S. karavajevi (with its synonyms S. pechi,
S. winterae and M. faniensis), S. lemasnei and S. kabylica. More recently, Radchenko

56
and Elmes (2003a) provided additional arguments for synonymisation of Sifolinia with
Myrmica, nevertheless Seifert (2007) again proposed to consider Symbiomyrma as a sub-
genus of Myrmica. We do not want to prolong a non-fruitful discussion so we leave matters
as they are. Nevertheless, if Symbiomyrma is considered a subgenus, we could with as much
justification, establish several subgenera from among the oriental Myrmica species, although
some would comprise only one or two species. Thus the question of formal subgenera seems
to us to reflect more an authors' personal approach rather than being important taxonomi-
cally. In this monograph we will not separate the genus Myrmica into subgenera rather we
prefer to use the less formal and more flexible concept of species groups and complexes.

Figure 4. Morphology of Myrmica males: A, head, frontal view: ocl - ocelli (other parts of head
- as in workers); B, antenna: ac - apical club; fn - funiculus; sc - scape; C, alitrunk (= mesosoma)
and waist (petiole + postpetiole), lateral view: mf - Mayrian furrow, mpl - mesopleuron,
pn - pronotum, pr - propodeum, ppt - postpetiole, pt - petiole, sct - scutum, seti - scutellum;
D, alitrunk and waist, dorsal view: mf - Mayrian furrows, pn - pronotum, pr - propodeum,
ppt - postpetiole, pt - petiole, sct - scutum, scti - scutellum; E-G, forewings; E, M. rubra, with "typical"
wing venation for the genus; F, M. laurae, with completely reduced vein 2+3Rs and fused cells l+2r
and rm; G, M. karavajevi, completely reduced vein 2+3Rs and fused cells l+2r and rm, and reduced
cross-vein m-cu and open cell mcu.

57
2.2. Species group concept in the genus Myrmica

When considering the range of variation between species within a genus it is usually ap-
parent that some species are very similar to each other and are quite dissimilar to other
clusters of species. It is normal practice in taxonomic studies to reflect this either by formally
separating the genus into subgenera, or by using the less formal concept of species groups
(note that the International Code of the Zoological Nomenclature does not regulate the
concept of species groups or species complexes). The problem with a subgeneric division of
the genus Myrmica is that even in the case of the niae-group, which appears to us to have
the clearest claim as a subgenus, we have found species intermediate to other clear groups;
the number of clear subgenera would be so small that the vast majority of species would
remain unassigned. The greater flexibility of the species groups concept makes it possible
to assign a majority of Myrmica species to a group, but even so, a number of taxa are so dis-
similar to all others that they can not be satisfactorily assigned to any group, or can be
considered as a mono-specific group.
Species groups. Ideally, as in any supraspecific taxon, the species groups should include
phylogenetically related species. We have assigned species to groups using characters of all
three castes. Species are distinguished from related ones by at least one morphological
autapomorphy, but in contrast species groups are defined by features the species hold in

58
 common and they can only be distinguished from other groups by a whole complex of fea-
tures, some of which may widely overlap. In some cases, especially when only a single caste
is known (usually workers) homoplasies (independently evolved similarities) will place un-
related species in the same group. Recently, the molecular studies of Jansen et al. (2009,
2010) showed that those of our proposed species groups that they included in their study, ap-
pear to be monophyletic; however, the same studies also indicate that generally the species
complexes suggested by us appear to be paraphyletic, i.e. artificial complexes based on mor-
phological similarities that do not necessarily reflect close relatedness. Even so, the match
between Jansen et al.'s molecular phylogeny and our species groups gives us confidence that
when the "gaps" in their phylogeny are "filled in" most of our species groups will be vali-
dated. In the mean time, the species group names are useful "shorthand" when discussing the
distribution and evolution of the genus.
Bondroit (1920a) was the first to propose a more or less harmonious concept of species
groups among the European Myrmica species: he recognized three groups, ruginodis,
scabrinodis and lobicornis, that include 24 species and infraspecific forms. Considering the
criticisms that Bondroit faced in his lifetime (see Chapter 1.6) it would have gratified him
that we retain most of his ideas on these groups. The principal exceptions are his placing
of M. schencki in the scabrinodis-group and the modern treatment of M. sabuleti as repre-
senting a separate species complex within the group.
Bernard (1967) was less successful when he established only two species groups
(scabrinodis and rubra) based on the length of the antennal scape of males. He noted
"... les Myrmica d'Europe se répartissent très nettement en 2 groupes, bien séparés par les
antennes des mâles et les adaptations écologiques des ouvrières" (loc. cit., p. 109). In the
scabrinodis-group he placed species with males having a short scape ("...à scape très court,
égalant 2 à 4 articles du funicule"), and species with males having a long scape ("...à scape
long et mince, noir ou brun, égalant 5 à 7 articles du funicule") he placed in the rafcra-group
(loc. cit., p. 110). While we generally agree with Bernard's treatment of the scabrinodis
group (only excluding from it M. schencki) his "rubra-group" now seems artificial,
containing not only M. rubra (laevinodis) and M. ruginodis, but also species that we
now consider represent several other groups (M. sulcinodis, M. vandeli, M. lobicornis,
M. myrmecophila and M. lemasnei). Conversely, Elmes and Thomas (1987) recognized four
species groups among the common European species, based solely on the shape of the
worker scape (the lobicornis-, scabrinodis-, rugulosa- and rubra-groups). While this
treatment was moderately useful in helping non-specialists identify the species, it produced
quite artificial heterogenous groups with only the rubra-group resembling our most recent
groupings (below).
Our current concept of species groups in Myrmica derives from the treatment proposed
.by Radchenko (1994b) who combined both male and female characters to define his groups.
Ignoring the socially parasitic species, Radchenko recognized eight species groups among
the Palaearctic Myrmica: the rubra-, rugosa-, scabrinodis-, lobicornis-, schencki-, luteola,
arnoldii- and niae-groups. This was the starting point for our work on Myrmica taxonomy
(Radchenko and Elmes 1998, 1999a, 2001b, 2003a, 2004, 2009a; Radchenko, Elmes and
Viet 2006, Radchenko et al. 2007). In this monograph we recognize 17 groups of Old World
Myrmica (number of species in parenthesis); in alphabetical order they are: the arnoldii
(1), dshungarica (4), inezae (3), karavaievi (3), kurokii (2), laurae (1), lobicornis (22),
luteola (2), myrmicoxena (1), pachei (14), ritae (21), rubra (4), rugosa (9), scabrinodis
(26), schencki (11), smythiesii (5) and tibetana (3) groups.

59
 Below, we list the species groups alphabetically and amplify the diagnostic characteris-
tics of each group, together with any special notes or observations, and give a list of species
assigned to the groups. In some groups it is possible to recognise several species complexes.
Generally, we do not recognise species groups based on a single species, except for the
arnoldii-, laurae- and myrmicoxena-gvoups; the first contains a most unusual free-living
species that has morphological characters often associated with social parasites, and the
latter pair contains poorly studied social parasites (see also Chapter 3.3). This approach
leaves ten free-living species (listed after the species groups data) that can not be placed in
any species group unambiguously.

arnoldii-group
Diagnosis. Workers (Fig. 28) : frontal carinae merging with the rugae that extend to the
occipital margin, they do not curve outwards and do not merge with rugae that surround
antennal sockets; frons wide, but frontal lobes extended. Scape quite strongly curved at the
base but not angled and with no trace of lobe or carina. Petiole and postpetiole with at least
small ventral lobes. Spurs on the middle and hind tibia reduced to various extents.
Males (Fig. 30): antennae 12-segmented, scape short, SIj < 0.40.
This monospecific group was established to contain a most unusual free-living species
from South Siberia and Mongolia that has in all castes morphological features often asso-
ciated with social parasites. At the moment it is unique worldwide, but possibly similar
species could be found in Tibet and China when these regions are more fully studied (see
also Chapters 3.3 and 4.3.2.2): arnoldii.

dshungarica-group

Diagnosis. Workers (e.g. Fig. 72): frontal carinae merging with the rugae that extend
to the occipital margin, they do not curve outwards and do not merge with rugae that
surround antennal sockets; frontal lobes slightly curved, frons wide, and frontal lobes not
extended. Scape very smoothly curved at the base, not angled and with no trace of lobe or
carina. Anterior clypeal margin convex, often prominent, with no medial notch.
Males (e.g. Fig. 74): scape long, SI2 > 1.00.
The species of this group are superficially similar to those of the rubra-group, but differ
from the latter by the shape of frontal carinae. The group includes four species, all of
which are restricted to the Middle Asian mountains and NE Afghanistan: dshungarica,
ferganensis, juglandeti, kryzhanovskii.

inezae-group

Diagnosis. Workers (e.g. Fig. 110): scape long, subequal to head length, smoothly
curved at the base, not angled and with no trace of lobe or carina; frontal lobes slightly
curved, frons wide and frontal lobes not extended. Anterior clypeal margin is distinctly
prominent with no medial notch. Petiole has a very long and thin peduncle and a quite short
and high node; postpetiole is subglobular. Propodeal spines very long; propodeal lobes
rounded, not pointed apically.
Males are unknown.

60
 The species of this group share several features with ritae-group species (e.g. long scape,
petiole and propodeal spines, and a slender body), but well differ from them by their strongly
prominent and not-notched anterior clypeal margin and not-pointed apically (more rounded)
propodeal lobes. We place three species from the Himalaya and south-western China in this
group: inezae, mixta, rigatoi.

karavajevi-group

Diagnosis. Workerless social parasite.

Queens (e.g. Fig. 126): frontal carinae merging with the rugae that extend to the
occipital margin, they do not curve outwards and do not merge with rugae that surround
antennal sockets; frons wide, frontal lobes not extended. Scape gradually but quite strongly
curved at the base, not angled and with no trace of lobe or carina. Petiole and postpetiole
have distinct ventral lobes. Spurs on the middle and hind tibia reduced to various extents.
Males (e.g. Fig. 127): antennae 12-segmented, scape relatively long, SIj > 0.70.
We place three species distributed in Europe and Algeria in this group, which equates
to the subgenus Symbiomyrma (sensu Seifert 2007): karavajevi, kabylica, lemasnei.

kurokii-group

Diagnosis. Workers (e.g. Fig. 145): overall appearance is large and robust. The frontal
carinae merge with the rugae that extend to the occipital margin, they do not curve outwards
and do not merge with rugae that surround antennal sockets. Scape gradually but quite
strongly curved at the base, not angled and with no trace of lobe or carina. Anterior clypeal
margin rounded, without a medial notch. Petiole has a short peduncle and massive, quite
high node. Head dorsum has a very dense but not coarse rugosity, surface between rugae
densely punctated.
Males (Fig. 147): scape long, SI1 > 0.90.
We placed to this group two species, distributed from Himalaya and Tibet to Japan.
Quite probably it will be found that this group comprises several more species when the
fauna of Tibet and China is better known: kurokii, kozlovi.

laurae-group

Diagnosis. Workerless social parasite.

Queens (Fig. 148): frontal carinae merging with the rugae that extend to the occipi-
tal margin, they do not curve outwards and do not merge with rugae that surround
antennal sockets; frons wide, frontal lobes not extended. Scape quite strongly but
gradually curved at the base with a horizontal lobe (the combination of a gradually
curved scape with a lobe is unique among all known Old World Myrmica species).
Petiole and postpetiole with ventral lobes. Spurs on the middle and hind tibia reduced
to a variable extent. Very hairy species; eyes with conspicuous hairs, length of the longest
hairs > 0.035 mm.
Males (Fig. 149): antennae 12-segmented, scape long, Sl1 > 0.75.
This group contains a single species found in the Italian peninsula: laurae.

61
 lobicornis-group

Diagnosis. Workers (e.g. Figs 5, 22, 152, 273, 315): scape strongly curved at the base,
often with vertical lobe or dent, but sometimes only angled or curved, without dent; the
lobe or dent, if present, never forms shield-like plate along the short (vertical) surface of
scape (foot of scape) (e.g. Fig. 152, D). Anterior clypeal margin widely rounded, with
a notch medially (except for M. xavieri that has no medial notch).
Males (e.g. Fig. 154): scape relatively long, SIj > 0.60.
The lobicornis-group is one of the three most diverse species groups of the Old World
containing 22 species that, taken together, are distributed throughout the Palaearctic
Region. We have divided this group into five species complexes, based on worker charac-
ters (see Table 2.1).

Table 2.1. Characteristics of the species complexes within lobicornis species group
(based on workers).

Anterior clypeal margin not-notched medially - xavieri-complex

Lateral portion of clypeus raised into sharp ridge in front of antennal insertions, so
that antennal sockets distinctly separated from clypeal surface - exce/sa-complex

Lateral portion Scape at the base with vertical lobe or at least dent -
Anterior clypeal margin

of clypeus lobicornis-complex
notched medially

not raised into Alitrunk with very coarse, straight longitudinal rugae;
sharp ridge Scape propodeal spines long, species' means ESLI > 0.40;
in front of gradually petiole with short peduncle, its anterior surface almost
antennal curved or straight, meets with dorsal surface at an almost right or
insertions, so angled at faintly blunt angle, dorsal plate well developed, flat-
that antennal the base, tened, not inclined posteriorly - sulcinodis-complex
sockets lay on without
vertical Alitrunk with less coarse sinuous longitudinal rugosity;
the same level propodeal spines shorter, species' means ESLI < 0.35;
with clypeal lobe or
dent petiole of various shape, but never with well developed,
surface flattened dorsal plate - kscenkoi-complex

The species are distributed among the complexes as follows:

lobicornis-complex

lobicornis, anatolica, eidmanni, forcipata, jessensis, kirghisorum, Lobulicornis,

pisarskii, saposhnikovi, wesmaeli, zojae.

excelsa-complex

excelsa, tamarae, transsibirica.

Cri
 kasczenkoi-compiex

kasczenkoi, commarginata, angulinodis, displicentia, kamtschatica.

sulcinodis-compiex

sulcinodis, ademonia.

xavieri-compìex

xavieri.

luteola-group

Diagnosis. Workers (e.g. Fig. 161): frontal carinae curved outwards to merge with the
rugae that surround antennal sockets; frontal lobes slightly curved, frons wide, and frontal
lobes not extended. Scape gradually though quite strongly curved at the base, not angled,
with no trace of lobe or carina. Large and very hairy species.
Males (Fig. 163): scape short, SIj < 0.50.
We place two species to this group, both are quite unusual compared with all other Myr-
mica species. The first is found in Eastern Asia, including Japan, while the second is known
only from Taiwan: luteola, mirabilis.

myrmicoxena-group

Diagnosis. Workerless social parasite.

Queens (Fig. 171): frontal carinae merging with the rugae that extend to the occipital
margin, they do not curve outwards and do not merge with rugae that surround antennal
sockets; frons wide, frontal lobes not extended. Scape gradually while quite strongly curved
at the base, not angled, without any trace of lobe or carina. Petiole and postpetiole with
ventral lobes. Spurs on the middle and hind tibia reduced in various extents.
Males (Fig. 171): antennae 13-segmented, scape short, SIj < 0.40.
This group contains a single species: myrmicoxena.

pachei-group

Diagnosis. Workers (e.g. Figs 186, 250): alitrunk dorsum at least partly with trans-
verse rugosity. Scape gradually though distinctly curved at the base, not angled, with no
trace of lobe or carina. Anterior clypeal margin rounded or slightly prominent with no me-
dial notch. Petiole with a relatively short peduncle.
Males (Fig. 188): scape long, SIj > 0.90 (but males of only M. pachei are known).
Previously we placed in this group only Himalayan species that we believed to be rare and
unusual (Radchenko and Elmes 2001b). However, following a recent examination of the
Myrmica of China (Radchenko and Elmes 2009a), we have shown that the pachei-group is
much more diverse than previously expected. It now contains 14 species that are found in the

63
Himalaya, south-western and southern China: pachei, heterorhytida, hlavaci, multiplex,
phalacra, pleiorhytida, polyglypta, sculptiventris, schulzi, taibaensis, varisculpta, villosa,
zveii, yunnanensis.

ritae-group

Diagnosis. Workers (e.g. Figs 52, 211): frontal carinae merging with the rugae that
extend to the occipital margin, they do not curve outwards and do not merge with rugae
that surround antennal sockets; frontal lobes slightly curved, frons wide, and frontal lobes
not extended. Scape very long, often longer than head, smoothly curved at the base, with-
out any trace of lobe or carina. Anterior clypeal margin slightly convex and distinctly
notched medially. Posterior head margin has narrow collar-like ridge and posterio-ventral
angles of head prominent. Petiole very long and low (Pll > 1.35, often > 2.0); postpetiole
fig-shaped (seen from above). Propodeal spines very long; propodeal lobes sharply pointed
apically.
Males (e.g. Figs 71, 253): scape of various lengths, but unfortunately males are known
only for a few species.
Usually it is quite easy to recognise species from the ritae-group because their over-
all appearance, particularly their elongated petiole, fig-shaped postpetiole, very long
propodeal spines and relatively long appendages are quite distinct from other
Myrmica species. It is probably the strongest candidate among all the groups to warrant
subgenus status (see above). Despite this, preliminary studies suggest that the behaviour
and ecology of its species is very similar to that of other species groups. Furthermore,
it is the only morphologically-based group that clearly resembles extinct fossil species
(see Chapter 3.4), although it should be remembered that the modern ritae-group species
shared a common ancestor with species from the other Myrmica species groups 36 Ma,
4-8 million years after the known fossil species lived (see Jansen et al. 2010). At one time
the ritae-group was considered "archaic" containing just a few "relict" taxa, but our
recent work has shown it to be rich in species, currently we recognise 21 extant species
from the temperate zones of high mountains in the Himalaya, Meghalaya, southern China,
Taiwan, Vietnam, Burma, Thailand and even Indonesia. We are confident that con-
siderably more niae-group species will be described when the high mountains of this
region are better investigated. We are able to separate the group into three species com-
plexes, but these will probably be revised when the males of all the species are properly
known:

ritae-complex

Diagnosis. Workers (e.g. Fig. 211): head, alitrunk and waist very coarsely rugose
or/and reticulate, surface of head dorsum between rugae not punctated, smooth, appears
shiny.
Males (e.g. Fig. 253): scape short, SI, < 0.50.
This complex contains the majority (14) of species in the group, also some of the largest
Myrmica species found to date belong to this complex (i.e. M. gigantea and M. titanica), but
not all species are large (for example M. weberi workers are about the same size as those of
M. ruginodis): ritae, alperti, angulata, emeryi, gigantea, indica, margaritae, pararitae,
pulchella, serica, sinensis, titanica, urbanii, weberi.

64
 boltoni-complex

Diagnosis. Workers (e.g. Fig. 42): head, alitrunk and waist finely sculptured, rugulose
or even striated, surface of head dorsum between rugae distinctly punctated, appearing dull.
Males are unknown.
boltoni, collingwoodi, martensi.

draco-complex

Diagnosis. Workers (e.g. Fig. 69): head dorsum and alitrunk rugose, but waist finely
striated and punctated; surface of head dorsum between rugae punctated.
Males (Figs 71, 248): scape of various lengths, SIj < 0.50 or > 0.90.
The sculpture of the workers of this species complex seems to be intermediate between
the ritae- and frotom'-complexes, but the males are known for only two of the species
(M. draco and M. schoedli), the former has long scape and the latter short one. The com-
position of this complex might be better understood when additional material, including
males of the other species, is obtained.
draco, poldii, schoedli, yamanei.

rubra-group

Diagnosis. Workers (e.g. Fig. 213) : frontal carinae curved outwards to merge with the
rugae that surround antennal sockets, frontal lobes slightly curved, frons wide and frontal
lobes not extended. Scape very smoothly curved at the base, not angled and with no trace
of lobe or carina. Anterior clypeal margin convex, not prominent and without a medial
notch. ,-
Males (e.g. Fig. 215): scape long, SI2 > 1.00.
The group includes four species. Two of them (M. rubra and M. ruginodis) have
a Transpalaearctic distribution, while the others occur in East Asia and Taiwan: rubra,
arisana, kotokui, ruginodis.

rugosa-group

Diagnosis. Workers (e.g. Figs 44, 219): frontal carinae merging with the rugae that
extend to the occipital margin, they do not curve outwards and do not merge with rugae that
surround antennal sockets; frontal lobes slightly curved, frons wide and frontal lobes not
extended. Scape very smoothly curved at the base, not angled and with no trace of lobe or
carina. Anterior clypeal margin is convex and prominent, without a medial notch.
Males (e.g. Fig. 221): scape relatively short, SIj < 0.60.
By the main diagnostic features of workers, the species of the ragosa-group are superfi-
cially similar to those of the dshungarica-group, but differ from them by their greater size and
coarse body sculpture, and the males have a distinctly shorter scape.
We place nine species in this group, and divide it into two species complexes, the
rugosa- and cachmiriensis-comp\ex. The distribution of all species of this group is
Himalayan.

65
 rugosa-complex

Diagnosis. Workers (e.g. Fig. 219): promesonotal dorsum (seen in profile) forms
a more or less regular arch, mesonotal dorsum is not impressed transversely and gently
curves down to the propodeum so that the metanotal groove generally is shallow. Body
sculpture is fairly coarse.
rugosa, aimonissabaudiae, ereptrix, foreliana, hecate, rupestris.

cachmiriensis-compìex

Diagnosis. Workers (e.g. Fig. 44): promesonotal dorsum (seen in profile) does not
form regular arch, mesonotal dorsum impressed transversely, often saddle-shaped and
curves down abruptly to the propodeum to form deep and wide metanotal groove. Body
sculpture is generally finer than in the ragosa-complex.
cachmiriensis, ordinaria, wardi.

scabrinodis-group

Diagnosis. Workers (e.g. Figs 36, 99, 158, 239, 264) : scape gradually but quite strongly
curved or even angled at the base. Usually the bend of the scape is embellished by a hori-
zontal lobe, ridge or carina, but some species have no such structures on the bend at all.
The anterior clypeal margin is widely rounded and, with the exception of M. vandeli, has
no medial notch.
Males (e.g. Figs 232, 241, 266): scape short, SIj < 0.60 (usually < 0.40).
This is the most diverse of the Old World species groups; currently we place 26 spe-
cies in it. These are mainly associated with Europe and West Asia (Europe, Caucasus,
Asia Minor, West Siberia, Kazakhstan and Middle Asia), but only a few species are
found east of the Enisei River and none have been recorded in the Far East. We have
divide this group to five species complexes based on both worker and male characters
(Table 2.2).

Table 2.2. Characteristics of the species-complexes within scabrinodis species group

(based on a combination of worker and male features).

Males: scape longer (SI, > 0.50) - sabuleti-complex

Males: legs with very long hairs - scabrinodis-complex
Workers: frons narrower, mean Fl < 0.40; scape sharply
Males: Males: angulated, with small to large lobe - spec/'o/des-complex
scape legs with
Workers: Workers: scape slightly angulated, often with fine
shorter much
frons wider, ridge or carina - rugulosa-complex
(SI, <0.45) shorter
hairs mean Workers: scape gradually curved, with no trace
Fl > 0.40 of a lobe - bergi-complex

66
 The scabrinodis-group species are assigned to complexes as follows:

scabrinodis-complex

scabrinodis, aloba, cagnianti, tulinae.

bergi-complex

bergi, divergens, gallienii.

rugulosa-compïex

rugulosa, hellenica, constricta.

sabuleti-compìex

sabuleti, bibikoffi, hirsuta, lonae, vandeli, spinosior.

specioides -complex

specioides, bakurianica, georgica, kozakorum, salina, slovaca, stangeana, tpbiasi,

turcica.
The males of M. symbiotica are unknown, therefore we cannot reliably place it into any
species complex.

schencki-group
Diagnosis. Workers (e.g. Fig. 60, 242): scape strongly angled at the base and with
vertical lobe or ridge; the lobe or ridge always forms shield-like plate along the short (ver-
tical) surface (foot of scape) (e.g. Figs 60, E; 242, D). Anterior clypeal margin widely
rounded and notched medially.
Males (e.g. Fig. 244): scape short, SIj < 0.50.
There are 11 species in the schencki-group. In general, this group seems to be associ-
ated with hot grasslands and may have originated in steppe-like habitats. Many of the
species have quite limited distributions and are probably local endemics (e.g. M. pelops and
M. siciliana), but as a group it is widespread in southern habitats ranging from the Atlantic
to Pacific Oceans, including Japan. M. schencki is the most widespread species being locally
common throughout Europe.
schencki, caucasicola, deplanata, inucta, koreana, obscura, onoyamai, pelops,
ravasinii, siciliana, sinoschencki.

smythiesii-group

Diagnosis. Workers (e.g. Fig. 262): frontal carinae curve outwards to merge with
the rugae that surround the antennal sockets, frontal lobes slightly curved, frons wide
and frontal lobes not extended. Scape smoothly curved at the base, not angled and with no

67
trace of lobe or carina. Anterior clypeal margin is distinctly prominent without a medial
notch.
Males (e.g. Fig. 90): scape long, SI, > 0.70.
By the main diagnostic features, the species of this group are superficially similar to
those of the dshungarica-group, but differ from that group by having frontal carinae that
curve outwards and merge with the rugae that surround antennal sockets. In this respect
they are similar to the шога-group, but differ from that group by their distinctly more promi­
nent anterior clypeal margin, and by the somewhat shorter scape of the males.
We place to this group five species, all are distributed in the Himalaya and Tibet:
smythiesii, bactriana, fortior, ruzskyana, wittmeri.

tibetana-group

Diagnosis. Workers (e.g. Fig. 284) : frontal carinae curved at their anterior third, frontal
lobes strongly extended, wide and subsquare, FLI > 1.2. Antennal sockets not surrounded
by concentric rugae. Scape gradually curved at the base, not angled and with no trace of lobe
or carina. Anterior clypeal margin prominent, without medial notch.
Males (e.g. Figs 283, 285): scape of various length SI1 < 0.40 or > 0.75 (see below).
The unusual, distinct shape of the frontal carinae and frontal lobes (see above) de­
fines this group, which comprises three species found in Tibet, NE Afghanistan and the
Middle Asian mountains. It is rather confusing that a group with such a distinct feature
in workers should have males with such variable length scape: males of M. tenuispina
and M. orthostyla have short scape (SIj < 0.40), while males of M. tibetana have much
longer scape (SIj > 0.75). Originally, M. tibetana was described only from workers, and
males were described later by Ruzsky (1915), so it is possible that they were collected
from a different nest to the workers and may not be conspecific with M. tibetana.
We investigated Ruzsky's material (9 w and 5 m) with the labels "yni- Хату, сев. Бурхан
Будда, 10800' Козлов, кон. VII.Ol" [gorge Khatu, N of Burkhan Budda, 10,800', end of
vii.01, leg. Kozlov] and concluded that the workers with no doubt belong to M. tibetana
and that the males indeed have relatively long scape: there being no evidence that these
specimens came from different nests but at the same time no firm indication that they
were collected from the same nest. If more material of M. tibetana becomes available and
if it is confirmed that the males have a long scape then we would need to revise our species
groups and we would place M. tenuispina and M. orthostyla into a new tenuispina-group.
tibetana, orthostyla, tenuispina.

Species with uncertain taxonomic position

The following 10 species have unique combination of features that distinguish them
from each other and all the above species groups:
afghanica, brancuccii, curiosa, jennyae, nitida, petita, rhytida, tschekanovskii, vittata,
williamsi.

68
CHAPTER 3. REVIEW OF SPECIES

3.1. Material studied and its deposition

From 1758 until now, 426 names were originally ascribed to the genus Myrmica from
the Old World and 123 of these taxa were later excluded from Myrmica (transferred to other
genera or considered incertae sedis in the subfamily Myrmicinae). Of the remaining 303
names 19 are nomina nuda, nine are treated incertae sedis in the genus Myrmica, four are
unnecessary proposed replacement names and nine are unavailable names which leaves 262
valid names that certainly belong to the Old World Myrmica. These 262 names include eight
replacement names for preoccupied names, 107 names considered as junior synonyms of
other Myrmica species, besides the 147 Myrmica taxa that we consider here to be good
extant and extinct Old World species.
During the course of this study we designated four neotypes and described 60 new
species, and examined the type specimens of 163 taxa: thus, about 85% of the types of the
Old World valid Myrmica names were examined. We did not see the type specimens of 36
taxa with valid names, though in many cases we saw equivalent non-type material. The
types of at least 10 taxa most probably are lost. In addition, we examined a large quantity
of material from the whole geographical range of the Old World Myrmica species. This was
a mixture of specimens collected by us from throughout West and East Europe, Asia Minor,
Caucasus, Middle Asia, Siberia, Russian Far East, Mongolia, Korea, Japan and Taiwan.
Overall we examined more than 100,000 specimens and measured several thousands of
them (see Chapter 1.5.3 for details). We also worked with the North American Myrmica
species (several thousands of specimens were checked), but these data are not
included in this revision.
The Old World specimens examined are preserved in about 50 Museums, Institutions
and private collections (see below). The collection of Graham Elmes (ELMES) is probably
the largest and most representative collection of Myrmica ants in the World, the Old World
part contains > 65,000 mounted specimens from > 5,700 series, representing ca. 130
species, of these there are > 1,750 type specimens from 162 series for 67 of these taxa.
In this monograph we abbreviate the names of the collections using the name of the city-
home of the Museum, or the name of the private owner rather than using the "traditional"
acronyms for Museums found in many specialist taxonpmic publications because, in
our opinion, this is more helpful .to non-specialist readers. The following list is compiled
alphabetically:
BASLE - Naturhistorisches Museum, Basle, Switzerland (collections of
Santschi and partly of Forel) ;
BEIJING - Chinese Academy of Forestry, Beijing, China (collection of Wu and
Wang) ;
BERLIN - Museum für Naturkunde der Humboldt-Universität, Berlin, Ger-
many (collections of Stitz and partly of Mayr and Viechmeyer);

69
 BOLOGNA - Zoological Museum of the Bologna University, Italy (collection of
Menozzi);
BRUSSELS - Institute Royal des Sciences naturelles de Belgique, Brussels, Bel-
gium (collection of Bondroit);
BUDAPEST - Hungarian Natural History Museum, Budapest, Hungary (rich Mon-
golian and North Korean material);
COPENHAGEN - Zoological Museum of the University of Copenhagen, Denmark
(collection of Meinert);
DRESDEN - Staatliches Museum für Tierkunde Dresden, Germany (collection of
Viehmeyer) ;
EDIRNE - Trakya University, Edirne, Turkey (collection of Aktac);
ELMES - Centre for Ecology and Hydrology, Wallingford, UK (private collec-
tion of Elmes);
ESPADALER - Autonomous University of Barcelona, Bellaterra, Spain (private col-
lection of Espadaler);
GENEVA - Museum d'Histoire Naturelle, Geneva, Switzerland (collection of
Forel, part.);
GENOA - Museo Civico di Storia Naturale "Giacomo Doria", Genoa, Italy
(collection of Emery and some type specimens of Ruzsky);
GÖRLITZ - Staatlisches Museum für Naturkunde, Görlitz, Germany (collection
of Seifert);
GUILIN - Guangxi Normal University, Guilin, China (collection of Zhou);
HAMBURG - Geological-Palaeontological Institute and Museum, Hamburg Uni-
versity, Germany (amber collection) ;
HARVARD - Museum of Comparative Zoology of Harvard University, USA
(many types plus collections of Wheeler, Finzi and Weber);
HELSINKI - Zoological Museum of University of Helsinki, Finland (collection of
Nylander) ;
HLAVAC - Kosice, Slovakia (private collection of Hlavac);
IKSAN - Wonk Wang University, Iksan, Korea (collection of B.-J. Kim);
KAGOSHIMA - Kagoshima University, Kagoshima, Japan (collection of Yamane);
KIEV - Shmalhausen Institute of Zoology of the Ukrainian National
Academy of Sciences, Kiev, Ukraine (collections of Karawajew and
Radchenko) ;
LAUSANNE - Museum de Zoologie, Lausanne, Switzerland (collection of Kutter
and partly of Forel) ;
LEIDEN - National Museum of Natural History, Leiden, The Netherlands (col-
lection of Stärcke);
LIVERPOOL - Liverpool City Museum, UK (collection of Collingwood);
LONDON - The Natural History Museum, London, UK (collections of
Donisthorpe and Bolton, and very rich material from the whole
World);
MAASTRICHT - Natuurhistorisch Museum Maastricht, The Netherlands (collections
of Wasmann and van Boven);
MARTENS - University of Mainz, Germany (private collection of Martens);
MEI - Istituto di Zoologia, Roma, Italy (private collection of Mei);
MELBOURNE - Museum Victoria, Melbourne, Australia (collection of Curtis);

70
 MILAN -Museo Civico.di Storia Naturale di Milano, Italy (collections of
Rigato and Poldi);
MOSCOW - Zoological Museum of the Moscow State University, Moscow, Rus-
sia (collections of Arnoldi, Dlussky, and partly of Nasonov and
Ruzsky);
MZ-WARSAW - Museum Z.iemi, Warsaw, Poland (amber collection);
OXFORD - The Hope Entomological Collections, University Museum, Oxford,
UK (collections of Rothney, F. Smith, etc.);
PARIS - Museum Nationale d'Histoire Naturelle, Paris, France (collections of
Bernard and partly of Mayr, Emery, Forel, etc.) ;
PETERSBURG - Zoological Institute of the Russian Academy of Sciences, St.-Pe-
tersburg, Russia (part of material of Ruzsky's and Arnoldi's collec-
tions);
PRAGUE - National Museum of Natural History, Prague, Czech Republic (col-
lection of Sadil);
PIN-MOSCOW -Paleontological Institute of the Russian Academy of Sciences,
Moscow, Russia (amber collection);
SCHULZ - Leilingen, Germany (private collection of Schulz) ;
STOCKHOLM - Naturhistoriska Riksmuseet, Stockholm, Sweden (very rich Central
Asian and Himalayan material);
VERONA - Museo Civico di Storia Naturale di Verona, Italy (collection of
Scupola);
VIET - Institute of Ecology and Biological Resources, Hanoi, Vietnam (col-
lection of Viet) ;
VLADIVOSTOK - Biological and Soil Institute of the Russian Academy of Sciences,
Vladivostok, Russia (collection of Kupyanskaya) ;
WARD - University of California, USA (collection of Ward);
WARSAW - Museum and Institute of Zoology of the Polish Academy of Sci-
ences, Warsaw, Poland (collections of Pisarski, Czechowska and
Czechowski) ;
WERNER - Research Institute of Crop Production, Prague, Czech Republic (pri-
vate collection of Werner) ;
XIANYANG - China Northwest Scientifically-Technological University of Agricul-
ture and Forestry, Xianyang, China (collection of Wei).

3.2. The 303 names definitely attributable to genus Myrmica

In the list below, we differentiate the names into good extant species (bold italic), good
fossil species (bold), their synonyms (italic), incertae sedis (underlined) and nomina nuda,
preoccupied names, unnecessary proposed replacement names and unavailable names (nor-
mal type). The lists of taxa excluded from the genus Myrmica and details of the taxa incer-
tae sedis and of the unavailable names are given later in this chapter (see sections 3.5, 3.6
and 3.7). It is interesting to note the prevalence for the tradition of dedicating species to
a person, about 40% of names of good species (extant and extinct) usually being that of the
collector; about a third of names indicate some morphological attribute of the species, about
20% indicate the place of origin while only < 5% of names derive from habitat preference
or some other behaviour. •

7i
 1. aborigenica Zhigulskaya, 1991 (junior synonym of kamtschatica)
2. ademonia Bolton, 1995 (replacement name for aspersa)
3. afghanica Radchenko et Elmes, 2003
4. ahngeri Karawajew, 1926 (junior synonym of scabrinodis)
5. aimonissabaudiae Menozzi, 1939
6. alajensis Arnoldi, 1976 (junior synonym of juglandeti)
7. albuferensis Lomnicki, 1925 (junior synonym of aloba)
8. aloba Forel, 1909
9. aiperti Elmes et Radchenko, 2009
10. alpestris Arnoldi, 1934 (junior synonym of lobicornis)
11. alpina Stärcke, 1927 (junior synonym of lobulicornis)
12. anatolica Elmes, Radchenko et Aktac, 2002
13. angulata Radchenko, Zhou et Elmes, 2001
14. angulinodis Ruzsky, 1905
15. angustifrons Stärcke, 1927 (junior synonym of lobicornis)
16. apennina Stärcke, 1927 (junior synonym of lobulicornis)
17. arduennae Bondroit, 1911 (junior synonym of lobicornis)
18. arisana Wheeler,1930
19. arnoldii Dlussky, 1963
20. aspersa Kupyanskaya, 1990 (preoccupied name, replacement name - ademonia)
21. atlantica Stärcke, 1942 (nomen nudum)
22. bactriana Ruzsky, 1915
23. baikalensis Pisarski, 1969 (junior synonym of angulinodis)
24. bakurianica Arnoldi, 1970
25. balcanica Sadil, 1952 (junior synonym of specioides)
26. barchanica Ruzsky, 1905 (junior synonym of bergi)
27. bergi Ruzsky, 1902
28. bergiana Tarbinsky, 1976 (junior synonym of saposhnikovi)
29. bessarabica Nasonov. 1889 (incertae sedis)
30. betuliana Ruzsky, 1946 (junior synonym of schencki)
31. bibikoffi Kutter, 1963
32. bicolor Kupyanskaya, 1986 (preoccupied name, replacement name - displicentia)
33. breviceps Smith, F. 1878 (incertae sedis)
34. boltoni Radchenko et Elmes, 1998
35. brancuccii Radchenko et Elmes, 1999
36. brevispina Kuznetsov-ugamsky, 1927 (unavailable name)
37. brunescens Karawajew, 1929 (junior synonym of lobicornis)
38. burtshakabramovitshi Karawajew, 1929 (junior synonym of lobicornis)
39. busoni Chapman et Capco, 1951 (nomen nudum)
40. cachmiriensis Forel, 1904
41. cadusa Kim B.-J., Park et Kim J.-H., 1997 (junior synonym of excelsa)
42. cagnianti Espadaler, 1996
43. carbonaria Forel, 1902 (preoccupied name, replacement name - foreliana)
44. carinata Kupyanskaya, 1990 (preoccupied name, replacement name - taediosa)
45. caucasica Arnoldi, 1934 (not necessary proposed replacement name)
46. caucasicola Arnoldi, 1934
47. chersonensis Arnoldi, 1934 (unavailable name)

72
48. chinensis Viehmeyer, 1922 (junior synonym of kurokii)
49. clandestina Foerster, 1850 (junior synonym of rugulosa)
50. collingwoodi Radchenko et Elmes, 1998
51. commarginata Ruzsky, 1905
52. constricta Karawajew, 1934 (replacement name for minuta Karawajew, 1929)
53. curiosa Radchenko, Zhou et Elmes, 2008
54. cursor Smith F., 1878 (incertae sedis)
55. curvithorax Bondroit, 1920 (junior synonym of slovaca)
56. dagmarae Sadil, 1939 (junior synonym of deplanata)
57. daurica Motschoulsky, I860 (nomen nudum)
58. debilior Forel, 1902 (junior synonym oirupestris)
59. dentata Kuznetsov-ugamsky, 1927 (unavailable name)
60. denticornis Curtis, 1854 (junior synonym of lobicornis)
61. deplanata Emery, 1921
62. derzhavini Ruzsky, 1920 (junior synonym of sulcinodis)
63. dicaporiaccoi Menozzi, 1939 (junior synonym of aimonissabaudiae)
64. diluta Nylander, 1849 (junior synonym of ruginodis)
65. displicentia Bolton, 1995 (replacement name for bicolor)
66. divergens Karawajew, 1931
67. dolens Stärcke, 1942 (junior synonym of specioides)
68. draco Radchenko, Zhou et Elmes, 2001
69. dshungarica Ruzsky, 1905
70. eidmanni Menozzi, 1930
71. emeryi Radchenko et Elmes, 2009
72. eocenica Radchenko, Dlussky et Elmes, 2007
73. ereptrix Bolton, 1988
74. europea Finzi, 1926 (junior synonym of rubra)
75. everesti Donisthorpe, 1929 (junior synonym of rupestris)
76. excelsa Kupyanskaya, 1990
77. exigua Ruzsky, 1915 (preoccupied name, replacement name - ruzskyana)
78. eximia Kupyanskaya, 1990 (junior synonym of sulcinodis)
79. faniensis Boven, 1970 (junior synonym of karavajevi)
80. ferganensis Finzi, 1926
81. forcipata Karawajew, 1931
82. foreli Santschi, 1931 (junior synonym of lobicornis)
83. foreliana Radchenko et Elmes, 2001 (replacement name for carbonaria)
84. foreliella Arnoldi, 1976 (junior synonym of kirghisorum)
85. formosae Wheeler, 1929 (preoccupied name, replacement name -pulchella)
86. fortior Forel, 1904
87. fortior Crawley, 1920 (preoccupied name, replacement name -persiana)
88. furva Ruzsky, 1915 (junior synonym of bactriana)
89. gallienii Bondroit, 1920
90. georgica Seifert, 1987
91. gigantea (Collingwood, 1962)
92. grandis Kuznetsov-ugamsky, 1927 (unavailable name)
93. granulinodis Nylander, 1846 (incertae sedis)
94. hecate Weber, 1947 , • .

73
 95. hellenica Finzi, 1926
96. helleri Viehmeyer, 1922 (junior synonym of kurokii)
97. heterorhytida Radchenko et Elmes, 2009
98. himalayana Weber, 1947 (junior synonym of smythiesii)
99. hirsuta Elmes, 1978
100. hlavaci Radchenko et Elmes, 2009
101. hyungokae Elmes, Radchenko et Kim, 2001 (junior synonym of jessensis)
102. incurvata Collingwood, 1976 (junior synonym of angulinodis)
103. indica Weber, 1950
104. inezae Forel, 1902
105. infuscata Kuznetsov-ugamsky, 1927 (unavailable name)
106. inornata Menozzi, 1941 {nomen nudum)
107. intermedia (Wheeler, 1915)
108. intermedia Kuznetsov-ugamsky, 1927 (unresolved secondary homonym, incertae
sedis)
109. iranica Arnoldi, 1948 {nomen nudum)
110. iskanderi Arnoldi, 1976 (junior synonym of dshungarica)
111.'inuctaRadchenko et Elmes, 2006
112. jacobsoni Kutter, 1963 (junior synonym of gallienii)
113. jennyae Elmes, Radchenko et Aktac, 2002
114. jessensis Forel, 1901
115. juglandeti Arnoldi, 1976
116. kabylica (Cagniant, 1970)
117. kamtschatica Kupyanskaya, 1986
118. kamyschiensis Arnoldi, 1934 (junior synonym of bergi)
119. karavajevi (Arnoldi, 1930)
120. kasczenkoi Ruzsky, 1905
121. khamensis Ruzsky, 1915 (incertae sedis)
122. kievensis Karawajevi, 1934 (junior synonym of lobicornis)
123. kirbii Stephens, 1829 (nomen nudum)
124. kirghisorum Arnoldi, 1976
125. kirgisica Ruzsky, 1903 (junior synonym of bergi)
126. kоrеапа Elmes, Radchenko et Kim, 2001
127. kotokui Forel, 1911
128. kozakorum, sp nov.
129. kozlovi Ruzsky, 1915
130. kryzhanovskii Arnoldi, 1976
131. kurokiiFord, 1907
132. kutteri Finzi, 1926 (junior synonym of schencki)
133. lacustris Ruzsky, 1905 (junior synonym of stangeana)
134. laevinodis Nylander, 1846 (junior synonym of rubra)
135. laevinodoruginodis Ruzsky, 1896 (nomen nudum)
136. laurae (Emery, 1907)
137. lemasnei Bernard, 1967
138. limanica Arnoldi, 1934 (junior synonym of gallienii)
139. lissahorensis Stärcke, 1927 (junior synonym of lobicornis)
140. littoralis Kuznetsov-ugamsky, 1928 (junior synonym of eidmanni)

74
141. lobata Kutter, 1977 (nomen nudum)
142. lobicornis Nylander, 1846
143. lobicornosulcinodis Forel, 1874 (nomen nudum)
144. lobulicornis Nylander, 1857
145. lonae Finzi, 1926
146. longiscapus Curtis, 1854 (junior synonym of rubra)
147. longispinosa Mayr, 1868
148. lucidula Smith F., 1857 (nomen nudum)
149. luctuosa Smith F., 1878 (incertae sedis)
150. lulakeranica Arnoldi, 1948 (nomen nudum)
151. luteola Kupyanskaya, 1990
152. lutescens Menozzi, 1939 (junior synonym of cachmiriensis)
153. macrogyna Brian M. et Brian A., 1949 (junior synonym oiruginodis)
154. margaritae Emery, 1889
155. martensi Radchenko et Elmes, 1998
156. medvedevi Arnoldi, 1970 (junior synonym of stangeana)
157. mekongi Ruzsky, 1915 (junior synonym of kozlovi)
158. melanocephala Mayr, 1855 (nomen nudum)
159. microgyna Brian M. et Brian A., 1949 (junior synonym of ruginodis)
160. microrubra Seifert, 1993 (junior synonym of rubra)
161. minuta Ruzsky, 1905 (junior synonym of dshungarica)
162. minuta Karawajew, 1929 (preoccupied name, replacement name - constricta)
163. mirabilis Elmes et Radchenko, 1998
164. mixta Radchenko et Elmes, 2008
165. moravica Soudek 1922 (junior synonym of deplanata)
166. multiplex Radchenko et Elmes, 2009
167. mutata Sadil, 1952 (junior synonym oiruginodis)
168. myrmecophila Wasmann, 1910 (junior synonym of sulcinodis)
169. myrmicoxena Forel, 1895
170. neglecta Stärcke, 1942 (nomen nudum)
171. nevodovskii (Karawajew, 1926) (junior synonym of specioides)
172. nigripes Ruzsky, 1896 (junior synonym of sulcinodis)
173. nitida Radchenko et Elmes, 1999
174. nodicornis Chappell, 1886 (nomen nudum)
175. obensis Arnoldi, 1970 (junior synonym of gallienii)
176. obscura Finzi, 1926
177. onoyamai Radchenko et Elmes, 2006
178. ordinaria Radchenko et Elmes, 1999
179. orientalis Karawajew, 1926 (junior synonym of kotokui)
180. orthostyla Arnoldi, 1976
181. pachei Forel, 1906
182. paradoxa Radchenko, Dlussky et Elmes, 2007
183. pararitae Radchenko et Elmes, 2008
184. pechi (Samsinak, 1957) (junior synonym oikaravajevi)
185. pelops Seifert, 2003
186. perelegans Curtis, 1854 (junior synonym of sulcinodis)
187. persiana Weber, 1947 (replacement name for fortior and junior synonym of bergi)

75
188. petita Radchenko et Elmes, 1999
189. phaìacra Radchenko et Elmes, 2009
190. pilosiscapus Bondroit, 1920 (junior synonym of scabrinodis)
191. pisarskii Radchenko, 1994
192. plana Karawajew, 1927 (junior synonym of deplanata)
193. pleiorhytida Radchenko et Elmes, 2009
194. poidii Radchenko et Rigato, 2008
195. polyglypta Radchenko et Rigato, 2008
196. pradella Kutter, 1977 (nomen nudum)
197. puerilis Stärcke, 1942 (junior synonym of specioides)
198. pulchella Santschi, 1937 (replacement name for formosae)
199. pyrenaea Bondroit 1918 (junior synonym of lobulicornis)
200. ravasinii Finzi, 1923
201. reticulata Stärcke, 1942 (junior synonym of scabrinodis)
202. rhynchophora Foerster, 1850 (incertae sedis)
203. rhytida Radchenko et Elmes, 1999
204. rigatoi Radchenko et Elmes, 1998
205. ritae Emery, 1889
206. rolandi Bondroit, 1918 (junior synonym of aloba)
207. rubra Linnaeus, 1758
208. rudis (Mayr, 1868)
209. ruginodiformis Karawajew, 1929 (junior synonym of dshungarica)
210. ruginodis Nylander, 1846
211. ruginodolaevinodis Forel, 1874 (junior synonym oiruginodis)
212. rugosa Mayr, 1855 (nomen nudum)
213. rugosa Mayr, 1865
214. rugososmythiesi Forel, 1902 (not necessary proposed replacement name)
215. rugulosa Nylander, 1849
216. rugulosoides Forel, 1915 (junior synonym of scabrinodis)
217. rugulososcabrinodis Karawajew, 1929 (junior synonym of hellenica)
218. rupestris Forel, 1902
219. ruzskyana Radchenko et Elmes nom. nov. (replacement name for exigua)
220. ruzskyi Kiseleva, 1925 (junior synonym of kurokii)
221. ruzskyi Weber, 1947 (not necessary proposed replacement name)
222. sabuleti Meinert, 1861
223. salina Ruzsky, 1905
224. samnitica Mei, 1987 (junior synonym of laurae)
225. sancta Karawajew, 1926 (junior synonym of specioides)
226. saposhnikovi Ruzsky, 1904
227. scabrinodis Nylander, 1846
228. scabrinodoides Sadil, 1952 (junior synonym of specioides)
229. scabrinodolobicornis Forel, 1915 (junior synonym of sabuleti)
230. scabrinodorugulosa Nasonov, 1892 (nomen nudum)
231. scabrinodosabuleti Sadil, 1952 (junior synonym of scabrinodis)
232. schencki Viereck, 1903
233. schenckioides Boer et Nordijk, 2005 (junior synonym of schencki)
234. schoedli Radchenko, Elmes et Viet, 2006

76
235. schulzi Radchenko et Elmes, 2009
236. sculptiventris Radchenko et Elmes, 2009
237. serica Wheeler, 1928
238. siciliana Radchenko, Elmes et Alicata, 2006
239. silvestrianum (Emery, 1924) (junior synonym of specioides)
240. silvestrii Wheeler, 1928 (junior synonym oìruginodis)
241. sinensis Radchenko, Zhou et Elmes, 2001
242. sinica Wu et Wang, 1995 (junior synonym of excelsa)
243. sinoschencki Radchenko et Elmes, 2009
244. slobodensis Arnoldi, 1934 (not necessary proposed replacement name)
245. slovaca Sadil, 1952
246. smythiesii Forel, 1902
247. sontica Santschi, 1937 (junior synonym of ruginodis)
248. specioides Bondroit, 1918
249. speciolosa Seifert, 1988: 14 (unavailable name)
250. specularis Donisthorpe, 1929 (junior synonym of kozlovi)
251. spinosior Santschi, 1931
252. stangeana Ruzsky, 1902
253. starki Karawajew, 1929 (junior synonym of lobicornis)
254. strandi Arnoldi, 1934 (unavailable name)
255. striata Finzi, 1926 (junior synonym of specioides)
256. subacuta Kuznetsov-ugamsky, 1927 (unavailable name)
257. subalpina Ruzsky, 1915 (junior synonym of kozlovi)
258. subaltaica Ruzsky, 1925 (unavailable name)
259. subbrevispinosa Ruzsky, 1915 (junior synonym of kozlovi)
260. subopaca Arnoldi, 1934 (junior synonym of schencki)
261. sulcinodis Nylander, 1846
262. sulcinodolobicornis Ruzsky, 1895 (nomen nudum)
263. sulcinodoruginodis Donisthorpe, 1915 (junior synonym of sulcinodis)
264. sulcinodorugulosa Nasonov, 1889 (junior synonym of rugulosa)
265. sulcinodoscabrinodis Ruzsky, 1895 (junior synonym of sulcinodis)
266. sulcinodoscabrinodis Forel, 1915 (junior synonym of sulcinodis)
267. svartshevskii (Karawajew, 1915) (junior synonym of dshungarica)
268. symbiotica Menozzi, 1925
269. taediosa Bolton, 1995 (replacement name for carinata and junior synonym of transsi-
birica)
270. taibaensisWei, Zhou et Liu, 2001
271. tamarae Elmes, Radchenko et Aktac, 2002
272. tarbinskii Arnoldi, 1976 (junior synonym of dshungarica)
273. tenuispina Ruzsky, 1905
274. testaceus Sadil, 1953 (nomen nudum)
275. tianshanica Arnoldi, 1976 (junior synonym of juglandeti)
276. tibetana Mayr, 1889
277. tipuna Santschi, 1937 (junior synonym of arisana)
278. titanica Radchenko et Elmes, 2001
279. tobiasi Radchenko et Elmes, 2003
280. transsibirica Radchenko, 1994

77
 281. tschekanovskii Radchenko, 1994
282. tschuliensis Arnoldi, 1976 (junior synonym of specioides)
283. tulinae Elmes, Radchenko et Aktac, 2002
284. turcica Santschi, 1931
285. urbanii Radchenko et Elmes, 1998
286. ussuriensis Kuznetsov-ugamsky, 1928 (incertae sedis)
287. vandeli Bondroit, 1920
288. varisculpta Radchenko et Rigato, 2008
289. vicaria Kuznetsov-ugamsky, 1928 (junior synonym oí sulcinodis)
290. villosa Radchenko et Elmes, 1999
291. vittata Radchenko et Elmes, 1999
292. wardi Radchenko et Elmes, 1999
293. weberi Elmes et Radchenko, 2009
294. weii Radchenko et Zhou, 2008
295. wesmaeli Bondroit, 1918
296. williamsi Radchenko et Elmes, 1999
297. wittmeri Radchenko et Elmes, 1999
298. winterae (Kutter, 1973) (junior synonym oí karavajevi)
299. xavieri Radchenko, Elmes et Savolainen, 2008
300. yamanei Radchenko et Elmes, 2001
301. yoshiokai Weber, 1947 (junior synonym of ruginodis)
302. yunnanensis Radchenko et Elmes, 2009
303. zojae Radchenko, 1994
Thirty seven species names have junior synonyms (see list below). It is interesting to
note that the widespread boreal species M. ruginodis, M. lobicornis and M. sulcinodis and
the more southerly distributed M. specioides have been most frequently described under
other names. The widespread M. scabrinodis, M. schencki and M. rubra have each 4 or 5
junior synonyms, while M. karavajevi is the social parasite with most synonyms. This no
doubt reflects the relative abundance of these species and the history of intensive study in
Europe and western Siberia. It is unlikely that any of the junior synonyms of these species
will be shown to represent valid species based solely on morphological criteria, but possibly
genetical study might show some to represent morphologically cryptic species. On the other
hand, the Asian species, M. kozlovi and M. dshungarica have 4 and 5 junior synonyms,
respectively, both species are widespread and insufficient material is available to determine
whether some of their junior synonyms might represent morphologically different species.
aimonissabaudiae (1): dicaporiaccoi
aloba (2) : rolandi, albuferensis
angulinodis (2) : baikalensis, incurvata
arisana (1): tipuna
bactriana (1) : furva
bergi (4) : barchanica, kamyschiensis, kirgisica, persiana
cachmiriensis (1): lutescens
deplanata (3): dagmarae, moravica, plana
dshungarica (5): iskandari, minuta, ruginodiformis, svartshevskii, tarbinskii
eidmanni (1): littoralis
excelsa (2): cadusa, sinica
gallienii (3) : jacobsoni, limanica, obensis

78
 hellenica (1): rugulososcabrinodis
jessensis (1): hyungokae
juglandeti (2): alajensis, tianshanica
kamtschatica (1) : aborigenica
karavajevi (3): faniensis, pechi; winterae
kirghisorum (1) : foreliella
kotokui (1) : orienta lis
kozlovi (4): mekongi, specularis, subalpina, subbrevispinosa
kurokii (3): helleri, chinensis, ruzskyi
laurae (1): samnitica
lobicornis (10): alpestris, angustifrons, arduennae, brunescens, burtshakabra-
movitshi, denticornis, forel, kievensis, lissahorensis, starki
lobulicornis (3): alpina, apennina, pyrenaea
rubra (4): europea, laevinodis, longiscapus, microrubra
ruginodis (8) : diluta, macrogyna, microgyna, mutata, ruginodolaevinodis, sil-
vestrii, sontica, yoshiokai
rugulosa (2): clandestina, sulcinodorugulosa
rupestris (2): debilior, everesti
sabuleti (1): scabrinodolobicornis
saposhnikovi (1): bergiana,
scabrinodis (5): ahngeri, pilosiscapus, reticulata, rugulosoides, scabrinodosabuleti
schencki (4): betuliana, kutteri, subopaca, schenckioides
slovaca (1): curvithorax
smythiesii (1) : himalayana
specioides (9) : balcanica, dolens, nevodovskii, puerilis, sancta, scabrinodoides,
silvestrianum, striata, tschuliensis
stangeana (2): lacustris, medvedevi
sulcinodis (9) : derzhavini, eximia, myrmecophila, nigripes, perelegans, sulcino-
doruginodis, sulcinodoscabrinodis Ruzsky, sulcinodoscabrinodis
Forel, vicaria

3.3. Review of extant Myrmica species of the Old World

3.3.1. Rewiev of species

Myrmica ademonia Bolton, 1995

(Figs 5-7)
Myrmica ademonia Bolton, 1995: 277 (replacement name for Myrmica aspersa Kupyanskaya, 1990:
105); Radchenko 2005: 138.
Myrmica aspersa Kupyanskaya, 1990: 105, w, q,- m, Russian Far East (junior primary homonym of
Myrmica aspersa Smith F., 1865: 72, transferred to Tetramorium, see Chapter 3.8); Radchenko
1994b: 43, 1994c: 139, 1994Î: 89.
Myrmica sulcinodis: Collingwood 1976: 301 {part.), misidentification.

Type localities and type specimens. "Голотип - рабочий, Приморский край, Уссурийский заповедник,
гора Емельянова, южный склон, 9.VII.1969 (Купянская); паратнпы - 70 рабочих, 1 самка, 28 самцов из той
же серии, кроме того: Приморский край, Владивосток, 13.Х.1959 (Онисимова), 55 рабочих: (Купянская),

79
1 рабочий, 1 самка; Кедровая Падь, 18.IX.1973, 5.IX. 1974, 24.VI.1975 (Купянская), 30 рабочих, 5 самок;
Анисимовка, гора Криничная, 29.VII.1974 (Дахно), 25 рабочих; Шкотовскос плато, 1.VI.1973 (Купянская),
18 рабочих; Тигровая падь, 2.VI.1973 (Купянская), 75 рабочих, 2 самки; Партизанское плато, 2 4 -
27.VI.1973 (Купянская), 25 рабочих; Бснсвскос, 12.VI.1981 (Купянская), 13 рабочих; Таежный, 9.VI.1973
(Купянская), 53 рабочих; водораздел Амбы и Крауноеки, 11.VI.1973 (Купянская), 32 рабочих; Евсссвка,
12.VII.1978 (Купянская), 13 рабочих; верховье Бол. Уссуркн, 2, 13.VIII.1972, 16.VII.1973 (Купянская),
42 рабочих, 1 самка; Красноармейский р-н, Восток, 4.VI.1973 (Купянская), 7 рабочих [Holotype -
worker, Primorsky Krai, Ussurijsky Natural Reserve, Mt. Emelianova, southern slope, 9.VÜ.1969
(Kupyanskaya); paratypes: 70 w, 1 q, 28 m from the same scries, additionally: Primorsky Krai, Vladi-
vostok, 13.x.1959 (Onisimova), 55 w: (Kupyanskaya), 1 w, 1 q, Kedrovaya Pad', 18.ix.1973, 5.ix.1974,
24.vi.1975 (Kupyanskaya), 30 w, 5 q; Anisimovka, Mt. Krinichnaya, 29.vii.1974 (Dahno), 25 w; Shko-
tovskoe Plateau, l.vi.1973 (Kupyanskaya), 18 w; Tigrovaya Pad', 2.vi.l973 (Kupyanskaya), 75 w, 2 q;
Partizanskoe Plateau, 24-27.vi.1973 (Kupyanskaya), 25 w; Benevskoe, 12.vi.1981 (Kupyanskaya), 13
w; Taezhnyi, 9.vi.l973 (Kupyanskaya), 53 w; watershed of riv. Amba and Kraunovka, 11.vi. 1973
(Kupyanskaya), 32 w; Evseevka, 12.vii.1978 (Kupyanskaya), 13 w; upper reaches of riv. Bol. Ussurka,
2, 13.viii.1972, 16.vii.1973 (Kupyanskaya), 42 w, 1 q; Krasnoarmeisky Distr., Vostok, 4.VÌ.1973
(Kupyanskaya), 7 w].
Material examined. Holotype, w, "Прим. кр., Уссурийск, зап., г. Емельянова, юж. склон, 9.vii.69,
Купянская" [Russia, Primorsky Krai, Ussurijsky Natural Reserve, Mt. Emelianova, southern slope,
9.VÜ.1969, leg. Kupyanskaya] (MOSCOW); paratypes: > 20 w, 1 q, 7 m from the nest of holotype; 11
w, 2 q, "Тигровая Падь, 2.VÌ.1973 (Купянская)" [Russia, Primorsky Krai, Natural Reserve "Tigrovaia
Pad'", 2.VÌ.1973, leg. Kupyanskaya]; 2 w, "Прим. Край, Супут. запов., Змеиная горка, юж. склон 9.VII.69,
Купянская [Primorsky Krai, Suputinsky Nat. Res., Zmeinaia gorka, Southern slope, 9.VÜ.69, leg.
Kupyanskaya] (MOSCOW, VLADIVOSTOK); non-type material: about 100 w, > 20 q and m from
Russian Far East and Korean Peninsula.
Distribution (Map 2). Southern part of Russian Far East, Korean Peninsula.
Etymology. M. ademonia: apparently Bolton made-up this replacement name, loosely
basing it on the word ademptio meaning "to take away" (Barry Bolton, pers. comm.). In-
terestingly, in medicine the name means sad, weary or depressed which may have described
Bolton's mood at having to find another replacement name, but more appropriately, the

Map 2. Distribution of M. ademonia.

80
Ademonia family have red and black among their heraldic colours. M. aspersa: derived from
the Latin word (f.) aspersa = rugged with bumps to describe its overall sculpture.
Notes. M. ademonia is most similar to M. sulcinodis and is a member of the sulcinodis
species complex of the lobicornis-group. It differs from M. sulcinodis by its longer propodeal
spines, less coarsely sculptured petiole and postpetiole, and well-developed petiolar peduncle.
Ecology. In the Russian Far East M. ademonia is a forest species, living in mixed and de-
ciduous forests and nesting mainly in decayed wood, rarely in soil. In Korea it lives in mixed
and deciduous forests (oak, maple, elm, pine, fir) mainly at altitudes between 600 and 1500
m a.s.l., although it also has been found living in relatively wet places at lower altitudes (ca.
2ÖÖ m). As in Russia, it usually nests in decayed wood and very rarely in soil under stones.
The preference for decaying wood appears to be one of the main ecological differences from
M. sulcinodis that usually nests directly in coarse, sandy soil or under stones.

Myrmica afghanica Radchenko et Elmes, 2003

(Figs 8, 9)

Myrmica afghanica Radchenko et Elmes, 2003b: 3, w, Afghanistan.

Myrmica tibetana: Collingwood 1961: 56, misidentification.

Type localities and type specimens. NE Afghanistan, for details see material examined.
Material examined. Holotype, w, "Afghanistan, Pashki Nuristan, 6:vi.(S/c!) 48, leg. K. Paludan"
(COPENHAGEN); paratypes: 2 w, 1 q (dealate, specimen without postpetiole and gaster), with the
same labels as holotype (COPENHAGEN).
Distribution (Map 3). The species is known only from the type locality in the Nuristãn
(also spelled as Nurestãn) Province, NE Afghanistan.
Etymology. M. afghanica: from the name of the country Afghanistan with the adjective
suffix for nouns ica (from the Greek IKO) = belonging to, from, it is probably endemie to that
country.

Map 3. Distribution of M. afghanica.

81
 1

Notes. Males of M. afghanica are unknown and without males it is impossible to deter-
mine membership of a Myrmica species group with certainty. The morphological features
of the female castes suggest that M. afghanica belongs either to the dshungarica-group or
to the smythiesii-group. It most resemble species of the dshungarica-group, which are dis-
tributed in Middle Asian Mountains, in which case M. afghanica, like M. tenuispina, might
have invaded Afghanistan from the north. On the other hand, if it belongs to the smythiesii-
group it is probably a true endemie of the region.
Ecology is unknown.

Myrmica aimonissabaudiae Menozzi, 1939

(Figs 10-12)

Myrmica aimonissabaudiae Menozzi, 1939: 286, w, q, Pakistan (also described as a new in Weber
1947: 464); Eidmann 1942: 245; Collingwood 1961: 56, m; Pisarski 1967a: 378, 1967b: 47; Bolton
1995: 277; Radchenko and Elmes 2001b: 248.
Myrmica dicaporiaccoi Menozzi, 1939: 289, w, Pakistan (also described as a new in Weber 1947: 465);
Radchenko 1994b: 42; Bolton 1995: 278. Synonymy by Radchenko and Elmes 2001b: 248;
confirmed here.

Type localities and type specimens. M. aimonissabaudiae: "Come tipo ho scelto quattro operaie
raccolte a Gund (Valle del Sind) a 2080 m., in un vallone umido scendente fra coltivati in mezzo a pen-
dii rocciosi ed aridi, e una femmina raccolta ad Askole a 3100 m. in un' oasi nella Valle del Braldo".
M. dicaporiaccoi: "Due esemplari, l'uno raccolto ad Alchori a 2300 m. e l'altro a Shigar a 2200 m. in
oasi ombrose e irrigue nella valle del Shigar".
Material examined. Lectotype of M. aimonissabaudiae, w (designated by Radchenko and Elmes
2001b), "Karakorum, Gund, Valle Sind, 2080 m, 9.iv.l929" (MILAN); paralectotypes: 3 w, with same
labels as lectotype; 1 q, "Askole, Braldo, 3100 m, 10.viii.1929" (MILAN); lectotype of M. dicaporiaccoi,
w (designated by Radchenko and Elmes 2001b), "Sped. Karakorum, Shigar, 23.viii.29, 2200 m"
(MILAN); non-type material: > 250 w, 16 q and 3 m from Afghanistan, Pakistan, Indian, Nepal and
Bhutan.
Distribution (Map 4). NE Afghanistan, NE Pakistan, India: Kashmir, Himahal Pradesh,
West Bengal (Darjeeling), Sikkim, Meghalaya; Nepal, Bhutan.
Etymology. M. aimonissabaudiae: the name derives from the first name Aimone com-
bined with Sabaudi (meaning Savoyan). It is dedicated to the leader of the Italian Expedi-
tion to the Karakorum, 1929, which collected the type material. He was an Italian Prince,
HRH Aimone Roberto Margarita Guiseppe Torino Prince of Savoy-Aosta, Duke di Spo-
leto, 4 t h Duke d'Aosta. He was later proclaimed HM Tomislav II of Croatia but abandoned
claims to the throne in 1943 and died in Buenos Aires in 1948. It is a curiosity that the
original labels of the lectotype and paralectotypes are written as "M. amedeussabaudiae"
in red ink in Menozzi's own hand and Menozzi used this name once in his discussion of the
species (Menozzi 1939: 289). At one time we thought that the name "aimonissabaudiae"
was misprinted in the original description (Radchenko and Elmes 2001b) but clearly it is the
correct version. Perhaps, when he labelled the type specimens, Menozzi confused the Duke
di Spoleto's first name with that his uncle (Amedeo Duke di Aosta from whom he inherited
this title), but rectified this at some stage during the publication of the description in 1939.
M. dicaporiaccoi: this species was dedicated to the arachnologist and expeditionist, Prof.
Lodovico di Caporiacco (1900-1951), who collected some ants for Menozzi.

82
 Notes. M. aimonissabaudiae belongs to the rugosa species group (Radchenko and Elmes
2001b); its junior synonym, M. dicaporiaccoi, was erroneously placed to the rubra-group by
Radchenko (1994b). It most resembles two other Himalayan species - M. rugosa and
M. hecate - particularly by the distinct (but not coarse) sculpture on the nodes of the petb
ole and postpetiole. However it differs from the former by having much better developed
reticulation on the head dorsum, and from the latter by its less multidentate mandibles
(7-9 teeth on the masticatory margin of mandibles vs. > 10 in M. hecate).
In addition to the type specimens listed above, we have examined a "paratype" worker
from HARVARD with the labels: "Dusso (Braldo), 18-8-1929, 2400 m", "M. amedeus-
sabaudiae Men. det." (red ink, written by Menozzi's own hand) and "M.C.Z. Paratypus
28766" (red printed label). This specimen can not be considered as a type because its lo-
cality label does not correspond with the type localities, given in original description. More-
over, this specimen is not M. aimonissabaudiae, we have identified it as M. wardi; no doubt
it was part of Menozzi's original material, but it should be remembered that he included only
part of all the material identified by himself as M. aimonissabaudiae to the type series
(Menozzi 1939:289).
We also examined two workers of M. dicaporiaccoi from MILAN labelled by Menozzi
as "M. dicaporiaccoi, typus". The locality label of the first specimen is "Sped. Karakorum,
Shigar, 23.8.29, 2200 m" and of the second one - "Sped. Karakorum, Askole (Braldo),
V-29, 3100 m". The first agreed with Menozzi's data (see above) and description, therefore
we designated it as the lectotype of M. dicaporiaccoi. However, the second specimen (from
Askole) neither corresponds with type locality given in the original paper, nor is it actually
M. dicaporiaccoi; we identified it as M: wardi. Possibly there is another specimen in MILAN
from "Alchori, 2300 m" that will be a paralectotype of M. dicaporiaccoi. Menozzi
considered that M. dicaporiaccoi could be separated from the other species of the rugosa-
group by its shorter propodeal spines and eyes that are situated distinctly anterior to the
mid-point of the sides of the head. However, the length of the propodeal spines is quite
variable in many of the species, and the eyes of M. aimonissabaudiae are anterior to the

Map 4. Distribution of M. aimonissabaudiae.

83
midlength of the head (measured from anterior point of the clypeus). After examining the
rich material of M. aimonissabaudiae, we could find no consistent distinguishing charac-
ters for M. dicaporiaccoi and therefore considered these names as synonyms (Radchenko
and Elmes 2001b).
Ecology. A little more is known about the ecology of M. aimonissabaudiae compared to
most other Himalayan species because it appears to be one of the most common, based on
published data and material in collections. It has been found at quite a range of altitudes,
1300 to 3450 m a.s.l. (most of Menozzi's material was collected between 2200 m and 2600
m), in both natural and disturbed habitats. The latter includes fields and parks where it
prefers relatively wet places being quite common in irrigated fields; Menozzi (1939: 289)
particularly mentions that it was found near oasi in the middle of quite arid regions. More
recent records suggest that M. aimonissabaudiae is most abundant in semi-natural habitats
where nests have been found in open grasslands with or without shrubs, and in deciduous
and coniferous forests, where it nests both in the ground and in rotten wood (Philip
Ward, pers. comm.). Based on these reports it appears to occupy a niche similar to that of
M. ruginodis in Europe.

Myrmica aloba Forel, 1909

(Figs 13-15)

Myrmica scabrinodis var. aloba Forel, 1909: 103, w, Spain; Santschi 1925: 340, w, q, nec m; Weber
1950: 204.
Myrmica rugulosoides var. aloba: Finzi 1926: 95, w, nec m.
Myrmica aloba: Santschi 1931b: 340; Collingwood and Yarrow 1969: 56; Collingwood 1978: 67;
Espadaler 1981b: 192; Tinaut and Espadaler 1987: 61, m; Seifert 1988: 29; Radchenko 1994b: 42;
Bolton 1995: 277; Espadaler 1996: 73.
Myrmica rolandi Bondroit, 1918: 101, w, q (part.), nec m; 1920b: 300, w, q; Finzi 1926: 89, w, q, nec
m; Bernard 1967: 115, nec Bondroit 1920a: 150, m. Synonymy by Seifert 1988: 29; see also Bolton
1995: 282; confirmed here.
Myrmica sulcinodis var. rolandi: Santschi 1919: 244; Menozzi 1922: 325.
Myrmica scabrinodis var. (st., subsp.) rolandi: Emery 1921: 40; Santschi 1925: 340; Santschi 193lb:
344; Weber 1950: 208.
Myrmica albuferensis Lomnicki, 1925: 2, w, Balearic Isl. Synonymy by Finzi 1926: 89 (as synonym of
M. rolandi); Santschi 1931b: 340 (as synonym of M. aloba); see also Bernard 1956: 257; Seifert
1988: 29; Bolton 1995: 278; the latter confirmed here.

Type localities and type specimens. M. aloba: "Ronda (Andalousie) et Sierra Guadarama".
M. rolandi: "Pyrénées-Orientales: Banyuls et Font-Romeu; aussi en Espagne" (see also Notes below).
M. albuferensis: "Altera w in Albufera 15.vi., altera in La Porassa 12.vii. collecta est."
Material examined. Lectotype of M. aloba, w (bottom specimen on the pin with 2 w, designated
here), "M. scabrinodis Nyl. v. aloba For. W type Andalousie] (Vogt)", "v. M. aloba For.", "Coll.
A. Forel" (GENEVA); paralectotype: upper w on the pin with lectotype (GENEVA); lectotype of
M. rolandi, w (upper specimen on the pin with 2 w, designated here), "Pyrénées Orientales, 1917",
"Coll. R. 1. Sc. N. В. France", "R. I. Sc. N. В. I. G. 21.400", "Myrmica Rolandi Bondr., type"
(BRUSSELS); paralectotypes: bottom w on the pin with lectotype; 3 w, "Pozuello, Espagne", "Coll. R.
I. Sc. N. B. Espagne", "R. I. Sc. N. B. I. G. 21.400", "Myrmica Rolandi Bondr., type"; 1 q, "Espagne",
"Coli. R. I. Sc. N. B. Espagne", "R. 1. Sc. N. B. I. G. 21.400", "Myrmica Rolandi Bondr., type"; 1 q (alate),
"Madrid", "Coli. R. I. Sc. N. B. Espagne", "R. I. Sc. N. B. I. G. 21.400", "Myrmica Rolandi Bondr., type";

84
1 q, " Carthagena, G. Schramm", "Coll. R. I. Sc. N. B. Espagne", "R. I. Sc. N. B. I. G. 21.400", "Myr-
mica Rolandi Bondr., type" (BRUSSELS); non-type material: > 100 w, 25 q and 15 m from Spain.
Distribution (Map 5). Iberian Peninsula, Balearic Isl., French Pyrenees.
Etymology. M. aloba: from a combination of loba derived from the Greek
= lobe [of ear] and the prefix a (a) = not or without, to indicate the mostly absent (or ex-
tremely reduced) lobe on the bend of the scape compared to its close relative M. scabri-
nodis. M. rolandi: presumably dedicated to a Mr. Roland, we have no information as to who
he was, just possibly, given Bondroit's militarist views (see biographies in Chapter 1.6) it
may have been the French flying ace, Roland Garros (1889-1918) who was shot down on
5 th October 1918. M. albuferensis: a combination of the name Albufera with the Latin suf-
fix ensis = place of origin, to indicate that the types were taken in Albufera, a region of the
island of Mallorca (now Natural Park Albufera).
Notes. We have investigated type specimens of M. aloba and M. rolandi (see above) and
confirm the earlier established synonymy. M. aloba clearly belongs to the scabrinodis species
group. It is most similar to M. scabrinodis: the female castes of the two species have
a strongly curved (though not angularly bent) scape, but M. aloba differs from
M. scabrinodis by the absence of a lobe on the bend of the scape having at most only a small
longitudinal ridge. Males were described relatively recently (Tinaut and Espadaler 1987)
based on nest samples containing both workers and males; the males of M. aloba are very
similar to those of M. scabrinodis by the length of scape and character of pilosity on the
scape and legs.
The taxonomic history of this species is somewhat confused being tied-up with earlier
attempts to describe the males. Santschi (1925) noted that the scape of M. scabrinodis
aloba males was similar to that of M. pilosiscapus (now a synonym of M. scabrinodis).
Then, a year later, Finzi (1926) described males of M. aloba and stressed that the scape is
as long as four or five basal funicular segments. Santschi (1931b) reinvestigated Finzi's
specimens and based on the male characters he believed the species to be related to M. sa-
buleti, whose males have relatively long scapes, and was not a variety of M. scabrinodis.
Consequently he raised M. aloba to the species level.

Map 5. Distribution of M. aloba.

85
 The history of the junior synonym M. rolandi is equally confused: Bondroit (1918) de-
scribed M. rolandi based on all three castes, but later Bondroit (1920a) noted that the male
from Font-Romeu that he had described as M. rolandi, was in fact a male of M. pilosiscapus,
so he redescribed males and excluded Font-Romeu from the type localities. These newly
described males had scapes as long as, or slightly longer than three basal funicular segments
and having rather short hairs; additionally, the hind tibiae had semierect hairs, being some-
what longer than the half of tibia's width. Finally he decided (Bondroit 1920b) that in both
previous papers he had mixed two species under the name "rolandi" and he redescribed
workers and queens of M. rolandi and believed that males were unknown. Finzi (1926) de-
scribed males of this species yet again, observing that the scape is equal to the length of
four or five basal funicular segments (most probably Finzi's male was M. sabuleti or even
M. vandeli).
Ecology. M. aloba is almost exclusively an Iberian form that is associated with hot, but
moist grassland conditions, where it nests in very similar places to M. scabrinodis living in
cooler climates. In the extreme south-west of Europe near cape St. Vincent in Portugal,
we observed many populous nests living in the tops of large grass tussocks in river flood
meadows that are grazed by cattle. They have been found on marshy grassland in north
Portugal, where they were hosts to the Large Blue Butterfly Phengaris alcon (Denis et
Schiffermüller) (I. Wynhoff, pers. comm.). In Spain in summer they are often found
foraging on rocks near dried-up water-courses, lakes and springs. In the mountains of Soria
we have found nests living in soil on cattle meadows near a spring. Here M. aloba lived
sympatrically with M. scabrinodis and both species appeared to be hosts to Ph. alcon. The
ecological separation of these species on such sites has not yet been studied in detail, but
it seemed that M. aloba avoided the very boggy conditions close to the spring, where some
M. scabrinodis nests were observed. Sexuais were found in nests from the end of July until
mid-September, which indicates that M. aloba swarms in late summer.

Myrmica alperti Elmes et Radchenko, 2009

(Fig. 16)

Myrmica alperti Elmes et Radchenko, 2009: 117, w, Nepal.

Type localities and type specimens. Nepal, for details see material examined.
Material examined. Holotype, w, Nepal, 3200 m, Thodung, 2-9.iv. 1973 (leg. J. Martens), Conifer-
enwald, "M. indica Weber det Radchenko and Elmes", [6] (BASLE); paratypes: 14 w with the same
labels; 9 w, Nepal, Jiri-Thodung, 28.V.1976 (leg. W. Wittmer and С Baroni Urbani), "M. indica Weber
det Radchenko and Elmes"; 1 w, Nepal, Shiralaybis, Jiri-grat, 2200 m, 8.vi.1973 (leg. J. Martens),
"M. indica Weber det Radchenko and Elmes" (BASLE, KIEV, ELMES).
Distribution (Map 6). The species is knows only from the type series from Nepal.
Etymology. M. alperti: this species was dedicated to our friend and colleague, the Amer­
ican Dr. Gary Alpert of Harvard University, USA, who gave us Myrmica samples that he col­
lected in Nepal and elsewhere.
Notes. M. alperti belongs to the ri/ae-complex of the ritae species group. It closely re­
sembles M. indica, but differs by its more stocky body and shorter appendages, by the longer
propodeal spines, by the shape of petiole, and especially by its bluntly rounded propodeal
lobes. Queens and males are unknown.

86
 Map 6. Distribution of M. alperti.

Ecology. Nothing is known of its ecology, except that it was found at altitudes between
2000 and 3000 m a.s.l.

Myrmica anatolica Elmes, Radchenko et Aktac, 2002

(Figs 17-19)

Myrmica anatolica Elmes, Radchenko et Aktac, 2002: 167, w, q, m, Turkey.

Type localities and type specimens. Anatolian part of Turkey, for details see material examined.
Material examined. Holotype, w, NE Turkey, Trabzon Reg., nr. Uzungöl, 40°34'29"N, 40°17'47"E,
alt. 2372 m, 15.viii.1999, TR-27 (leg. J. E. and G. W. Elmes) (EDIRNE); paratypes: 12 w, 4 q, 3 males
from the nest of holotype; 30 w, same locality, site, date and collectors, TR-26; 320 w, 1 q, 3 q, 36
males (12 colonies), same locality, date and collectors, but co-ordinate 40°38'33"N, 40°16'21"E, alt.
2397 m, TR-10, 11, 12, 13, 14, 15, 16, 17, 18, 19, 22, 23; 72 w (4 colonies), NE Turkey, Trabzon Reg.,
ca 5 km S Uzungöl, up mountain, alt. 1800 m, 17.viii.2000, TR-154, 155, 156, 157 (leg. G. W. Elmes);
13 w, NE Turkey, Trabzon Reg., ca 5 km S Uzungöl, up mountain, alt. 1800 m, 17.viii.2000, No.
00/171a (leg. K. Kiran); 4 w, NE Turkey, Trabzon Reg., Hamsiköy, 41 km S Trabzon, alt. 1400 m,
25.V.1974, No. 42, 61 (leg. N. Aktac); 8 w, same locality and collector, 8.VÌ.1975, No. 117; 2 w, N
Turkey, Erzurum Reg., 113 km NE Erzurum, Oltu-Uzunoluk, alt. 1900 m, 23.ix. 1979, No. 2122 (leg.
N.Aktaç); 10 w, NE Turkey, Rize Reg., SofMadenli (Sof Cayeli), S of Kaptanpasa, between Cataldere
and Inçesu, alt. 1800 m, 12.viii.2000, TR-86 (leg. G. W. Elmes); 53 w, 4 males (3 colonies), NE Turkey,
Rize Reg., S of Madenli (S of Cayeli) ca. between Inçesu and Baltas/Balikli Gölu Mts., alt. 2400 m,
12.viii.2000, TR-88, 89, 90 (leg. G. W. Elmes) ; 83 w (5 colonies), NE Turkey, Rize Reg., S of Madenli
(S of Cayeli), ca. east of Baltas/Balikli Gölu Mts., nr. Baskoy, alt. 2300 m, 12.viii.2000, TR-91, 93, 94,
95, 96 (leg. G. W. Elmes); 18 w, NE Turkey, Rize Reg., Ça irán yaylasi, alt. 1850 m, ll.viii.2000, No.
00/4, (leg. K. Kiran); 1 worker, NE Turkey, Rize Reg., Yenisu, alt. 1850 m, 12.viii.2000, No. 00/39
(leg. K. Kiran); 2 q, NE Turkey, Rize Reg., Basköy, alt. 2300 m, 12.viii.2000, No. 00/41 (leg. K. Kiran);
79 w, 1 q (9 colonies), NE Turkey, Rize Reg., Inçesu, alt. 2300 m, 12.viii.2000, No. 00/42, 44, 46, 47,
48, 50, 54, 55, 56 (leg. K. Kiran); 2 w, NE Turkey, Rize Reg., Rüzgarli Köyü, alt. 2300 m, 13.viii.2000,

87
No. 00/88 (leg. К. Kiran); 44 w, N Turkey, Erzincan - Refahiye, 50 km W Erzincan Gülendag, alt. 1800
m, 14.vi.1982, No. 2225 (leg. N. Aktac); 14 w, Central Turkey, 130 km N Ankara, Cankir, 18.vi.1985,
No. 103 (leg. N. Aktac) (EDIRNE, KIEV, WARSAW, ELMES).
Distribution (Map 7). Anatolian part of Turkey.
Etymology. M. anatolica: from the name Anatolia (a region of Turkey) with the adjec-
tive suffix for nouns ica (from the Greek IKO) = belonging to, from, to indicate that it is
probably an endemic of that region.
Notes. M. anatolica belongs to the lobicornis species group and well differs from
M. lobicornis by its much smaller lobe at the scape base, wider frons and more rounded
petiole node (seen in profile).
Ecology. M. anatolica has been found only on mountains in Anatolia at altitudes be-
tween 1800 and 2400 m a.s.l., where it is fairly common on both intensively grazed and
more natural alpine meadows. It is also found in damper, shaded hay meadows at slightly
lower altitudes and on steep slopes at the edge of mixed and pine forests. It nests in the soil,
usually under stones. M. anatolica frequently lives sympatrically with M. lobicornis, and in
the field (using a hand lens), it could be mistaken for a species from the scabrinodis-group.
Sexuais were found in nests in mid-August indicating late summer swarming.

Map 7. Distribution of M. anatolica.

Myrmica angulata Radchenko, Zhou et Elmes, 2001

(Figs 20, 21)

Myrmica angulata Radchenko, Zhou et Elmes, 2001: 213, w, S China; Radchenko, Elmes and Viet
2006: 42, q.
M. margaritae: Wei С et al. 2001: 561, misidentification.

Type localities and type specimens. Southern China and northern Vietnam, for details see mate­
rial examined.

88
 Material examined. Holotype, w, S China, Guangxi Province, Xing An, 500 m, 7.VÜ.1994, No. 2 (leg.
S. Zhou) (LONDON); paratypes: 4 w from the same nest; 1 w, S China, NE Guangxi Province, Hua Ping,
840 m, 16.viii.1998, HP-25 (leg. J. Fellowes); 2 w, S China, Guangxi Province, Da Yao Shan, 1020 m,
19.ix.1998, DYS-108 (leg. J. Fellowes) (KIEV, GUILIN, ELMES); non-type material: 2 w, China, Hubei
Prov., Xiang Feng County, No. 10, 10.viii.2004 (leg. Wei Wang); 2 w, China, Pingbaying Natural
Reserve, 10.viii.2004 (leg. Wei Wang); 45 w, N Vietnam, Lai Chau Prov., western slopes of Mt. Fan Si
Pan (W Cong Troi), 2100-2200 ra, 29.iv-6.v.2002, Nos. Eg 02-VN-179, 313, 315 (leg. K. Eguchi, from
several nests); 103 w, 1 q, N Vietnam, Cong Troi, Sa Pa distr., Lao Cai province, 28.iv-5.v.2002, 2000-
2200 m, Nos Eg02-VN-159, 183, 253, 308, 311 (leg. K. Eguchi, from several nests).
Distribution (Map 8). Southern China and northern Vietnam.
Etymology. M. angulata: from the Latin word angulata = angular, with angles, to reflect
the more strongly angled base of the scape, compared to most other niae-group species.
Notes: M. angulata belongs to the ritae-complex of ritae species group, and well differs
from all other known species in this complex by the combination of coarse rugosity and
dense punctation on the petiole and postpetiole. Records of M. margaritae from China (Wei
С et al., 2001) in fact refer to M. angulata (material examined). Males are unknown.
Ecology. In southern China M. angulata nests were found in rotten wood in forests at
altitudes 500-1000 m. In northern Vietnam it also lives in mixed forests between 1700 and
2200 m a.s.L, nests being found in rotten wood, in soil under logs and pieces of bark, or
under stones. The altitude difference between the China and Vietnam populations most
probably relates to regional climatic differences.

Map 8. Distribution of M. angulata.

Myrmica angulinodis Ruzsky, 1905

(Figs 22-24)

Myrmica scabrinodis angulinodis Ruzsky, 1905: 689, w, q, Russia; Emery 1921:41; Ruzsky 1936: 94;
Weber 1950: 204.

89
Myrmica angulinodis: Collingwood 1962: 217; Pisarski 1969a: 227, 1969b: 296; Dlussky and Pisarski
1970: 86; Collingwood 1976: 302; Dmitrienko, Petrenko 1976: 16; Pisarski, Krzysztofiak 1981:
155; Radchenko 1994b: 43, 1994c: 139, 1994f: 84, m, 1994g: 109; Bolton 1995: 277; Wu and Wang
1995: 94; Wei С et al. 2001: 561; Kim B.-J. 1996: 179, 2003: 2; Radchenko 2005: 138; Pfeiffer et
al. 2007: 4, nec Collingwood 1976: 302, misidentification.
Myrmica saposhnikovi subsp. baikalensis Pisarski, 1969a: 228 (first available use ai Myrmica scabri­
nodis subsp. saposhnikovi var. baikalensis Karawajew, 1931a: 28, w, q, Russia); Bolton 1995:
277. Synonymy by Radchenko 1994f: 84; confirmed here.
Myrmica baikalensis: Dmitrienko 1979: 94; Dmitrienko, Petrenko 1976: 10.
Myrmica incurvata Collingwood, 1976: 301, w, North Korea; Radchenko 1994b: 43, 1994c: 139.
Synonymy by Radchenko 2005: 301; confirmed here.
Myrmica forcipata: Kupyanskaya 1990: 107, misidentification.
Myrmica kasczenkoi: Kupyanskaya 1979: 128, 1985: 77, 1986b: 95, 1990: 113, misidentifications.

Type localities and type specimens. M. angulinodis: "Иркутская губ., ок. р. Тупки, бескр. самка
поймана на берегу реки 17.VIII.98 (Иловайский); речка Улунтай, правый прит. р. Слюдянки, конец VI.
1902 (Дравсрт)" [Russia, Irkutsk Prov., near riv. Tupka, dealate queen on the riverbank, 17.viii.1898,
leg. Ilovajsky; riv. Uluntai, right inflow of the riv. Slyudianka, end of vi.1902, leg. Dravert].
M. baikalensis: "Strand des Baikalsees bei dem Dorf Listwenitschnoje, 15.VII. (Nr. 5272), 11 w und
1 flügelloses q. -Ebenda, 18.VII. (Nr. 5259), 4 w. -Tanchoj-Mischicha, ostlicher Strand des sudlichen
Teils des Baikalsees, 1-6.VIII (Nr. 5238), 7 w". M. incurvata: "Locality No: 202. Ryang-gang Province:
Chann-Pay plateau, Sam-zi yan, 1700 m, 12 w. Described from 12 w taken at 1700 m in forest litter at
Sam-zi-yan on the Chann-Pay plateau, Ryang-gang Province by J. Papp and S. Horvatovich, 26 August
1971. Holotype and paratypes in the Hungarian Natural History Museum, Budapest".
Material examined. Neotype of M. angulinodis, w (designated by Radchenko, 1994f), "Иркутская
обл., Качукский р-н, Ацикан, No. 285, 28.08.86 (Плсшанов)" [Russia, Irkutsk Prov., Kachuksky Distr.,
Atsykan, No. 285, 28.viii.1986, leg. Pleshanov] (MOSCOW); syntypes of M. baikalensis: 9 w, 2 q,
"Бср. Байкала, с. Лиственничное, 15.vii.30, Г. Шпст" [shore of Lake Baikal, vil. Listvcnnichnoe,
15.vii.1930, leg. G. Shpet], "5272 Coll. Karawajewi", "M. (Myrmica) saposhnikovi Ruzs. var. baikalen­
sis Karaw. typ", "Syntypus Myrmica saposhnikovi var. baikalensis Karaw."; 4 w, same labels as above,
but collecting date 18.vii.30; 4 w, "Танхой-Мишиха, вост. бср. южн. Байкала, Г. Шпст" [Tankhoi-Mishi-
kha, eastern shore of southern Lake Baikal, leg. G. Shpet], "5238 Coll. Karawajewi", "M. [Myrmica)
saposhnikovi Ruzs. var. baikalensis Karaw. dct.", "Syntypus Myrmica saposhnikovi var. baikalensis
Karaw." (KIEV, MOSCOW); holotype of M. incurvata, w, "North Korea, Prov. Ryang-gang, Plateau
Chann-Pay, Sam-zi-yan, 1700 m, 29 Aug. 1971", "No. 202, leg. S. Horvatovich and J. Papp ", "Holo-
typus w Myrmica incurvala Collingwood С A." (BUDAPEST); paratypes: 9 w with same label as holo­
type (BUDAPEST, MOSCOW); non-type material: > 200 w, several tens of q and m from Siberia,
Mongolia, Russian-Far East and Korean Peninsula.
Distribution (Map 9). Siberia, from Kuznetsky Alatau until Pacific Ocean, Mongolia,
N China, Korean Peninsula.
Etymology. M. angulinodis: name from a combination of the Latin words angularis =
angled and nodus = knot or lump, to describe its sharply angled petiolar node (seen in pro­
file). Myrmica baikalensis: from Baikal, the region of Siberia where the types were found,
combined with the Latin suffix ensis = place of origin. Myrmica incurvata: from the Latin
word incurvus = bent with the adjectival suffix ata, to describe both the incurved propodeal
spines and curved, but not angled, base of its scape: it appeared to be "a rather curvy
species" (Cedric Collingwood, pers. comm.).
Notes. M. angulinodis belongs to the lobicornis species group. Unfortunately, Ruzsky's
original types seem to be lost and Radchenko (1994Í) designated a neotype for this species.
Female castes are characterised by i) a scape that is distinctly angled at its base (not

90
gradually curved), without a vertical lobe or dent on the bend, ii) its sharply angled, sub-
triangular petiolar node, seen in profile, and iii) propodeal spines that curve inwards when
viewed from above. By the last two features, M. angulinodis is similar to M. forcipata, but
the latter has at least a small vertical dent at the scape base. On the other hand, by the
shape of scape M. angulinodis resembles M. kamtschatica, but differs by its subtriangular
petiolar node compared to the more rounded node of the latter.
We have examined abundant material, collected in various areas of Siberia, Mongolia
and Korea, including the type localities. We compared the syntypes of M. baikalensis with
this material and also with Ruzsky's original description of M. angulinodis, and concluded
them to be synonyms.
Collingwood (1976) described M. incurvata from North Korea and compared it with
M. forcipata, from which it differed by the absence of a vertical dent at the scape bend. On
the other hand, M. incurvata is very similar to M. angulinodis, with which it shares all the
main diagnostic features, including morphometries. The only feature that might separate
M. incurvata from M. angulinodis, is a less angled base of the scape. However, we found
many specimens with less angled scape than "typical" among the abundant M. angulinodis
material examined, even among specimens from the same nest, and even among the
paratype series of M. incurvata some specimens have strongly angled scape. We conclude
that M. incurvata falls within the infraspecific variability of M. angulinodis, and therefore
consider these names as synonyms.
Prior to Radchenko (1994b, c, f), the taxonomy of the East Palaearctic Myrmica was
poorly understood and misidentifications were made by Kupyanskaya (1979, 1985, 1986b,
1990) who determined specimens of M. angulinodis as M. kasczenkoi or M. forcipata, and
by Collingwood (1976 - repeated by Kim B.-J. 1996), who identified three workers of
M. kamtschatica from the Chang-Pay plateau (North Korea) as M. angulinodis.
Ecology. M. angulinodis is a typical boreal species associated predominantly with dif-
ferent kinds of taiga forest (spruce, larch, fir, birch and mixed forests), preferring areas with
a more open canopy and often living in clearings and glades; in some old forests it is one of

Map 9. Distribution of M. angulinodis.

91
the commonest ant species accounting for 60% of ant nests (see Berman et al. 2010 and ref-
erences therein). In the north of its range M. angulinodis lives at lower altitudes, from about
sea level to some hundred meters, preferring open south facing mesoxerophyous habitats
that are relatively rare in the Kolyma region, where the largest colony excavated contained
ca. 1700 workers and 30 queens (Berman et al. 2010). In the Korean Peninsula it inhabits
mountain forests and subalpine meadows at altitudes between 1100 and 2500 m a.s.l., where
it builds nests in rotten wood and in soil, often under stones; on peat bogs it nests in moss
mounds. A large series of > 80 nests were taken between 1999 and 2000 in northern Mon-
golia, at about 1000 m a.s.l., by the Polish entomologist Micha! Woyciechowski, who de-
posited representative nest series in the ELMES collection. About 80% of these series lived
in mountain forest biotopes and ecotones between this forest and the herb-meadow biotopes
found adjacent to rivers, and between forest and dry-steppe biotope at the other extreme (for
description of these biotopes see Muehlenberg et al. 2000). In the first two habitats two
thirds of nests were in rotten wood (usually tree stumps), and the remainder split between
grass tussocks, soil and under stones; in the latter drier steppe-like ecotone only 15% of
colonies nested in rotten wood, the majority nesting in soil or under stones. The remaining
20% of colonies were evenly split between the herb-meadow biotopes and riparian-wood-
land biotopes where they preferred to nest in rotten stumps or moss and grass tussocks. All
these data show that M. angulinodis is principally a forest species that nests in rotten wood.
Mating swarms were observed on a hilltop (c. 1050 m) on 10 th and 12 th August 1999, these
swarms contained a mixture of M. angulinodis and M. forcipata and a few M. divergens.

Myrmica arisana Wheeler W. M., 1930

(Figs 25-27)

Myrmica rugosa var. arisana Wheeler W. M., 1930a: 95, w, Taiwan.

Myrmica rugosa subsp. arisana: Weber 1947: 462; Chapman and Capco 1951: 128; Bolton 1995: 277.
Myrmica arisana: Elmes and Radchenko 1998: 221.
Myrmica kurokii st. tipuna Santschi, 1937: 367, w, Taiwan. Synonymy by Elmes and Radchenko 1998:
221; confirmed here.
Myrmica kurokii subsp. tipuna: Weber 1947: 470; Chapman and Capco 1951: 127; Bolton 1995: 284.

Type localities and type specimens. M. arisana: "Five workers from Arisan". M. kotokui tipuna:
"Formose: 4 w (K. Sato), (n° 727)".
Material examined. Lectotype of M. arisana, w (bottom specimen on the pin with 2 w, designated
by Elmes and Radchenko 1998), "Arisan, Formosa, 4.24.1928, No 48, leg. R. Takahashii", "Typus",
"Wm. M. Wheeler", "MCZ Cotype 20557", "Myrmica rugosa var. Arisana Wheeler - Syntypes" (HAR-
VARD); paralectotype, upper w on the pin with lectotype (HARVARD); lectotype of M. kurokii st.
tipuna Santschi, w (bottom specimen on the pin with 4 w, designated by Elmes and Radchenko 1998),
"Formosa, K. Sato", "Formosa K. Sato 727", "Typus", "Sammlung der Dr. F. Santschi, Kairouan"
(BASLE); paralectotypes, 3 w on the pin with lectotype (BASLE); non-type material: > 300 w, 10 q,
65 m from Taiwan.
Distribution (Map 10). Taiwan.
Etymology. M. arisana: named after the Arisan region of Taiwan where the type speci-
mens were collected. M. tipuna: named for Tipun, an alternative name of Taiwan for the lan-
guage spoken by the Pyuma aboriginal peoples of Taiwan, who live in the southeast of the
Island.

92
 Notes. M. arisana female castes have frontal carinae that curve outwards and merge
with the rugae, which surround the,antennal sockets, and its males have long antennal
scape. On these characters it clearly belongs to the rubra species group and is not related
to M. rugosa. Workers and queens differ from other species of this group by the distinctly
prominent anterior clypeal margin; additionally, males well differ by the much coarser sculp-
ture of the head dorsum, especially by the rugulose clypeus.
The type specimens of st. tipuna slightly differ from those of M. arisana by their smaller
size, slightly more truncated petiolar dorsum and feebly sculptured alitrunk, petiole and
postpetiole. Study of additional material from Taiwan (see above) confirmed our opinion
that these differences represent infraspecific variation and are not sufficient for a separation
as species.
Since queens and males were unknown, we described them below (based on 10 queens
and 28 males, collected by G. Elmes in Taiwan, Hsien in 2002).
First description of queens: resembles workers by the shape of the head and scape, but
differ by having relatively shorter propodeal spines and longitudinally rugose scutum and
scutellum, without reticulation.
First description of males.
Head slightly longer than broad, with widely rounded occipital corners, lateral and oc-
cipital margins convex, anterior clypeal margin narrowly rounded and prominent, without
a medial notch. Scape long and very weakly curved at the base, antennae 13-segmented;
second funicular segment about 1.5 times longer than the third one. Alitrunk relatively long,
scutum slightly convex, scutellum does not project dorsally above scutum when seen in
profile. Propodeum with short blunt denticles. Petiole relatively short, with almost straight
and steep anterior surface and widely rounded node dorsum (seen in profile) ; postpetiole
higher than long, with convex dorsum.
Head dorsum (including clypeus) with dense but not coarse longitudinal rugulae, sur-
face between rugae densely though very finely punctated, but appears shiny. Scutum smooth
or at most finely longitudinally striated posteriorly, central part of scutellum smooth and

Map 10. Distribution of M. arisana.

93
shiny, lateral parts with rugosity. Sides of pronotum very finely longitudinally striated and
punctated, and mesopleura smooth, sides of propodeum rugose. Petiole and postpetiole
smooth and shiny.
Head margins and mandibles with numerous, relatively long suberect hairs. Alitrunk,
waist and gaster with numerous fine short hairs. Scape, tibiae and tarsi with quite dense
subdecumbent to suberect pilosity. Body colour blackish-brown, appendages brownish.
Ecology. Wheeler gave no ecological details when he described this species as a variety
of M. rugosa. We found a large population of M. arisana living under stones on a south-
facing bank by a roadside at 1700 m on Yungfen, Nantou (Taiwan). The stones were close
together and almost all housed part of the population. The impression was the stones were
connected and that the population comprised several large polycalic nests reminiscent of
how M. rubra lives in some European habitats. However, no mature queens were caught
(as would be expected if it was M. rubra) indicating that M. arisana is probably not an
especially polygynous species. The day was hot and the workers were very aggressive and
stung freely. In Chiayi near the Alishan Recreational Park (2000—2500 m a.s.l.), a few nests
were taken under stones and in tree stumps at the forest edge and in glades in the forest,
these were not particularly large and reminded us more of typical M. ruginodis nests.
Sexuais were observed leaving the nest (presumably to join a mating swarm - not seen) on
11 th August 2002.

Myrmica arnoldii Dlussky, 1963

(Figs 28-30)

Myrmica arnoldii Dlussky, 1963: 191, w, Russia; Pisarski 1969a: 227; Dlussky and Pisarski 1970: 85;
Dmitrienko, Petrenko 1976: 20; Dmitrienko 1979: 95; Pisarski, Krzysztofiak 1981: 155; Francoeur
1981: 758; Radchenko 1994b: 44, 1994c: 134, 1994e: 76, 1994g: 105; Bolton 1995: 277; Radchenko
and Elmes 2003a: 222; Pfeiffer et al. 2007: 4.
Myrmica (subgen. Dodecamyrmica) arnoldii: Arnoldi 1968b: 1803, q, m.

Type localities and type specimens. "Голотип: один рабочий, 12 IX 1957, гнездо № 302, Читинская
обл, Сретенский район, Дунасвскос лесничество, Балаганная падь (Длусский). Паратипы: 40 рабочих из
того же гнезда, что и голотип; 10 рабочих, собранных 7 IX 1957, гнездо № 275, Читинская обл., Сретенский
район, Дунасвскос лесничество, падь Кулинда (Длусский) [Holotype: one worker, 12.ix. 1957, nest No.
302, Chitinskaya Prov., Sretensky Distr., Dunaevskoe forestry department, Balagannaya pad', leg.
Dlussky. Paratypes: 40 w from the nest of holotype; 10 w, collected 7.ÌX.1957, nest No. 275, Chitinskaya
Prov., Sretensky Distr., Dunaevskoe forestry department, pad' Kulinda, leg. Dlussky].
Material examined. Holotype, w, "Читинская обл., Сретенский р-н, Усть-Дунасвскос, 12.9.1957,
Длусский" [Chitinskaya Prov., Sretensky Distr., Ust'-Dunaevskoe, 12.ix.1957, leg. Dlussky] (PE­
TERSBURG); paratypes: 8 w from the nest of holotype; 6 workers, same locality and collector, but
date 7.ix. 1957; 1 w, "302", "Myrmica arnoldii Dlussky Длусский опр.", "Усть-Дунасво, Читин. обл.,
Длусский, 12.ix.957" [Ust'-Dunaevo, Chitinskaya Prov., Dlussky, 12.ix.l957" (MOSCOW); non-type
material: > 100 w, about 50 q and m from different regions of South Siberia and Mongolia.
Distribution (Map 11). South Siberia from Tuva to eastern Transbaikalia, Mongolia.
Etymology. M. arnoldii: this species was dedicated to the famous Russian myrmecolo-
gist K. V. Arnoldi (see biography in Chapter 1.6).
Notes. This unusual species has a rather uncertain taxonomic position: its strange com­
bination of morphological features (see below) led us to erect the monotypic arnoldii species

94
group. On the other hand, a recent molecular analysis locates it in the incompleta species
group of North-American Myrmica (Jansen et al., 2009, 2010). Although this result was
quite unexpected it seems plausible because M. arnoldii shares several morphological fea-
tures with the species of incompleta -group, e.g. shape of scape and frontal lobes in the fe-
male castes, and short scape of males. M. arnoldii well differs from almost all other
free-living Palaearctic Myrmica species by several features, many of which are considered
as the "socially-parasitic syndrome": males have 12-segmented antennae (this feature led
Arnoldi 1968b erect subgenus Dodecamyrmica); the petiole and postpetiole of all castes
have ventral lobes, which are most developed in queens; the spurs on tibiae of mid- and
hind legs are often reduced (more frequently in female castes); they are small, queens are
little larger than workers (AL of workers, queens and males < 1.5, 1.8 and 1.6 mm, re-
spectively) (see also Radchenko and Elmes 2003a).
Ecology. Despite its "socially-parasitic" features, M. arnoldii has only been found living
in independent colonies. Dlussky (1963) reported that in Siberia most colonies lived in
larch forests with some being found in steppe-like habitats, nests were usually built in de-
caying wood and rarely in the soil. This accords with the more recent samples from Mon-
golia taken by M. Woyciechowski (see notes for M. angulinodis above): of > 20 colonies
almost all were found on northern and western slopes in the mountain forest biotopes at
about 1000 m a.s.l., nests were usually found in rotten tree stumps, but sometimes in the
soil or moss, they were quite populous and usually highly polygynous. Dmitrienko (1979)
reported that in Buryatia (Russia, just north of Mongolia) M. arnoldii was especially abun-
dant in forest clearings in larch and pine forests, where colonies exceeded 50% of all
ant-nests.
We suggested that M. arnoldii might be a temporary social parasite, or represent a hy-
pothetical step in the evolution of social parasites (Radchenko and Elmes 2003a). In any
case, this interesting species would make an ideal subject for further study in the context
of the evolution of social parasitism.

Map 11. Distribution of M. arnoldii.

95
 Myrmica bactriana Ruzsky, 1915, stat. rev.
(Figs 31, 32)

Myrmica smythiesi var. bactriana Ruzsky, 1915: 438, w, m, Tibet; Weber 1947: 457.
Myrmica bactriana: Collingwood 1970: 373.
Myrmica smythiesi subsp. bactriana: Bolton 1995: 277.
Myrmica tibetana var. furva Ruzsky, 1915: 441, w, Tibet; Menozzi 1939: 293; Weber 1947: 466.
Synonymy by Radchenko 1994b: 41; Bolton 1995: 279 (as synonym of M. tibetana); synonym of
M. bactriana: syn. nov.

Type localities and type specimens. M. smythiesi var. bactriana: "окр. уроч. Дариндо, дол. р. Дза-
чю, Кам, верховья р. Голубой (Янт-зцы), 8.VI11.1900 (mm, ww), (Козлов)" [Tibet, vicinity of Darindo,
valley of riv. Dza-chju, upper reaches of riv. Yangtze, 8.viii. 1900 (mm, ww), leg. Kozlov]. M. tibetana
var. furva: "1) p. И-чю (пр. Дза-чю), Кам, басе. р. Голубой, кон. III.1901 (рабочие); 2) р. Дза-чю, Кам,
басе. р. Голубой, 12-12.5000', нач. V. 1901 (рабочие). (Козлов)" [1) riv. I-chju (tributary of riv. Dza-
chju), Kam, basin of riv. Yangtze, end of iii.1901 (ww); 2) riv. Dza-chju, Kam, basin of riv. Yangtze,
12-12.500', beginning of v.1901 (ww), leg. Kozlov].
Material examined. Lectotype of M. bactriana, w (designated here), "окр. ур. Дариндо, Кам, верх.
Голубой, Козлов, 1/3.VIII.O0" [Tibet, vicinity of Darindo, Kam, upper reaches of riv. Yangtze, leg. Ko­
zlov, 1/3.viii.1900], "Myrmica smythiesi For. var. bactriana Ruzsky", "M. smythiesi v. bactriana Lec-
totypus design. Radchenko, 2008" (PETERSBURG); paralectotypes: 14 w, 3 m with the same locality
label as the lectotype (PETERSBURG, MOSCOW); lectotype of M. tibetana var. furva, w (designated
here), "p. Дза-чю, Кам, Голубая, 12-12500', Козлов, нач. v.Ol" [riv. Dza-chju, Kam, Yangtze, 12-12500',
leg. Kozlov, begin, v.1901], "Myr. tibetana var. furva Ruzsky" (PETERSBURG); paralectotypes: 14 w,
with the same labeis as lectotype (PETERSBURG, MOSCOW); non-type material: 1 w, Mobaishin,
east Nan-Shan Mts., leg. Kozlov 6-ll.viii.1908 (MOSCOW); 1 w, Tibet, Lhasa, Chunsap, 90°58'E,
29°28'N, 20.V.1999, 4200 m, taken foraging on sand by fast flowing mountain stream - with Formica
spp., leg. L. Johannessen (ELMES); 9 w, China, E Qinghai Prov., Hcnan, S env., 3500 m, 34°41'N,
101°36'E, 20.vi.2005, leg. J. Kalab, meadows, grasslands (MILAN).
Distribution (Map 12). Tibet and adjacent regions of Qinghai Prov.
Etymology. M. bactriana: Bactria (or Bactriana) was an old province of Greater India
centred on modern Tajikistan that has lent its name to many taxa from that region. We are
not sure why Ruzsky chose the name bactriana for a Tibetan species unless he had in mind
the Bactrian camels that live wild in parts of Tibet. M. furva: from the Latin word furva =
dark or swarthy, presumably to indicate the body colour of the types.
Notes. Collingwood (1970) recorded M. bactriana from several localities in Nepal
(Ringmo-Junbesi, 2800 m; Yaral, Pangboche, 3900 m; Thate, 2900 m) and raised it to the
species level with no supporting comments. We have not seen this material and can
not comment on the identification of these samples. However, based on the original
type material in PETERSBURG and MOSCOW we are sure that M. tibetana (see notes
on that species) is quite distinct from the previously enigmatic taxon M. bactriana.
On the other hand, we think that M. tibetana var. furva and M. bactriana are synonyms.
Queens are unknown, but based on the characteristics of the males and workers we
place it in the smythiesii species group, as defined by Radchenko and Elmes (2001b).
M. bactriana is very similar to M. ruzskyana, differing only by its distinctly longer
scape (SI 2 > 0.93 vs. < 0.91) with more abundant and suberect hairs, and it is quite
possible this represents different populations of the same species. This question can
not be resolved until more material from Tibet and adjacent regions of China becomes
available.

96
 Ruzsky's original description of both var. bactriana and var. furva are incomplete and
do not include many diagnostic feature that are used in the modern taxonomy of the genus
Myrmica, therefore we redescribed the workers and males.
Redescription of workers.
Head distinctly longer than broad, with subparallel sides, straight occipital margin and
narrowly rounded occipital corners; anterior clypeal margin narrowly rounded, distinctly
prominent with no medial notch. Upper latero-ventral corners of head rounded, not pointed
(seen in profile). Mandibles with 6-7 teeth. Frontal carinae curved outwards to merge with
rugae that surround antennal sockets. Frons wide (FI > 0.41), frontal lobes not extended
(FLI < 1.15). Scape distinctly shorter than head width (SI2 < 0.91), weakly curved at the
base, without any trace of lobe or carina. Alitrunk quite short, metanotal groove distinct,
deep. Propodeal lobes rounded apically. Propodeal spines short (ESLI < 0.24, mean 0.19),
straight, thin, directed backward at an angle about 45°. Petiole with relatively short but dis-
tinct peduncle, its anterior surface slightly concave, dorsum of node rounded. Postpetiole
higher than long (PPIj < 0.87), its anterior and dorsal surfaces forming a regular arch.
Spurs on middle and hind tibiae well developed and pectinate.
Frons with quite dense, fine, slightly sinuous longitudinal rugosity, numbering ca. 20 be-
tween frontal carinae level with the eyes, occiput and temples with fine reticulation, surface
between rugae at most with superficial micropuntures, but appears shiny; clypeus with weak
longitudinal carina and very fine rugulae, surface shiny; frontal triangle smooth and shiny;
mandibles longitudinally rugose. Alitrunk with longitudinal, slightly sinuous but not coarse
rugae. Surfaces between rugae on alitrunk smooth and shiny. Petiole and postpetiole with
very fine longitudinal rugulosity, appearing shiny. Gaster smooth and shiny.
Head margins above the eyes with subdecumbent hairs, genae with a few long hairs;
alitrunk dorsum with longer hairs; petiole with 6-8 long hairs. Scape and tibiae with sub-
decumbent to decumbent pilosity. Body colour reddish to reddish-brown, appendages some-
what lighter.

Map 12. Distribution of M. bactriana.

97
 Redescription of males.
Head distinctly longer than broad (CI 1.16-1.20), with rounded occipital corners, sides
and occipital margin feebly convex, anterior clypeal margin narrowly rounded and promi­
nent, with no medial notch. Scape long (SI, 0.85-0.86, SI 2 1.00-1.02) and very weakly
curved at the base, antennae 13-segmented; second funicular segment subequal in length to
the third one. Alitrunk relatively long (AI 1.64-1.66), scutum slightly convex, scutellum
does not project dorsally above scutum when seen in profile. Propodeum without denticles,
at most with small tubercles. Petiole relatively short, with slightly concave and steep ante­
rior surface and widely rounded node dorsum (seen in profile) ; postpetiole higher than long,
with convex dorsum.
Head dorsum (excluding frontal triangle) densely punctated, frons also finely longitu­
dinally rugulose, surface of clypeus with reduced sculpture, appearing shiny. Scutum lon­
gitudinally rugulose, scutellum and posterior part of scutum quite coarsely punctated. Sides
of pronotum and mesopleura finely longitudinally striated and punctated, sides of
propodeum coarsely rugose and finely punctated. Petiole and postpetiole very finely super­
ficially punctated but appear shiny.
Head margins and mandibles with numerous, relatively long, curved suberect hairs.
Alitrunk, waist and gaster with sparser hairs. Scape, tibiae and tarsi with quite dense de­
cumbent to subdecumbent pilosity. Body colour dark brown to blackish-brown, appendages
reddish-brown.
Ecology. Little is known, except that this species was found at high altitudes, on grass­
lands between 3500 and 3800 m a.s.l. Almost certainly, it nests in the soil.

Myrmica bakurianica Arnoldi, 1970

(Figs 33-35)

Myrmica pilosiscapus subsp. bakurianica Arnoldi, 1970: 1837, 1842, w, q, m, Georgia.

Myrmica bakurianica: Seifert 1988: 22; Bolton 1995: 277.
Junior synonym of M.. bessarabica: Radchenko 1994d: 78; not confirmed, see Notes below.

Type localities and type specimens. "Найдена на Кавказе в окрестностях Бакуриани на высотах
1500-1700 м; описана по 5 W, среди них голотип - 4 сентября 1927 г. (Арнольди), и по 4 mm (андротип,
тоже) и 2 qq" [Found in Caucasus in the vicinity of Bakuriani at altitudes 1500-1700 m; described
based on 5 w, among them holotype - 4th September 1927 (leg. Arnoldi), and on 4 m (androtype, the
same) and 2 q].
Material examined. Neotype, w (designated by Radchenko 1994d), "Borzhomi, 1700 m,
20.VIII.1927, No 5012, K. Arnoldi", "Myrmica pilosiscapus Bondr. K. Arnoldi det", "Myrmica baku­
rianica Arnoldi Det. Seifert 1988" (MOSCOW); non-type material: 5 w, 3 q and 3 m with the same la­
bels (MOSCOW) (see also Notes below).
Distribution (Map 13). Georgia.
Etymology. M. bakurianica: from the name Bakuriani with the adjective suffix for nouns
ica (from the Greek то) = belonging to, from, to indicate that it is a resident of vicinity of
Bakuriani (Georgia), where the types were found.
Notes. Arnoldi (1970: 1837) gave very brief descriptionofM . pilosiscapus bakurianica
in the Key, translated from Russian it reads: "Frontal carinae less curved [i.e. than in M. slo­
vaca and M. salina], i.f. = 2.40-2.65 [i.e. our FI 0.377-0.416]. Body colour dark, reddish-

98
brown, head and gaster black (w). Hairs light, very numerous, especially on gaster. Sculp­
ture coarse, especially on alitrunk and waist. Lateral parts of frons with coarse irregular
longitudinal rugae, area near eyes with reticulation. Metanotal groove very sharp, propodeal
spines long. Males have numerous long hairs on scape and tibiae". Additionally, one un­
derstands from the Key that the petiole of workers does not have a distinct dorsal plate.
We can not locate the original type material in any collection examined by us, and we
suggest that it is most probably lost. However, б w, 3 q, 3 m, originally determined by
Arnoldi as M. pilosiscapus and determined by Seifert (1988) as M. bakurianica, are pre­
served in MOSCOW (see Material examined, above). Although these specimens can not be­
long to the type series because the date and locality data do not correspond with those given
by Arnoldi in his original description of M. pilosiscapus bakurianica, a worker from this se­
ries was designated as a neotype of M bakurianica by Radchenko (1994d). Radchenko
(loc. cit.) then synonymised subsp. bakurianica with M. bessarabica (see also Notes to
M. specioides, below, and to M. bessarabica in Chapter 3.5).
We now believe that M. bakurianica is a good species. By the shape of their frontal cari­
nae, frontal lobes and petiole workers and queens of this species are similar to those of
M. specioides, but by the shape of scape lobe and by the length of propodeal spines they re­
semble M. scabrinodis. Similarly, males of M. bakurianica share features of both species;
thus, by the length and pilosity of the scape they are similar to M. specioides, but have much
longer standing hairs on the tibiae and tarsi of the middle and hind legs, similar to but some­
what shorter than those of the males of M. scabrinodis.
Since Arnoldi's original description is very short and incomplete, we redescribe all castes
of M . bakurianica.
Redescription of workers.
Head longer than broad, with rounded occipital corners, sides and occipital margin
weakly convex. Anterior clypeal margin broadly rounded, not prominent, without medial
notch. Frontal carinae not strongly curved, frons relatively wide and frontal lobes
moderately extended (similar to that of M. specioides). Scape strongly angled at its base,
with a narrow, but distinct horizontal lobe.
Alitrunk with convex promesonotal dorsum; promesonotal suture indistinct from above;
metanotal groove distinct and deep. Propodeal spines relatively long, acute and straight, di­
vergent (seen from above), projecting backwards at an angle ca. 45°. In profile, anterior
surface of petiole concave, its dorsal surface without plate, gradually sloping posteriorly.
Postpetiole shorter than high, with feebly convex dorsum. Spurs on middle and hind tibiae
well developed and pectinate.
Only frons with sinuous, quite coarse longitudinal rugae, remainder parts of head dor­
sum with reticulation. Antennal sockets surrounded by fine concentric rugae. Clypeus with
longitudinal rugae. Alitrunk with longitudinal, slightly sinuous rugae. Petiolar node with
short sinuous rugae and reticulation, postpetiole with longitudinally-concentric rugae. Sur­
face between rugae on the body at most with fine microsculpture, appears shiny.
Head margins with abundant, relatively long, but subdecumbent hairs. Scape with very
abundant suberect hairs. Alitrunk dorsum with numerous straight erect to suberect hairs,
petiolar node with 7-9 straight long hairs and a few shorter ones, postpetiole with abundant
hairs. Gaster with abundant hairs. Generally; hairs on the body quite coarse and whitish,
coarser and more abundant than in related species (e.g. M. specioides, M. scabrinodis,
M. turcica etc).
Body colour reddish-brown (not so dark, as was mentioned by Arnoldi).

99
 Redescription of queens.
Relatively small compared to their workers, about the same size or even smaller than
some (the mean HL and HW is less than that of workers). Nevertheless, they resemble the
workers by the shape of the head, frontal carinae and lobes, scape, petiole and postpetiole,
as well as by the general appearance of body sculpture and pilosity.
Redescription of males.
Head sides and occipital margin convex, occipital corners gradually rounded, head-
length equal to width or very slightly longer; anterior clypeal margin widely rounded, not
prominent and with no medial notch. Antennal scape short (similar to that of M. specio-
ides), antennae 13-jointed, antennal club 4-segmented; second funicular segment only
slightly longer than the third one.
Alitrunk relatively long, scutum slightly convex, scutellum does not project dorsally above
scutum when seen in profile. Propodeum with distinct but short, blunt denticles. In profile,
petiole relatively long and low, with widely rounded node dorsum, its anterior surface slightly
concave, not steep; postpetiole distinctly higher than long, with weakly rounded dorsum.
Head dorsum (except for clypeus) densely punctated, fine striation presents only near
ocelli. Central part of scutum behind Mayrian furrows with longitudinal striation, remain-
der of surface smooth and shiny; scutellum with fine longitudinally-concentric rugulosity.
Propleura smooth, mesopleura with fine striation, sides of propodeum rugulose. Petiolar
node partly with fine striation, postpetiole smooth. Whole surface of alitrunk appears shiny.
Head margins and mandibles with long, curved erect hairs. Alitrunk, petiole, postpeti-
ole and gaster with somewhat shorter erect to suberect hairs. Tibiae and tarsi on both an-
terior and posterior surfaces with long suberect to erect hairs; the longest hairs on tibiae
distinctly longer than their maximal width, those on the basitarsus more than twice longer
than maximal tarsal width. Scape and 8 following funicular segments.with long hairs (how-
ever, hairs on the legs and scape shorter than those in M. scabrinodis or M. tulinae).
Body colour brownish-black, appendages somewhat lighter.
Ecologyis unknown.

Map 13. Distribution of M. bakurianica.

100
 Myrmica bergi Ruzsky, 1902
(Figs 36-38)

Myrmica bergi Ruzsky, 1902b: 473, w, Kazakhstan (also described as a new by Ruzsky, 1902c: 12);
Ï903: 314, 1905: 675, w, q, m; Emery 1908a: 166, 1921: 37; Kuznetsov-ugamsky 1927: 191, 1929:
44; Arnoldi 1934: 160; Weber 1947: 472; Sadil 1952: 244; Arnoldi 1970: 1839, 1976: 556; Tarbin-
sky 1976: 45; Seifert 1988: 11; Radchenko 1994b: 42, 1994c: 143, 1994d: 76; Bolton 1995: 277;
Radchenko et al. 2002: 413, nec Sadil 1952: 244.
Myrmica rubra L. r. bergi: Forel 1904a: 374.
Myrmica rugosa var. kirgisica Ruzsky, 1903: 314, w, Russia; 1905: 659; Emery 1908a: 168, 1921: 40;
Weber 1947: 464. Synonymy by Seifert 1988: 11; Radchenko 1994d: 76; Radchenko et al. 2002:
413; confirmed here.
Myrmica rubra var. kirgisica: Forel 1907: 18.
Myrmica bergi subsp. kirgisica: Arnoldi 1970: 1839.
Myrmica bergi var. barchanica Ruzsky, 1905, 678, w, Russia; Emery 1908a: 173, 1921: 37; Weber
1947: 473. Synonymy by Arnoldi 1970: 1839; Seifert 1988: 11; Radchenko 1994d: 76; Radchenko
et al. 2002: 413; confirmed here.
Myrmica bergi var. fortior Crawley, 1920: 163, w, Iran (junior primary homonym of Myrmica smythiesi
var. fortior Forel 1904b: 22).
M. bergi subsp: persiana Weber 1947: 474 (replacement name for Myrmica bergi var. fortior Crawley,
1920: 163); Bolton 1995: 281. Synonymy by Radchenko 1994d: 76; Radchenko et al. 2002: 413;
confirmed here.
Myrmica kamyschiensis Arnoldi, 1934: 159, w, q, Ukraine; synonymy by Seifert 1988: 11; Radchenko
1994d: 76; Radchenko et al. 2002: 413; confirmed here.
Myrmica bergi subsp. kamyschiensis: Karawajew, 1936: 272, w, m; Weber 1947: 474; Arnoldi 1970:
1838.

Type localities and type specimens. M. bergi: "окр. Аральского моря, Тас-Булак на западн. берегу,
5.VII.1900 (Берг); устье Сыр-Дарьи, Раим, 7-8.V.1901 (он же); Ак-Джулпас на северо-вост. бср., 26.
V.01 (он же); окр. Казалинска, берег Сыр-Дарьи, 13.VI.1901 (он же)" [vicinity of Aral Sea, Tas-Bulak,
W shore, 5.VÜ.1900, leg. Berg; mouth of the riv. Syr-Darya, Raim, 7-8.V-1901, same collector; Ak-
Dzhulpas, NE shore, 26.V.1901, same collector; vicinity of Kazalinsk, bank of the riv. Syr-Darya,
13.vi.1901, same collector] (Ruzsky 1902a); "Fundorte: Tas-Bulak an Westufer des Aralsees; Mün-
dung des Syr-Darja; Ack-Dshulpas am nordöstlischen Ufer des Aralsees (H. L. Berg, 1900-1901)
(Ruzsky 1902b). M. rugosa var. kirgisica: "Найдена около Астрахани. Кроме того, еще один экземпляр -
в песках близ Ханской Ставки" [Was found near Astrakhan'. Additionally, one more specimen - on
sandy areas near Khanskaya Stavka]. M. bergi var. barchanica: "пески около Ханской Ставки
(Астраханская губ.)" [sandy areas near Khanskaya Stavka (Astrakhan' Prov.)]. M. bergi var. fortior:
"Enzeli, N.W. Persia, 1919 (Buxton)". M. kamyschiensis: "Krim, Kertsch-Meeres-enge, sandiger Salz­
strand des Brackwasserbeckens Kamysch-Burun am Meer. Nistet im feinen Muschelsand im Juncus-
bestand, IV.1923 (Nr. 888 meiner Sammlung); auch am südlichen Rande der Ukrajnischen Steppen,
Salzmoor bei Sivaschufer (Medvedev, 1931)".
Material examined. Lectotype of M. bergi, w (middle specimens on the pin with 3 w, designated by
Arnoldi 1970; see Notes below), "986, Aral see, L. S. Berg, 1901", "Lectotype" (MOSCOW); paralec-
totypes: 2 w on the ; pin with lectotype; 1 w, "Kazalinsk, 1901, Berg" (MOSCOW); 1 w, "Аральское
море, Л. С. Берг, 1901" [Aral Sea, L. S. Berg, 1901], "5630 Coll. Karawajewi", "M. (Myrmica) bergi
Ruzs. typus", "Paratypus Myrmica bergi Ruzs." [KIEV]; 2 w (bottom specimen without head), "Myrm.
Bergi m, w" (written by Ruzsky's own hand), "Myrmica bergi Ruzski [Sicl] Lago Arai" (GENOA); lec­
totype of M. bergi var. barchanica, w (designated by Radchenko et al. 2002), "Myrmica bergi var. bar­
chanica Ruszs, Ханская Ставка, рынь-псски, 6.VI. 1902, M. Рузский" [Khanskaya Stavka, ryn'-peski,
6.VÌ.1902, M. Ruzsky] (written by Ruzsky's own hand) (MOSCOW); paralectotype: w, "Myrmica bergi

Ю1
var. barchanica Ruszs., Ханская Ставка, пески, 5.VI.1902, M. Рузский" [Khanskaya Stavka, peski,
5.VÌ.1902, M. Ruzsky] (written by Ruzsky's own hand) (PETERSBURG); lectotype of M. bergi var.
fortior, w (designated by Radchenko et al. 2002), "NW Persia, 1919, P. A. Buxton", "Myrmica bergi var.
fortior Crawley" (OXFORD); paralectolypes: 10 w with same labels as lectotype (OXFORD,
LONDON, MOSCOW); syntypes of M. kamyschiensis (designated by Radchenko et al. 2002; see Notes
below): 1 w, "A 888", "M. bergi kamyschiensis", "Holotype" (all written by Arnoldi's own hand); 2 w,
"A 888", "M. bergi kamyschiensis", "Paratype" (all written by Arnoldi's own hand) (MOSCOW); 1 w,
"No. 888, Керчь, Камыш-Бурун, 21.IV.23, К. Arnoldi" [Kerch', Kamysh-Burun, 21.iv.23, К. Arnoldi",
"5629 Coll. Karawajewi", "Myrmica (Myrmica) kamyschiensis Arnoldi typus", "Paratypus Myrmica
kamyschiensis K. Arn." (KIEV); non-type material: > 200 w, several tens of q and m from the whole
area of the species.
Distribution (Map 14). Steppe Zone from the delta of Danube riv. to Altai Mts.,
Transcaucasus, NE Iran, plains and mountains of Middle Asia.
Etymology. M. bergi: this species was named for its collector, Prof. Lev Semenovich
Berg, who was a distinguished Russian natural scientist, most famous for his work on ma­
rine biology and the theory of nomogenesis (the initials H. L. Berg in Ruzsky 1902b
probably means "Herr L. Berg"). M. kirgisica: from the name "Kirgis" with the adjective suf­
fix for nouns ica (from the Greek IKO) = belonging to, from, which indicates that it is
a resident of the "Kirgis Steppe". At the beginning of 20th century the name Kirgis (spelled
also as Kirgiz) signified the vast area of dry steppe and semi-deserts that extended from the
lower Volga river through modern Kazakhstan, to Dzhungaria near the Chinese border. At
that time, the modern Kazakh nation were called Kirgiz (to avoid confusion with the
Russian and Ukrainian Kazaks), while the modern Kirgiz peoples (living in Kyrgyz­
stan) were called the Kara-Kirgiz (Black Kirgiz). M. barchanica: from the name barchan
(winged-shaped sand dunes) with the adjective suffix for nouns ica (from the Greek IKO)
= belonging to, from, which indicates that it is a resident of barchans. The types were found
in sandy areas near Astrakhan', where there are many barchans. M. persiana: named for
Persia (modern Iran), where the types were found. M. kamyschiensis: from the name of
the type locality, Kamysh-Burun, with the Latin suffix ensis = place of origin.
Notes. We made a full appraisal of M. bergi and concluded that all of its subspecific
forms are synonyms of M. bergi, although several were sufficiently distinct to be considered
as geographic races (Radchenko et al. 2002). M. bergi is a member of the scabrinodis species
group and is most similar to M. gallienii, differing from the latter by the shorter propodeal
spines, lower petiole, darker colour, etc.; males differ by relatively longer scape.
Arnoldi (1970: 1839) wrote concerning M. bergi: "As lectotypes (Sici) I designate
worker: Tas-Bulak near Aral Sea (L. S. Berg), male - Frunze, Kirgizia (Arnoldi)". Re­
garding the male Arnoldi was in error because he added to the type series a specimen col­
lected by himself many years after the first description of the species; furthermore we could
not find specimens with this data either in MOSCOW or in PETERSBURG. However, we
found three workers from the Aral Sea, on a pin labelled by Arnoldi as "Lectotype", and for­
mally designated the middle specimen as the lectotype of M. bergi (Radchenko et al. 2002).
The types of var. kirgisica most probably are lost. We found in MOSCOW a pin with 3
workers from "Астраханский заповедник, 10.VI.60, поды, Писарев" ["Astrakhan' Natu­
ral Reserve, 10.vi.60, pody, Pisarev"], labelled by Arnoldi as "Lectotype, M. bergi kirgisica".
Again Arnoldi was in error, considering as "lectotypes" specimens collected 58 years after
the first description, at best one could only be considered as a neotype. Despite labelling
these specimens, Arnoldi (1970) did not formally designate a lectotype or neotype for var.
kirgisica.

Ю2
 We consider Arnoldi's type specimens of M. kamyschiensis as syntypes (Radchenko et
al. 2002), because he did not designate any type specimens in his original description of
this species (Arnoldi 1934). We are sure that Arnoldi added his type labels much later (see
above), probably when he revised Myrmica in the 1960s, because he followed his later
opinion on the taxonomic status of this form, labelling the specimens "M. bergi kamyschien-
sis" rather than "M. kamyschiensis" as in his original description.
Material from Czechoslovakia, identified by Sadil (1952) as M. bergi, in fact belongs to
M. gallienii (Arnoldi 1970).
Arnoldi's (1948) M. bergi iranica is nomen nudum. However, we investigated workers
from Zuvand (Talysh, Azerbaijan) (MOSCOW), labelled by Arnoldi as "M. bergi iranica"
or "M. bergi persiana"; all of them belong to M. bergi.
Ecology. Although M. bergi is distributed in semi-arid and even arid regions, within
these it dwells exclusively in intrazonal, damp and almost always salted habitats, where it
can be very abundant locally. Arnoldi (1934: 161) states that it is "Characteristic for salty
bogs". The populations of M. bergi from the steppe zone of southern Ukraine have been
most studied (Bondar et al. 1998; Bondar 1999; Bondar, Rusina 2003; Stukalyuk, Rad-
chenko 2008, and personal observations). Here it nests in the reed, sedge and grass asso-
ciations that border the sea shore (salt marshes and lagoon shores) and surround the
numerous salted lakes of differing sizes. Around salted lakes workers can be seen foraging
over the caustic encrusted, white, salty deposits even in full sunshine. Nests are built shal-
lowly in the soil but almost always have a large mound of soil above them in which the ma-
jority of ants live. Usually the mound is constructed around plant stalks, especially sedges,
and in times of flooding the entire colony migrates up the stems where it can construct tem-
porary nest high above the water level, covered with soil and pieces of vegetation (Bondar
et al. 1998), and the foraging workers actively swim, sometimes for several tens of meters.
The other studies cited above have shown that colonies can be large and polygynous,
containing several thousand workers, and in optimal conditions they can be polycalic

Map 14. Distribution of M. bergi.

103
comprising several tens of nests. Possibly this may be a local adaptation to high related-
ness, it being probably that populations living around isolated lakes are quite inbred. We
have seen small colonies nesting more typically in the soil (for scabrinodis-group.species),
living in competition with other ant species in short moist, grazed grassland on the raised
banks of the river Dnepr where flooding is less likely.
Tarbinsky (1976) studied M. bergi living in Kirgizia; he wrote (loc. cit., p. 46) that it
"Lives up to altitude 1600 m, along river and stream banks, near Salix and Hippophaë
shrubs, or near Phragmites, in semi-shaded places. It nests in sandy soil, sometimes with
small mounds, but usually without them. In the morning and evening ants forage on open
sand but during the day, in shade under shrubs or Phragmites". In this respect it is quite
similar to the Ukrainian populations. However, Tarbinsky (loc. cit.) also said that colonies
are "quite large compared to other Myrmica species" (which agrees with our observations)
"and monogynous" which is atypical for south Ukrainian populations. Ruzsky (1905)
recorded that var. barchanica nested in sandy soils with rich vegetation, mainly Populus and
Salix.

Myrmica bibikoffi Kutter, 1963

(Figs 39-41)

Myrmica bibikoffi Kutter, 1963: 130, w, q, m, Switzerland; 1973: 255, 1977: 65; Seifert 1988: 41;
Bolton 1988: 4, 1995: 277; Radchenko and Elmes 2003a: 227, 2004: 232; Seifert 2007: 200;
Garcia et al. 2008.

Type localities and type specimens. "Material: 5 ww, lq, 4 mm. Holotypus der w in meiner Samm-
lung, alle Geschlechtstiere, sowie 4 Paratypen der w in der Summlung Bibikoff im Britischen Naturhi-
storischen Museum, London. Als Fundort wird angegeben: Vaulion Kanton Waadt, 5 August 1949,
Sammler M. Bibikoff".
Material examined. Paratypes, 4 w, 1 q, 3 m, "Vaulion, Canton Vaud, 5.viii. 1949, M. Bibikoff"
(LONDON); non-type material: 1 w, 1 q, 2 m, "Germany, Schwarbische Alb, Erpfingen, limestone
grassland, sunny hill slope, beneath stone, EU 489, 02.viii.1998, leg. K. Rosciszewski" (ELMES).
Distribution (Map 15). Until now, this rare species has been found only four times in
Switzerland, Germany and Spain.
Etymology. M. bibikoffi: named for the collector Mr. Michel Bibikoff, a Russian émigré
who studied entomology in Paris and was later famous for his work on training guide dogs.
Notes. This quite conspicuous species belongs to the sabuleti-complex of the scabri-
nodis-group (Radchenko and Elmes 2004). M. bibikoffi is characterized by strongly reduced
spurs on the hind and mid tibiae, a coarse reticulated sculpture on head and alitrunk, the
wide postpetiole of workers and queens, and a hairy body. It is similar to M. hirsuta and
shares the same host, M. sabuleti. Queens are larger than those of M. hirsuta and also dif-
fer by their body sculpture and narrower frons. Males have a relatively long scape, similar
to that of M. sabuleti.
Ecology. The original collector (Mr. M. Bibikoff) found all castes of this species living
in an independent nest, but the other samples were collected from nests of M. sabuleti and
its close relative M. spinosior. In these cases only workers of the host species were found
which suggests that M. bibikoffi might be at least a temporary social parasite, which in some
ecological situations can only survive as a permanent social parasite.

104
 Map 15. Distribution of M. bibikoffi.

Myrmica boltoni Radchenko et Elmes, 1998

(Fig. 42)

Myrmica boltoni Radchenko et Elmes, 1998: 18, w, Nepal; 2001b: 242.

Type localities and type specimens. Nepal, for details see material examined.
Material examined. Holotype, w, "Nepal, Dhorpantan, 3000 m, 20.V.1973, leg. T. Martens"
(BASLE); paratypes: 15 w with the same label (probably same nest) as holotype; 3 w, "Umg. Goropani,
w. Pokhara, Zentral Nepal, Sept.-Oct. 1971, leg. H. Franz"; 1 w, "Nepal, 2 mis. S.E. Sikha, 7800-
8000 fts., 21-22.V.1954, leg. J. Quinlan"; 1 w, "Nepal, 18 km NNE Baglung, 28°24'N, 83°42'E, 2540
ni."; 1 w, "Nepal-23, Prov. Kosi, Distr. Sankhuwasawa, Vallee d'Induwa Koa, 2000 m, 16.iv. 1984, leg.
Lobi and A. Smetana"; 1 w, "Nepal-140, Prov. Manang, Marsyandi, 2550 m, 14-17.iv.1980, leg. J. Mar-
tens and Ausobsky"; 2 w, "Nepal-161, Prov. Mustang, Lethe, 2450-2600 m, 30.iv.1980, leg. J..Martens
and Ausobsky"; 1 w, "Nepal-233, Prov. Gorkha, Chuing Khola, Meme Kharka, 3300-3400 m, leg. J.
Martens and W. Schwaller" (BASLE, LONDON, KIEV, ELMES, WARD, MARTENS, SCHULZ).
Distribution (Map 16). Nepal.
Etymology. M. boltoni: this species was dedicated to our friend and colleague, Dr. Barry
Bolton of the British Museum of Natural History, London, the most famous of modem ant
taxonomists.
Notes. M. boltoni belongs to the boltoni-complex of the ritae species group. It differs
from all members of the níae-complex by the much finer rugosity on its body, and by the dull,
dense and coarsely punctated surface of its head. From M. collingwoodi it differs by its
densely punctated surface of the head dorsum and waist; by the characteristics of its body
sculpture M. boltoni resembles M. martensi, but differs from the latter by the distinctly longer
and not-divergent propodeal spines, and by the relatively longer and lower petiole. Queens
and males are unknown.

105
 Ecology is almost unknown, but one worker was collected under rotten wood in
a Quercus-Rhododendron forest (P. S. Ward, pers. comm.). All samples were collected at
altitudes between 2000 and 3400 m a.s.l.

Map 16. Distribution of M. boltoni.

Myrmica brancuccii R a d c h e n k o , Elmes et Collingwood, 1999

(Fig. 43)

Myrmica brancuccii Radchenko, Elmes et Collingwood, 1999: 30, w, Nepal, Pakistan (in: Radchenko
and Elmes 1999a); Radchenko and Elmes 2001b: 260.

Type localities and type specimens. Nepal, Pakistan, for details see material examined.
Material examined. Holotype, w, "Nepal, Utrot, 13.V.83, leg. M. Brancucci" (LONDON); paraty-
pes: 5 w, with the same label as holotype; 14 w, "Nepal, Lawarai, 21.05.83, leg. M. Brancucci"; 1 w,
"Nepal, Lumie, vi.88, leg. Collingwood"; 9 w, "Pakistan, Chitral V., between Dir and Lavari Pass, 2400
m, ll.viii.1994, leg. S. Dacatra" (LONDON, BASLE, LEEDS, MILAN, KIEV, ELMES).
Distribution (Map 17). Nepal, NE Pakistan.
Etymology. M. brancuccii: this species was dedicated to the well-known coleoptero-
logist, Prof. Michael Brancucci, of the Natural History Museum Basle, who collected the
type material.
Notes. We can not place this species with certainty into any species group. M. bran-
cuccii is well discriminated from almost all Old World Myrmica by its striato-punctated
first gastral tergite, sharing this unusual feature only with M. sculptiventris from SW
China, but the latter species clearly differs from M. brancuccii by the transversai
rugosity on the mesonotal and propodeal dorsum. Workers are very variable in size, so
that this could be a relatively polymorphic species in comparison with other Myrmica
species. Queens and males are unknown.
Ecologyis unknown, except that one sample was collected at an altitude 2400 m a.s.l.

toó
 Map 17. Distribution of M. brancuccii.

Myrmica cachmiriensis Forel, 1904

(Figs 44-46)

Myrmica smythiesii r. cachmiriensis Forel, 1904b: 23, w, India; Ruzsky 1905: 661; Forel 1906: 87;
Emery 1921: 41; Menozzi 1939: 292, m; Weber 1947: 457; Chapman and Capco 1951: 130; Bolton
1995: 278, nec Eidmann 1941: 15, misidentification.
Myrmica cachmiriensis: Radchenko and Elmes 2001b: 251.
Myrmica smythiesi var. lutescens Menozzi, 1939: 292, w, q (first available use of Myrmica smythiesii
x. cachmiriensis var. lutescens Forel, 1904b: 23, w, India; Ruzsky 1905: 661; Forel 1906: 87; Emery
1921: 412; Weber 1947: 458; Chapman and Capco 1951: 130); Bolton 1995: 281. Synonymy by
Radchenko and Elmes 2001b: 251; confirmed here.

Type localities and type specimens. M. cachmiriensis: "Sind Valley, Cachemire (Wroughton), 2286
à 2438m." M. smythiesii r. cachmiriensis var. lutescens: "Cachemire (Smythies)".
Material examined. Lectotype of M. cachmiriensis, w (designated by Radchenko and Elmes 2001b),
"Kashmir, Sind Valley, 9500 ft (Wroughton)", "M. smythiesii v. kashmiriensis [Sic\] Forel, type"
(GENEVA); paralectotypes: 2 workers with same labels as lectotype (GENEVA); 1 w, "M. Smythiesii
For. var. kaschmiriensis [Sicì] w type Sind Valley, 7500' Kaschmir (Wroughton) Jh. 6" (GENOA);
lectotype of M. smythiesi var. lutescens, w (designated by Radchenko and Elmes 2001b), "Cachmire
(Smythies)", "M. smythiesii For. r. cachmiriensis For. v. lutescens For., type" (GENEVA); paralecto-
types: 2 w with same labels as lectotype (GENEVA); non-type material: about 50 w, 7 q, 3 m from
India and Pakistan.
Distribution (Map 18). India (Kashmir), NE Pakistan. A record for China (Eidmann
1941) almost certainly belongs to another species.
Etymology. M. cachmiriensis: named from the Himalayan region of Cachmire (alter-
native spelling of Kashmir) combined with the Latin suffix ensis = place of origin, to indi-
cate where the type material was collected. M. lutescens: from a combination of the Latin
words lute = yellow and the suffix escens = -ish, to describe its yellowish colour (see Notes
below).

107
 Map 18. Distribution of M. cachmiriensis.

Notes. This species belongs to the cachmiriensis-comp\ex of the rugosa species group. It most
resembles M. wardi, but differs from the latter by the much finer sculpture on the head dorsum.
The type specimens of var. lutescens differ from the types of M. cachmiriensis only by
the lighter yellow or somewhat orange-yellow colour of their bodies. However, among the
other non-type material studied, we found that the colour of specimens, even from the same
nest, varied from pale yellow to yellowish-red. Consequently, we consider var. lutescens to
be a junior synonym of M. cachmiriensis.
Menozzi (1939: 292) recorded the following additional localities forM. cachmiriensis:
Chokpiong, 2600 m; Hoto, 2900 m; Askole, 3100 m (valley Braldo); Smiuntsa, 3200 m;
Tsok, 3490 m (valley Punmah); Kagan, 2000 m (all are in northeast of Pakistan, Karako-
rum). Menozzi labelled a specimen from one of these locations (Askole, 3100 m, valley
Braldo) as a type of M. dicaporiaccoi, but we identified this specimen as M. wardi (see notes
to M. aimonissabaudiae). Therefore, we suspect that the material determined by Menozzi
as M. cachmiriensis {loc. cit.) will after detailed examination, be shown to contain several
species from the cachmiriensis -complex. Furthermore, Eidmann (1942: 15) recorded
M. cachmiriensis from West China and Tibet (alt. 1000-3600 m). We suspect that a revi-
sion of Eidmann's material would show this identification to be incorrect.
Ecology. M. cachmiriensis has been found at altitudes between 2100 and 3500 m, but
its ecology is poorly known. Philip Ward (pers, comm.) found it in open pine and fir-pine
forests, while Menozzi (1939) noted this species was found on irrigated fields in oases in
arid territories (but note comments on Menozzi's material - above). Alate queens and males
have been collected in mid July.

Myrmica cagnianti Espadaler, 1996

(Figs 47, 48)

Myrmica cagnianti Espadaler, 1996: 68, w, q, m, Morocco.

108
 Type localities and type specimens. "Holotype: worker, Morocco, Oukaimeden, Higher Atlas, from
a colony collected under stone at 2450 m, near a small stream, 12 V 1983, Espadaler leg., in author's
collection. Paratypes: 46 workers, 18 females, 4 males, Morocco, Oukaimeden, Higher Atlas, from
a colony collected under stone at 2450 m, near a small stream, 12 V 1983, Espadaler leg. Two worker
paratypes and one female at the following institutions: Museum of Comparative Zoology, Harvard;
Museum d'Histoire Naturelle, Genève; Muséum d'Histoire Naturelle, Paris".
Material examined. Paratypes, about 30 w, 2 q, "Morocco, Oukaimeden, Higher Atlas, 12.V.1983,
Espadaler leg." (ESPADALER).
Distribution (Map 19). Morocco, Algeria (Atlas Mts.).
Etymology. M. cagnianti: the author dedicated this species to his friend, the myrmeco-
logist Prof. Henri Cagniant of The Paul Sabatier University, Toulouse, France.
Notes. M. cagnianti belongs to the scabrinodis-complex of the scabrinodis species
group. It is probably endemic to the Atlas mountains and is closely related to M. aloba, dif-
fering from that species only by its fine sculpture, by the shape of petiole, etc. (for details
see Espadaler 1996).
Ecology. Probably very similar to that of M. aloba. However, the only recorded data is
that it inhabits humid biotopes in mountains between 1650 and 2600 m a.s.l. and its
colonies are polygynous (Espadaler 1996).

Map 19. Distribution of M. cagnianti.

Myrmica caucasicola A r n o l d i , 1 9 3 4
(Figs 49-51)

Myrmica schencki nat. caucasicola Arnoldi, 1934: 172, w (part.), m, Azerbaijan, Armenia.
Myrmica schencki var. caucasicola: Weber 1948: 298.
Myrmica schencki subsp. caucasicola: Arnoldi 1970: 1838.
Myrmica caucasicola: Radchenko 1994b: 44, 1994c: 140, 1994e: 79; Arakelyan 1994: 24; Bolton 1995:
278; Seifert 2003: 147; Radchenko, Elmes and Alicata 2006: 519, q.

109
 Type localities and type specimens. "In Transkaukasien - Armenien, Azerbeidzhan, Talysh ver­
breitet; wohnt im Gebirge 700-1600 m. hoch." (see also Notes below).
Material examined. Lectotype, w (designated by Seifert, 2003, who erroneously noted the type lo­
cality as "Tallish Mountains, Nodus-galasi"), "Нюднс-галаси, Талыш, Арнольди, 14.vii.1929, А 4286"
[Nudis-galasi, Talysh, 14.vii.l929, a 4286, leg. Arnoldi"] (MOSCOW); paralectotypes: 39 w, "Нюдис-
галаси, Талыш, Арнольди, 14.VII.1929" [Nudis-galasi, Talysh, 14.vii.1929, leg. Arnoldi], Nos. a 4224,
a 4232, a 4237, a 4240, a 4242, a 4282, 4286, a 4294; No. 7164, VI.1936; 3 w, "Алазапин, 800-1000
M, Талыш, VI.1929, Арнольди" [Alazapin, 800-1000 m, Talysh, a 4240 (Siel), vi.1929, leg. Arnoldi]; 1 q,
2 m, "Армения, Караулис, ix.30, A 4680, Аветисян" [Armenia, Karaulis, ix.30, a 4680, leg. Avetisian"
(MOSCOW, PETERSBURG); non-type material: about 30 w, 1 q from Talysh.
Distribution (Map 20). Azerbaijan, Armenia.
Etymology. M. caucasicola: from the name Caucasus with the Latin suffix incola - in­
habitant of, to indicate that this is a species of the Caucasus region that includes the type
localities.
Notes. M. caucasicola belongs to the schencki species group and most resembles
M. schencki. Workers and queens differ from the latter mainly by their somewhat wider
frons and by a much smaller lobe at the scape bend. Males of M. caucasicola have very long,
erect hairs on the tibiae and tarsi and are hardly distinguishable from those of M. ravasinii.
On the other hand, this feature well discriminates males of both species from all other
species of the schencki-group.
Among Arnoldi's original type material we found 6 workers (KIEV, LONDON), labelled
as "4810, Армения, Дилижан, VII.30, К. Арнольди" [Armenia, Dilizhan, vii.30, К.
Arnoldi], "Myrmica schencki caucasicola Arnoldi typ.", but in fact they are M. schencki.
Therefore, we excluded them from the syntype series.
Unfortunately, males are known only from three paralectotypes from Armenia, Karaulis
that were collected without workers. They are practically indistinguishable from the males
of M ravasinii, and we can not be sure whether they really belong to M. caucasicola (or to
M. ravasinii). This question can only be resolved when a nest sample of M. caucasicola
containing both workers and males is found.

Map 20. Distribution of M. caucasicola.

110
 Ecology is poorly known, except that this species inhabits mainly dry mountain forests
at an altitude 700-1600 m a.s.l.

Myrmica collingwoodi Radchenko et Elmes, 1998

(Figs 52, 53)

Myrmica collingwoodi Radchenko et Elmes, 1998: 23, w, q, Bhutan; 2001b: 243.

Type localities and type specimens. Bhutan, for details see material examined.
Material examined. Holotype, w, "Dorjula, 3100 m, 26.06" [Bhutan],."Nat.-Hist. Museum BASLE
- Bhutan Expedition 1972, Alk. No 56" (BASLE); paratypes: 9 w, 1 q with the same label (probably
same nest) as holotype; 6 w, with the same label, but date of collecting 6.06; 6 w, "Dorjula, 2450-3100
m, 6.06", "Nat.-Hist. Museum BASLE - Bhutan Expedition 1972, No 24-36" (BASLE, LONDON,
KIEV, ELMES).
Distribution (Map 21). Bhutan.
Etymology. M. collingwoodi: dedicated to our good friend, the eminent British ento­
mologist Cedric A. Collingwood, who at the time of writing is one of the oldest active
myrmecologists.
Notes. M. collingwoodi belongs to boltoni-complex of the ritae species group. It differs
from the other species of this complex by its much more finely punctated head dorsum, by
the finer sculpture of its alitrunk dorsum, by possession of outstanding hairs on the temples
and genae, and by its lighter colour. Additionally, it differs from M. martensi by its distinctly
longer propodeal spines, and from M. boltoni by its straight propodeal spines, which are di­
rected backwards and somewhat upwards. Males are unknown.
Ecologyis unknown, except that this species was found at altitudes 2450-3100 m a.s.l.

Map 21. Distribution of M. collingwoodi

Ш
 Myrmica commarginata Ruzsky, 1905
(Fig. 54)

Myrmica commarginata Ruzsky, 1905: 708, w, Russia; Emery 1908a: 166, 1921: 37; Ruzsky 1925: 46,
1936: 94; Weber 1947: 466; Radchenko 1994b: 43, 1994c: 138,1994f: 82, 1994g: 105; Bolton 1995:
278; Pfeiffer et al. 2007:4.

Type localities and type specimens. "Мсстонахожд. : Забайкальская обл., песчаная степь по дороге из
ел. Кяхты в г. Троипкосавск (в 23 в. от города), рабочие, 27. VII. 1901 (Моллссон)" [Locality: Transbaikal
Prov., sandy steppe by the road from Kyakhta to Troitskosavsk (23 km from the town), ww 27.vii.1901,
leg. Molleson].
Material examined. Neotype, w (designated by Radchenko 1994f), "100 км севернее Канска, No. 1,
1963 (В. Дмитриснко)" [100 km N of Kansk, No. 1, 1963, leg. V. Dmitrienko] (MOSCOW); non-type
material: 1 w, Mongolia, Hovd Aimak, Durgun soum, Chonoharaihiin gol. Deris, 1154 m, leg.
A. Stubbe, 27.viii.2003.
Distribution (Map 22). South Siberia, Mongolia.
Etymology. M. commarginata: from the Latin word (F.) marginata = enclosed with
edges or borders, with the prefix com = together with, to give "with margins" describing the
unusual shape of the alitrunk (see below).
Notes. M. commarginata is an enigmatic taxon that was placed to the lobicornis species
group by Radchenko (1994b). It has an uncertain taxonomic status because except for the
most unusual morphological feature after which it was named (see below) it is similar to
sympatric fob¿corm's-group species, such as M. kasczenkoi, M. displicentia, or even
M. divergens. Furthermore, it appears to be unusually rare for a Siberian Myrmica species,
having been found in only seven localities since its first description: Krasnoyarsk Krai,
Cbitinskaya and Amurskaya Prov, of Russia, and in Mongolia (Ruzsky 1925, 1936; Rad-
chenko 1994f; Pfeiffer et al. 2007).

Map 22. Distribution of M. commarginata.

112
 The mesonotum and propodeum of M. commarginata specimens are strongly constricted
laterally, so that their dorsal surface is narrow and delineated laterally by sharp ridges, which
merge with the outer bases of propodeal spines. This feature is not seen in any other known
Myrmica species (although we have occasionally seen something similar when young
workers with soft cuticles have been deformed by being held too tightly in forceps). We can
not exclude the possibility that such deformed individuals or perhaps naturally aberrant
workers (even from several different species) have been included in the name
M. commarginata. This will only be resolved when good consistent, colony samples con­
taining queens and males are found. Queens and males are unknown.
Ecology is poorly known, except that it inhabits steppes and dry meadows, often with
sandy soil.

Myrmica constricta Karawajew, 1934

(Figs 55-57)

Myrmica rugulosa var. constricta Karawajew, 1934: 74 (replacement name for Myrmica rugulosa var.
minuta Karawajew, 1929: 204). Junior synonym of M. rugulosa: Arnoldi 1970: 1840; of M. hel­
lenica: Radchenko 2009: 70; revived from synonymy and raised to species: Seifert et al. 2009: 72.
Myrmica rugulosa' var. minuta Karawajew, 1929: 204, w, m, Ukraine (junior primary homonym
of Myrmica minuta Say, 1836: 294 and Myrmica laevinodis var. minuta Ruzsky, 1905: 670).
Synonymy by Radchenko 2009: 70 (as synonym of M. hellenica); not confirmed, see Seifert et al.
2009 and Notes below.
Myrmica rugulosa var. slobodensis Karawajew, 1936: 274 (first available use of Myrmica rugulosa
rugulosa var. slobodensis Arnoldi, 1934: 162) (not necessary proposed replacement name for Myr­
mica rugulosa var. constricta Karawajew, 1934: 74). Junior synonym of M. rugulosa: Arnoldi 1970:
1840; of M. hellenica: Radchenko 2009: 70; of M. constricta: Seifert et al. 2009: 72; the latter
confirmed here.

Type localities and type specimens. Myrmica rugulosa var. minuta: "Umgegend von Kiew, jenseits
des Dnjepr, Kiefernwald hinter der Nikolskaja Slobodka, 1.VIII.1919 (Nr. 2770), W. Karawajew, ww
und einige mm aus dem Nest".
Material examined. Syntypes oi Myrmica rugulosa var. constricta: 71 w, 9 m, "окр. Киева, Николь­
ская слободка, сосновый лес за Днепром" [vicinity of Kiev, Nilolskaya slobodka, pine forest on the other
side of Dnieper], "2770 Coll. Karawajewi", "M. (Myrmica) constricta Karaw. Typus", "Syntypus Myr­
mica rugulosa var. constricta" (KIEV); 1 w, "Cotype", "M. rugai, v. slobodensis Kar." (MOSCOW);
non-type material: about 50 w, 1 q, 1 m from Poland and Finland.
Distribution (Map 23). This species is quite widely, but sporadically distributed in Cen­
tral and partly southern Europe: Ukraine, Poland, southern Finland, Germany, Austria,
Switzerland, Czech Republic, Romania, Italy, Serbia, Bulgaria and Greece.
Etymology. M. constricta: from the Latin constricta = small, limited in size, or marked
by contraction, tightening, or cornpressed, contracted; as this is a replacement name for
minuta, the meaning "small" seems the most reasonable. M. minuta: from the Latin mi­
nuta = small or minute, to indicate its relatively small size for a Myrmica species.
M. slobodensis: from the name Slobodka combined with the Latin suffix ensis = place of
origin, to indicate that the type locality was Nikolskaja-Slobodka, Ukraine.
Notes. M. constricta belongs to the rugulosa-comp\ex of the scabrinodis species group
(see also Radchenko and Elmes 2004). It most resembles M. rugulosa but differs from the

113
latter by the presence of a small narrow ridge at the base of its scape. Female castes differ
from the morphologically similar M. hellenica by having a less developed lobe on the scape
base, and males differ from the latter by having shorter standing hairs on the legs (for more
details see Seifert et al. 2009).
Karawajew (1929) described from the vicinity of Kiev M. rugulosa var. minuta. How-
ever, this name was preoccupied (junior primary homonym of M minuta Say, 1836 and
M. laevinodis var. minuta Ruzsky, 1905) and Karawajew (1934) proposed the replacement
name - M. rugulosa var. constricta. Simultaneously Arnoldi (1934) also gave the replace-
ment name for Karawajew's var. minuta - var. slobodensis, referring it to "Karavajev,
1932"; most probably Arnoldi expected that a paper by Karawajew would be published in
1932 using this name because he had discussed the matter with Karawajew and in private
correspondence (see Karawajew 1936: 274). Later Karawajew (1936) believed that
Arnoldi's name slobodensis had priority and used it instead constricta. However, Arnoldi's
(1934) name Myrmica rugulosa rugulosa var. slobodensis is unavailable (quadrinomen)
and therefore the first available use of the name M. rugulosa var. slobodensis was made by
Karawajew (1936). Consequently, the name M. rugulosa var. constricta Karawajew, 1934
has priority and "slobodensis" should be considered as an unnecessary proposed replace-
ment name. Later Arnoldi considered var. slobodensis (= constricta) to be junior synonym
of M. rugulosa (Arnoldi 1970; see also Radchenko 1994d), whereas Radchenko (2009) be-
lieved it to be a synonym of M. hellenica. Most recently, Seifert et al. (2009) showed that
M. constricta is morphologically distinct from (but related to) M. hellenica and can be
considered to be a good species (see also Notes to M. hellenica). We accept this latest
treatment.
Ecology. M. constricta inhabits rather xerothermic habitats, usually open places with
sandy soils, and relatively sparse vegetation in full exposure to the sun. In semi-natural
habitats it builds nests in the ground, occasionally under stones, but more often among
grass roots, in dry sandy meadows that border rivers, lakes, ponds and seashores. In urban
environments it is quite common in lawns and flower borders in city parks and around

Map 23. Distribution of M. constricta.

114
public buildings. However, although such habitats are superficially very dry, especially in
mid and late summer, they are generally moister than it appears; urban habitats tend to be
artificially watered and natural habitats tend to have a fairly high water table and nests
often penetrate the soil to a depth where it becomes dampish. Colonies may consist from
several hundred to more than 1500 workers and several queens, and the species can
probably form very large populations or super colonies (see notes to M. rugulosa). Nuptial
flights are in August-September.

Myrmica curiosa Radchenko, Zhou et Elmes, 2008

(Figs 58, 59)

Myrmica curiosa Radchenko, Zhou et Elmes, 2008: 769, w, q, China (in Radchenko et al. 2008).

Type localities and type specimens. China, for details see material examined.
Material examined. Holotype, w, "China, Sichuan Prov., Hailugou Nat. Park, 29°37'N, 102°05'E,
1700 m, 14X.88, leg. С Carpenter et N. J. Nordensten" (HARVARD); paratypes: 1 w with the same
label as the holotype; 1 w, "China, Hunan Prov., Ba Da Gong Mountain, No. 9, 15.x.2003, leg.
]. Huang"; 1 q, "China, Yunnan, Hobasan Mts., 2800-3150 m, 5-13.vii.2002, 27°20'N, 100°09'E, leg.
S. Becváf and R. Fouqué" (KIEV, GUILIN, MILAN).
Distribution (Map 24). The species is known from the type series, collected in Hunan,
Sichuan and Yunnan Provinces of China.
Etymology. M. curiosa: the name derived from the Latin word curiosa (f.) = "taking
care with" or "diligence", but used here in the modem sense of curious = puzzling or un-
usual, to indicate its curious combination of morphological features.
Notes. We have nothing to add to the notes we made in the original description of this
species (Radchenko et al. 2008) which we reproduce here: "M. curiosa possesses many fea-
tures that are intermediate between those of different species groups, and we can not assign

Map 24. Distribution of M. curiosa.

115
it to any of them. Its relatively large size, slender and long alitrunk and scape recall species
of the ritae-group, but it clearly differs from them by its not-notched anterior clypeal mar­
gin, a much shorter petiolar node, not-pointed upper latero-ventral corners of the head, and
rounded propodeal lobes. On the other hand, M. curiosa resembles species from the inezae-
group by the shape of its petiole, but differs by its much shorter propodeal spines. In addi­
tion, in many respects it has some features found in several species of the rabra-group (e.g.
shape of frontal lobes and clypeus, etc.). Moreover, the long, multidentate, masticatory mar­
gin of mandibles, with more than ten teeth, is one of the diagnostic features of Myrmica's
"sister" genus - Manica Jurine. This feature distinguishes M. curiosa from all other known
Myrmica species except for the Himalayan M. hecate, from which it differs by having frontal
carinae that curve outwards to merge with the rugae that surround antennal sockets" (see
also Radchenko and Elmes 2001b). When males of M. curiosa are discovered, its taxonomic
position could become clearer. Males are unknown.
Ecologyis unknown.

Myrmica deplanata Emery, 1921

(Figs 60-62)

Myrmica lobicornis var. deplanata Emery, 1921: 38, w (first available use of Myrmica scabrinodis
subsp. lobicornis var. deplanata Ruzsky, 1905: 700, Ukraine, Russia, Georgia; Emery 1908a: 179);
Karawajew 1927a: 283, 1927b: 259; Weber 1948: 276.
Myrmica schencki var. deplanata: Finzi 1926: 111.
Myrmica deplanata: Arnoldi 1934: 169; Karawajew 1934: 83, q, m, 1936: 275; Stitz 1939: 105; Sadil
1940: 102; Arnoldi 1970: 1842; Tarbinsky 1976: 31; Arnoldi, Dlussky 1978: 535; Agosti and
Collingwood 1987a: 53, 1987b: 268; Seifert 1988: 36; Dlussky et al. 1990: 184; Bolton 1995: 278;
Radchenko and Elmes 2009b: 511.
Myrmica moravica Soudek, 1922: 45, w, q, Czechia (Moravia) (also described as a new by Soudek,
1923: 107); 1925: 35, m; Finzi 1926: 104; Karawajew 1936: 275; Sadil 1939: 106, 1940: 102; Weber
1950: 213. Synonymy by Karawajew 1934: 83; Seifert 1988: 36; Bolton 1995: 281 (as synonym of
M. deplanata); Radchenko 1994e: 78; Seifert 2003: 150; Radchenko, Elmes and Alicata 2006: 516
(as synonym of M. lacustris); Radchenko and Elmes 2009b: 511 (as synonym of M. deplanata); the
latter confirmed here.
Myrmica deplanata moravica: Novak and Sadil 1939: 106, 1941: 80; Kratochvil, in Kratochvil et al.
1944: 40; Sadil 1952: 263; Arnoldi 1970: 1843.
Myrmica moravica var. dagmarae Sadil, 1939: 108, w, q, m, Czechia, syn. nov.
Myrmica lobicornis var. plana Karawajew, 1927a: 283, w, Ukraine; 1927b: 259; Arnoldi 1934: 170, q,
m; Karawajew 1936: 276; Weber, 1948: 302. Synonymy by Karawajew 1929: 208, 1934: 83; Arnoldi
1970: 1843, Seifert 1988: 36; Bolton 1995: 282 (as synonym of M. deplanata); Radchenko 1994e:
78, Seifert 2003: 150; Radchenko, Elmes and Alicata 2006: 516 (as synonym of M. lacustris);
Radchenko and Elmes 2009b: 511 (as synonym of M. deplanata); the latter confirmed here.
Myrmica lacustris: Radchenko 1994b: 44, 1994c: 140, 1994e: 77; Bolton 1995: 280; Seifert 2003: 150;
Radchenko et al. 2005: 168; Radchenko, Elmes and Alicata 2006: 516, nec Ruzsky 1905: 686,
Emery 1908a: 177, 1921: 40, Weber 1950: 206; not confirmed, see Notes to M. stangeana, below.

Type localities and type specimens. M. deplanata: "Мсстонахожд.: Крым, колон. Гсрцснбсрг,
12.VI.1901 (Муралсвич). Кавказ, на южн. склон главн. хребта, ст. Они, Воснно-Осстин. дор., VII. 99;
окр. Батума (1900). Саратовская губ., окр. Царицына, степь (VI. 1900). Оренбургская, степь на пути от
Орска до ст. Магнитной, 4.VII.94. Саратовская, ок. с. Орловки, Цариц, у. на вале, 15.VII.03 (Келлер)"

116
 [Localities: Crimea, colony Gertsenberg, 12.vi.1901, leg. Muralevich; Caucasus, S slope on the Main
Range, station Oni, Voyenno-Osetinskaya road, vii.99; vicinity of Batumi, 1900. Saratovskaya Gubcr-
nia, vicinity of Tsaritsyn, steppe, vi.1900. Orenburgskaya Gubernia, steppe, on the road from Orsk to
station Magnitnaya, 4.vii.94. Saratovskaya Gubernia, vicinity of vil. Orlovka, Tsaritsyn Distr., on a wall,
15.viii.03, leg. Keller] [now: Crimea, Volgograd and south of Chelyabinsk Provinces of Russia, Geor-
gia]. M. moravica: "Nalczl jsem na jizní Morave" [Moravia, no precise locality given]. M. lobicornis var.
plana: "Askania Nova (kontinentaler Teil des ehemaligen Taurischcn Gouvernements), Steppe, Spät
und Dobrschansky, 1923, einige ww". M. moravica var. dagmarae: "Praha".
Material examined. Neotype of M. deplanata, w, "Ukraine, vicinity of Kherson, valley of riv.
Vir'ovchyna, steppe, 3.ÌX.2000, leg. S. Bondar" (designated by Radchenko and Elmes, 2009b) (KIEV);
syntypes of M. moravica: 4 w on the same pin, "Myrmica moravica Soudek", "Brno Moravia" (most
probably both labels written by Soudek), "Museo Genoa coll. С Emery (dono 1925)" (new label,
printed by F. Penati, 2007); ); 7 workers (4 and 3 on two pins), "Brno Moravia" (most probably both
written by Soudek)", "Museo Genoa coll. С Emery (dono 1925)" (new label, printed by F. Penati)
(GENOA); syntypes of M. lobicornis var. plana: 4 workers, "Askania-Nova, Dobrschanskiy, 1923"
(KIEV, MOSCOW); non-type material: about 100 w, about 30 q and m from whole area.
Distribution (Map 25). Central (Austria, Czech Republic, Slovakia, SW Poland) and
Southern (former Yugoslavia, Romania, Bulgaria, Thrace) Europe, south of Ukraine and Eu­
ropean part of Russia (to the north as far as Nizhniy Novgorod), Transcaucasus, Iran, Turk­
menistan, south of West Siberia, northern Kazakhstan, Tien-Shan, to the east as far as the
Altai Mts.
Ruzsky (1925, 1936) recorded M. deplanata for Transbaikalia: Verkhneudinsk (now -
Ulan-Ude, Buryatia), vil. Kudara; vicinity of Chita. We can not find corresponding material
in any collections, and based on our current understanding of the distribution of
M. deplanata, these identifications seem doubtful and may belong to several Siberian
species from lobicornis-group, which are superficially similar to M. deplanata.
Etymology. M. deplanata: derived from the word deplanate meaning flattened (above
and below), which itself derives from the Latin word complanatus, it presumably refers to

Map 25. Distribution of M. deplanata.

117
the "more flattened" propodeum dorsum compared to M. scabrinodis (see below).
M. moravica: from the name Moravia with the adjective suffix for nouns ica (from the Greek
IKO) = belonging to, from, to indicate that the types were collected in that region of the
Czech Republic. M. plana: derived from the Latin word plana (f.) = flat or level, pre­
sumably to describe the shape of its propodeum. M. dagmarae: this species was presumably
dedicated to a female friend or relative (wife?) of Sadil, the given name Dagmar is quite
common in the Czech Republic.
Notes. M. deplanata belongs to the schencki species group and differs from sympatric
M. schencki, M. caucasicola and M. koreana by its wider frons and less extended frontal
lobes; the males well differ from all other schencki-group species by their very short scape
(for details see Radchenko, Elmes and Alicata 2006, and Key, Chapter 5.2).
The name M. deplanata was widely used in myrmecological literature until
Radchenko (1994e) synonymised it with M. lacustris. He did this because at that time the
types of both M. deplanata and var. lacustris seemed lost and based on the descriptions
these names appeared to be synonyms (see below). The name M. scabrinodis subsp. lobi­
cornis var. deplanata Ruzsky, 1905 was unavailable (quadrinomen) consequently the name
M. scabrinodis var. lacustris Ruzsky, 1905 had priority. M. lacustris was accepted as the
senior synonym of M. deplanata in recent publications (e.g. Seifert 2003; Radchenko et al.
2005,2006).
Apart from the missing types, the very brief and at least partly ambiguous description of
M. scabrinodis var. lacustris (Ruzsky 1905: 686) always was a problem. Translated from
Russian it reads: "Workers. Anterior clypeal margin with the small notch medially.
Propodeal spines are short, equal to 1/2 or at most 2/3 of the length of dorsal surface of
propodeum. Frontal triangle striated distally. Antennal scape at the base is with a small, in­
clined dentiform lobe. Metanotal groove weak, not deep, as a result mesonotum and
propodeal dorsum more flattened (in typical M. scabrinodis they are more convex). Body
rugosity is finer. Body colour is as in the type [i.e. M. scabrinodis], but the whole gaster, ex­
cept of its tip, is dark brown". This description could apply to many Myrmica species and
one of the most important diagnostic features, the shape and direction of the lobe at the base
of antennal scape, is hardly understandable: it can be interpreted either as horizontal (as in
species of the scabrinodis-group) or as vertical (as in species of the schencki- and lobicor­
nis -groups). On the other hand, a medially notched anterior clypeal margin is a significant
feature of the latter two groups. Finally, in the course of his revision of the Asian Myrmica,
Radchenko (1994e) found in MOSCOW a queen that had an original Ruzsky label:
"H.-Петровский, Акмолинский уезд, M. scabrinodis var. lacustris Ruzs." [N.-Petrovsky,
Akmolinslky uezd; now - Akmola Province of Kazakhstan]. This specimen could not belong
to the type series of var. lacustris (which was described only from workers) and Radchenko
believing that Ruzsky at least determined his "own" species correctly, designated this queen
as the neotype of M. lacustris.
Recently, the discovery in the GENOA of Ruzsky's original types of M. lacustris (two
workers) completely negated the above approach because these specimens clearly belong to
the scabrinodis-group and were attributed to M. stangeana (Radchenko and Elmes 2009b).
We therefore revived from synonymy the name M. deplanata that had been in regular use
for this species prior to Radchenko's (1994e) revision, and designated a neotype of this
species from a nest series taken in Ukraine.
Also, we found 7 workers (syntypes) of M. moravica in Emery's collection (GENOA)
and were able to confirm the earlier synonymy. Additionally, we examined 2 workers

118
labelled as "Hubochepy, Praha, Bohemia, J. Sadil, 1938", "Myrmica moravica Soud., var.
п., cotype" (MOSCOW), and 6 workers with the labels "Morava, Sadil leg.", "Myrmica
moravica Soud. det." (KIEV); these specimens, which can not be attributed formally to the
type series of M. moravica, all conformed toM. deplanata.
There remains three names to be considered. First, M. moravica var. dagmarae described
from Prague by Sadil (1939). This name was completely forgotten by all subsequent au­
thors and has been never used after 1939. We have not examined type specimens of this
form, but Sadil (1939) wrote that workers differ from M. moravica only by the slightly
coarser and more regular rugosity on the alitrunk, queens are like M. moravica, and males
- like M. deplanata. We are almost certain that this is just a variant of M. moravica and as
that species is a junior synonym of M. deplanata we are quite confident that var. dagmarae
is conspecific with M. deplanata. Secondly, the nomen nudum created by Arnoldi (1948),
M. deplanata subsp. lulakeranica. In the MOSCOW collection we found corresponding
material from Talysh (SE Azerbaijan) and similar nest series from Kopet Dagh. Apart from
a smaller reduced vertical dentiform lobe on the scape bend these series appeared to be
typical for M. deplanata and probably represent a regional variant. Thirdly, the unavailable
name M. scabrinodis subsp. lobicornis var. grandis Kuznetsov-ugamsky, 1927, described
from Osh Province, Kyrgyzstan can be attributed to M. deplanata having been synonymised
under the name M. lacustris (Radchenko 1994e).
Ecology. M. deplanata is one of the most thermo-xerophilous species of the genus Myr­
mica. Although it has a widespread distribution, nests of M. deplanata are comparatively
rare, it has never been reported as being abundant even in apparently optimal habitats.
It inhabits mostly steppes and steppe-like habitats at low altitudes, where it prefers stony
sites with low, often sparse vegetation. Nests are built in the soil, often under stones in
places that become very hot in summer months. Although it is reportedly tolerant of high
soil salinity it is not particularly associated with the damper steppe areas that can become
quite salted. Sexuais have been found in nests from July-September.

Myrmica displicentia Bolton, 1995

(Figs 63-65)

Myrmica displicentia Bolton, 1995:278 (replacement name for Myrmica bicolor Kupyanskaya, 1986b: 94).
Myrmica bicolor Kupyanskaya, 1986b: 94, w, q, m, Russia (junior primary homonym of Myrmica
bicolor Heer, 1867: 31); 1990: 106; Radchenko 1994b: 43, 1994c: 138, 1994f: 86.

Type localities and type specimens. "Голотип: п-ов Камчатка, Эссо, близ горячего источника, из почвы,
17.VIII.1976, 1 рабочий (Купянская). Паратипы: п-ов Камчатка, Эссо, 17.VIII 1976, 8 рабочих, 1 самка,
1 самец из той же гнездовой серии; Пущино, обочина дороги, из почвы, 3.VIII.1976, 4 рабочих (Купянская);
Магаданская обл., верховья Колымы, Дстрин, песчаные косы, 17.VII. 1963, 10 рабочих, 2 самки, 2 самца
(Жигульская). Голотип и часть паратипов хранятся в Зоологическом музее Московского государственного
университета, остальные паратипы - в Биолого-почвснном институте ДВНЦ АН СССР (Владивосток)"
[Holotype: Kamchatka, Esso, near hot stream; from soil, 17.viii.l976, 1 w, leg. Kupyanskaya; paratypes:
Kamchatka, Esso, 17.viii.1976, 8 w, 1 q, 1 m, from the nest of holotype; Puschino, roadside, from soil,
3.VÍÜ.1976, 4 w, leg. Kupyanskaya; Magadan Prov., upper reaches of riv. Kolyma, Detrin, sandy banks,
17.vii.1963, 10 w, 2 q, 2 m, leg. Zhigulskaya. Holotype and part of paratypes are preserved in Zool-
ogical Museum of Moscow State University, the rest of paratypes - in Biological and Soil Institute of
FESC Ac. Sci. of USSR in Vladivostok].

119
 Material examined. Holotype, w, "Камчатка, окр. Эссо, близ горячего источника, из почвы, 17.8.76,
Купянская" [Kamchatka, Esso, near hot stream, from soils, 17.viii.1976, Kupyanskaya] (MOSCOW);
paratypes: 9 w, 1 m, from the nest of holotype; 6 w, "Камчатка, окр. Пущино, обочина дороги, из почвы,
3.8.76, Купянская" [Kamchatka, vicinity of Puschino, roadside, from soil, 3.viii.1976, Kupyanskaya];
2 w, 2 m, "Магаданская обл., верховья Колымы, Дстрин, 17.VII.1963, Жигульская" [Magadan Prov.,
upper reaches of riv. Kolyma, Detrin, No. 321, 17.vii.1963, leg. Zhigulskaya] (MOSCOW, VLADI­
VOSTOK); non-type material: about 50 w, 5 q, 9 m from Kamchatka and Magadan Province..
Distribution (Map 26). Kamchatka Peninsula and Magadan Prov, of Russia.
Etymology. M. displicentia: the Latin word displicentia means dissatisfaction or dis­
content. We are not sure whether B. Bolton intended this or whether he made-up a word
from the English di (two) and spliced (joined together) to represent the bicoloured body.
M. bicolor: from the Latin word color = colour with the prefix bi derived from bis = twice,
to describe its bicoloured body.
Notes. M. displicentia belongs to the lobicornis species group and is characterized by the
strongly curved (not angled) bend at the base of the scape that has no vertical lobe or dent.
By this feature it is similar to some other species form this group (e.g. M. sulcinodis,
M. ademonia or M. kasczenkoi), but differs from them by its distinctly bicoloured body,
with the yellowish-red alitrunk, contrasting with much darker head and gaster; additionally,
M. displicentia well differs from M. sulcinodis by its much less coarse sculpture and by the
shape of its petiole (for more details see Key, Chapter 5.3).
Ecology. M. displicentia prefers well insolated and warm habitats such as roadsides,
open slopes, glades, forest clearing, etc. where it builds nests in the soil, often under stones
(see Kupyanskaya 1986b, 1990; Berman et al. 2010). Colonies examined comprise a few
hundreds to 1500 workers with a maximum of 3 queens. Everywhere it is quite a rare
species. Gynes and males were observed in June-August.

Map 26. DistributionofM . displicentia.

120
 Myrmica divergens Karawajew, 1931
(Figs 66-68)

Myrmica bergi var. divergens Karawajew, 1931b: 105, w, Russia; Weber 1947: 473; Pisarski 1969a:
227; Dlussky and Pisarski 1970: 85; Pisarski and Krzysztofiak 1981: 155; Bolton 1995: 278.
Junior synonym of M. bergi: Radchenko 1994d: 76, 1994g: 109. Revived from synonymy and raised to
species: Radchenko et al. 2002: 411, w, q, m; Pfeiffer et al. 2007: 4.

Type localities and type specimens. "Insel Char-jalach, 50 km unterhalb Olekminsk, 11.6.1925,
L. Bianchi, 1 arb.".
Material examined. Holotype, w, "о-в Харъялах, 50 км ниже Олскминска, 11.6.1925, Л. Бианки",
Якутск. Экспсд АН", "М. bergi Ruz. var. divergens Kar. W. Karawajew det.", "HolotypusMyrmica bergi
divergens Karaw." (KIEV); non-type material: > 200 w, 17 q, 13 m from South Siberia and Mongolia.
Distribution (Map 27). South and East Siberia (to the west - until Altai Mts.),
Mongolia.
Etymology. M. divergens: from the Latin word divergens - divergent, to indicate the
characteristic kind of rugosity on the head dorsum.
Notes. Based on the morphological characters of the female castes, M. divergens could
be placed either in the scabrinodis or lobicornis species groups. For example, the same nest
sample may contain some individuals with a broadly rounded anterior clypeal margin and
no medial notch, a feature of the scabrinodis-group, and others that have a shallowly
notched clypeus, as seen in species of the lobicornis-group. In addition, M. divergens has
antennal scapes that are strongly curved or even slightly angled at the base but with no
trace of a carina or lobe, this feature distinguishes M. divergens from all schencki-group
species, which are typified by the presence of vertical lobe (or at least denticle) on the base
of the antennal scape. However, males have a short antennal scape (SIj < 0.40, SI2 < 0.43),
which is a feature of the scabrinodis and sc/ie«c/n'-groups and completely distinguishes
M. divergens from any species from the lobicornis-group (SI, > 0.65, SI2 > 0.68 ). This
combination of male and female features led us to place M. divergens in the scabrinodis-
group (Radchenko et al. 2002).
However, this approach has recently been contradicted by the very interesting results,
obtained by Jansen et al. (2010) based on molecular data. These showed that M. divergens
is not closely related to any species of the scabrinodis-group, including the morphologically
similar M. bergi and M. gallienii, but forms a common clade with some Siberian species of
the lobicornis-group. If this result is confirmed by further study, then because we believe
a short male scape to be ancestral, we suppose that M. divergens might be a relict species,
close to the ancestral branch of the lobicornis-group as a whole. This idea would also
explain the Zoogeographie puzzle as to why in South and East Siberia M. divergens should
be the only member of the scabrinodis-group, whose origin and centre of diversity most
probably is connected with the Euro-Caucasian region (see Radchenko 1994b). The range
M. divergens is dominated by lobicornis-group species.
Ecology. Based on collecting data it seems probable that M. divergens' preferred habi­
tat is the intrazonal woodland between riparian meadows and steppe with rivers providing
the main routes for connection between populations. The type specimen was taken towards
the north of its range on an island in the river Lena. The best data is from collections made
in north Mongolia by Micha! Woyciechowski (see also notes to M. angulinodis). He col­
lected most colonies of M. divergens from "Riparian Woodland" at about 1000 m a.s.l. on
the south-facing bank of the Jeröö river, and some from "Shrublands" about 1100 m a.s.l.

121
 Map 27. Distribution of M. divergens.

that border the steeper valley sides (for vegetative details and photographs of these habitats
see Muehlenberg et al. 2000). Nests were constructed directly in the soil or in grass and
moss tussocks; one nest was found in the soil in a very wet peat bog in a river valley.
M. Woyciechowski found sexuais in the nest at the end of July and in August; a single male
was found participating in a mating swarm of M. angulinodis on 10th August 1999. Some-
what confusingly Pisarski (1969a) said that in Mongolia nests were found in the soil, under
and in moss and frequently in grass tussocks which conforms to the observations above, but
he then gives the habitat of M. divergens as "dry steppe, nests in the soil" (loc. cit., p. 227).
We have two nest samples collected by Bogdan Pisarski in 1964, from Larix forest near
Ulan Bator, but there is no further data. M. Woyciechowski collected very few dealate
queens in the Mongolian M. divergens nest samples, which suggests that their colonies are
not highly polygynous.

Myrmica draco Radchenko, Zhou et Elmes, 2001

(Figs 69-71)

Myrmica draco Radchenko, Zhou et Elmes, 2001: 214, w, q, m, China; Radchenko, Elmes and Viet
2006: 39.

Type localities and type specimens. China (Guangxi Prov.), for details see material examined.
Material examined. Holotype, w, "S. China, NE Guangxi Prov., Mao Er Shan, 1930 m, MES-125,
22.viii.1998, leg. J. Fellowes" (LONDON); paratypes: 1 w from the same nest as holotype; 1 q, 1 m,
same label, but No. MES-122; 4 w, same label, but No. MES-124; 1 w, 1 q (alate), same label, but No.
MES-128 (all leg. J. Fellowes); 6 w, same location, 29.viii.1997, No. 4, leg. S. Zhou; 5 w, same loca-
tion, 8.ÌX.1997, No. 3, leg. S. Zhou (KIEV, GUILIN, ELMES); non-type material: 1 w, China, Shaanxi
Prov., Quingling Nat. Res., 9.VÜ.2003, leg. Ru Tie; 1 w, China, N Yunnan, Dali Bai Nat. Aut. Pref.,
Diancang Shan, 4 km W Dali old town, 2900-3000 m, 25°41.4'N, 100°06.7'E, under stone, 31.viii.2003,
leg. Wrase.

122
 Distribution (Map 28). The species is known from the central and southern China
(Guangxi, Shaanxi and Yunnan Provinces).
Etymology. M. draco: this species is named for the dragon of Chinese mythology. Draco
is a Latin word derived from the Greek браксоу (dracon or dragon).
Notes. Originally Radchenko et al. (2001) placed M. draco to the ritae species group. We
discussed how the taxonomic position of this species is somewhat obscure because it shares
features of both the ritae- and botom-complexes of this group: M. draco has sculpture of
the alitrunk that is typical for the species from the ritae-complex, but sculpture of its head
and waist similar to species from the boZtom-complex (see also Radchenko and Elmes 1998).
However, after our more recent discovery of some similar new species from northern Viet­
nam, M. yamanei (Radchenko and Elmes, 2001) and M. schoedli (Radchenko et al., 2006),
we recognized a separate draco-complex, which contains these three species.
The males of M. draco have a long scape. Previously males were known only of M. se­
rica and M. indica, both of which have short scape and we assumed that this might be true
for all niae-group species. Until males from more species are discovered and described we
can not take this discrepancy any further. However, we speculate that males of all the
species from the draco-complex have long scapes while those of the riiae-complex are short;
in which case we would probably separate the ritee-group into two groups.
Ecology. Little is known about ecology of this species, except that it nests under rotten
wood, in the soil, in forests at altitudes of about 2000 m a.s.l.

Map 28. Distribution of M. draco.

Myrmica dshungarica Ruzsky, 1905

(Figs 72-74)

Myrmica rugosasubsp.dshungarica Ruzsky, 1905: 661, w, Kazakhstan; Kuznetsov-ugamsky 1927:

191, w, q, m.
Myrmica smythiesii dshungarica: Emery 1908a: 169, 1921: 42; Weber 1947: 458.

123
Myrmica dshungarica: Arnoldi 1976: 551; Tarbinsky 1976: 29, w, q, m; Marikovsky 1979: 165; Rad­
chenko 1994b: 41, 1994c: 137, 1994e: 73; Bolton 1995: 278; Schultz et al. 2006: 206, nec Whee­
ler G. С and Wheeler J. 1953: 123.
Myrmica laevinodis var. minuta Ruzsky, 1905: 670, w, Tajikistan (unresolved junior prirnary homo­
nym of Myrmica minuta Say, 1836: 294); Weber 1947: 455. Synonymy by Radchenko 1994e: 73;
confirmed here.
Myrmica rubra laevinodis var. minuta: Emery 1908a: 170; 1921: 39 (unavailable name).
Myrmica minuta: Arnoldi 1976: 548; Bolton 1995: 281.
Leptothorax svartshevskii Karawajew, 1916: 500, w, q; Bolton 1995: 245. Synonymy by Radchenko
1994e: 73; confirmed here.
Myrmica (Myrmica) rugulosa var. ruginodiformis Karawajew, 1929: 204, w, q, m, Kyrgyzstan; Weber
1948: 307. Synonymy by Arnoldi 1976: 551; Radchenko 1994e: 73; Bolton 1995: 282; confirmed
here.
Myrmica minuta subsp. iskanderi Arnoldi, 1976: 548, w, m, Tajikistan, Uzbekistan; Bolton 1995: 279.
Synonymy by Radchenko 1994e: 73; confirmed here.
Myrmica minuta subsp. tarbinskii Arnoldi, 1976: 549, w, m, Kyrgyzstan; Bolton 1995: 284. Synonymy
by Radchenko 1994e: 73; confirmed here.
Myrmica rugosa subsp. smythiesi: Ruzsky 1905: 659; Kuznetsov-ugamsky 1927: 193, misidentifica-
tions.

Type localities and type specimens. Myrmica rugosa subsp. dshungarica: "Мсстонахожд.:
Семирсчснская обл., долина р. Юй-Тас, южн. склон Джунгарск. Алатау, вые. 1000 м., близ границы
альпийских лугов, 29.VI1.1902 (Сапожников)" [Localities: Semirechenskaya Prov., valley of riv. Juj-Tas,
southern slope of Dzhungarian Alatau, alt. 1000 m, near border of alpine meadows, 29.vii.1902, leg.
Saposhnikov]. Myrmica laevinodis var. minuta: "Мсстонахожд.: Памир, Андсманын, перевал на
Памирском плоскогорий, рабочие, 28.VI.95 (Богоявленский)" [Localities: Pamir, Andermanyn, pass on
Pamir Plateau, workers, 28.vi.95, leg. Bogoyavlensky]. Leptothorax svartshevskii: "Киргиз-ата
Ферганской обл., 10.VII.14, Б. Сварчсвский, около 2 десятков рабочих и 4 крылатых самки" [Kirgiz-ata,
Fergana Prov., 10.vii.14, leg. В. Svarchevsky, about 20 workers and 4 alate gynes]. Myrmica rugulosa
var. ruginodiformis: "Karkara, östlich von Issyk-kul, am Oberlauf des Flusses Kegeni (Nebenfluss von
Iii), etwa 1950 Meter hoch, Gebirgsteppe mit Festuca, 16.VIH.1925 (Nr. 4003), N. Kuznctsov, ww,
2 q und mm". Myrmica minuta subsp. iskanderi: "Гиссарский хребет, Алай. Голотип (w), 20 паратипов,
аллотип (m): Искандср-Куль (О. Крыжановский); Джиргиталь, w, m (он же), Ллайский хребет
(Г. Длусский, Ю. Тарбинский), Юго-Западный Узбекистан, Байсун-Тау (Арнольди - 1942)" [Gissarsky
Range, Alai. Holotype (w), 20 paratypes, allotype (m): Iskander-КиГ, leg. O. Kryzhanovsky; Dzhirgi-
taP, w, m (same collector), Alaisky Range, leg. G. Dlussky and Yu. Tarbinsky; SW Uzbekistan, Bajsun-
Tau, leg. Arnoldi, 1942]. Myrmica minuta subsp. tarbinskii: "Таласский хребет, голотип (w) и 3 паратипа
(Ю. Тарбинский); хребет Каржантау (Обухова)" [Talassky Range, holotype (w) and 3 paratypes, leg. Yu.
Tarbinsky; Karzhantau Range, leg. Obukhova].
Material examined. Lectotype of Myrmica dshungarica, w (designated here), "Семирсчснская обл.,
1902, M. Р., Сапожников" [Semirechenskaya Prov., 1902, M. R., leg. Saposhnikov], "Myrmica dshun­
garica Ruz. w" (both labels written by Ruzsky's own hand), "Lectotypus, design. K. Arnoldi"
(MOSCOW); paralectotype: 1 w "Семирсчснская обл., 1902, M. P, Кожевников" [Semirechenskaya
Prov., 1902, M. R., leg. Kozhevnikov], "M. dshungarica M. Ruzskij d e t " , "Myrmica dshungarica Ruz.
w" (all three labels written by Ruzsky's own hand), "Lectotypus Myrmica dshungarica Ruz. design.
Arnoldi" (see Notes below) (MOSCOW); 1 w, "Myrmica dshungarica Ruz. w Semiretschinsk, Alatau,
M. R." (written by Ruzsky's own hand) (GENOA); lectotype of Myrmica laevinodis var. minuta, w
(designated by Arnoldi 1976), "Андсманын, VI. 95, Богоявленский" [Andermanyn, vi.95, leg. Bogoy­
avlensky] (both labels written by Ruzsky's own hand), "^Myrmica laevinodis var. minuta", "Lectoty­
pus, design. K. Arnoldi" (MOSCOW); syntypes of Leptothorax svartshevskii: 19 w, 4 q, "Киргиз-ата,
Фсрганск., Сварчсвский" [Kirgiz-ata, Fergana Prov., leg. Svarchevsky], "4241 Coll. Karawajewi",

124
"Leptothorax svartzevskii [Sic\] Karaw. typus" (KIEV, MOSCOW); syntypes of Myrmica rugulosa
var. ruginodiformis: 8 w, 2 q, 9 m, "Каркара, Кузнецов" [Karkara, leg. Kuznetsov], "4003 Coli. Karawa­
jewi", "Myrmica rugulosa ruginodiformis Karawajew typus", "Syntypus Myrmica rugulosa ruginodi­
formis Karaw." (KIEV, MOSCOW); holotype of Myrmica minuta subsp. iskanderi,vi, ""Iskander-Kul,
16.VIII.36, O. Kryzhanovkij", "HolotypeMyrmica minuta subsp. iskanderi K. Arnoldi" (MOSCOW);
paratypes: 1 w, 4 m, "Джиргиталь, Таджик., 11.VIII.36, О. Крыжановский" [DzhirgitaP, Tajikistan,
ll.viii.36, O. Kryzhanovsky]; 1 w, "Кашка-су, 30 км W Сарыташ, Алайская долина, 63-341, VI.1963,
Г. Длусский" [Kashka-su, 30 km W Sarytash, Alai valley, 63-341, vi.1963, G. Dlussky]; 6 w, "Iskan­
der-Kul, 16.viii.36, О. Kryzhanovskii"; 2 w, "No. 6221"; 1 w, 1 m (male without head), "No. 6222";
2 w, "No. 6224"; 9 w, "No. 6225" (MOSCOW); non-type material: about 50 w, 7 q and 8 m from
Middle Asian Mts.
Distribution (Map 29). Mountain ranges of Dzhungarian Alatau, Tien-Shan and
Pamiro-Alaj on the territory of Kazakhstan, Kyrgyzstan, Uzbekistan and Tajikistan.
Etymology. M. dshungarica: named from Dshungaria with the adjectival suffix for nouns
ica (from the Greek ¡ко) = belonging to, to indicate that the type specimens were taken in
the Dshungarian Alatau (mountains) of eastern Kazakhstan. Modern Dshungaria (Dzhun-
garia) is a region lying mostly within Xinjiang province of northeast China and extending
into western Mongolia and eastern Kazakhstan. M. minuta: from the Latin word (F.) mi­
nuta = small, to indicate its small size. Leptothorax svartshevskii: this species most probab­
ly was dedicated to Prof. Boris Alexandrovich Svarchevsky (1872-1930), the well-known
Russian freshwater zoologist, one of the founders of the Irkutsk University. M. ruginodi­
formis: name a combination of the Latin words ruga = wrinkle, nodus = knot or lump and
the suffix formis = to be in the shape of, or form, to describe its petiole node. M. iskanderi:
named after Iskander Kul (lake) in Tajikistan, the type locality, which itself was dedicated
to Iskander (Alexander the Great). M. tarbinskii: dedicated to the well known Soviet
myrmecologist Prof. Yury Serafimovich Tarbinsky (1937-2003), who collected the type ma­
terial. Tarbinsky published "The ants of Kirgizia" in 1976 and was the Head of the Ento­
mological Department of the Institute of Zoology in Frunze (now Bishkek, Kirgizia).
Notes. Arnoldi (1976) placed M. dshungarica to the "rubra-smythiesii" species group,
and Radchenko (1994b) - to the rabra-group. Now we consider that this species represents
a different group - the dshungarica-group (see also Radchenko and Elmes 2001b). It is
similar to, and often hard to distinguish from several sympatric Myrmica species, e.g.
M. juglandeti and M. ferganensis, but differs from them by its shorter and somewhat higher
petiole, distinctly convex side of head, etc.
The main diagnostic features of M. dshungarica are quite variable compared to many
Myrmica species and this no doubt provoked different authors to describe the quite large
number of infraspecific forms. Direct comparison of the types of all taxa now attributed to
this species (see above) shows that although there are minor differences, the range of
characters (including morphometries) overlap so we found it impossible to clearly define
even a pair of "sister" species. Moreover, the males where they are known for the "sub­
species", are practically identical. Therefore, we believe that the earlier decisions to
synonymise all the names mentioned above under M. dshungarica are correct. Thus,
M. dshungarica appears to be a widely distributed species in the mountains of Middle Asia
that is fragmented into many isolated populations. Commonly in such mountain regions
isolated populations evolve small morphological variations, in the case of M. dshungarica
these are not yet sufficiently different to consider any population as separate subspecies.
We found in MOSCOW 2 workers on separate pins that were determined by Ruzsky as
M. dshungarica. Each pin has one of Arnoldi's labels saying "Lectotypus, design. Arnoldi",

125
but he never published any data concerning a lectotype of M. dshungarica. Equally con-
fusingly the specimens have the same collection data except for a different collector's name:
Kozhevnikov instead Saposhnikov. We can not explain this discrepancy; possibly it is just
a printing error but on the other hand, Prof. Kozhevnikov also collected material in Middle
Asia, maybe even working together with Prof. Saposhnikov, and could equally well have
provided materia! for Ruzsky. Another specimen had one of Ruzsky's labels reading "Semi-
rechenskaya Prov., 1902, M. R., leg. Saposhnikov"; as this fully corresponds with the de-
scription of M. dshungarica (see above) we designate this specimen as the lectotype. We
designate the second specimen that has Ruzsky's label "Semirechenskaya Prov., 1902,
M. R., leg. Kozhevnikov", as a paralectotype; recently we found one more paralectotype
worker in Emery's collection (GENOVA).
In the original paper describing M. minuta subsp. iskanderi (Arnoldi 1976: 549), under
the section on characteristics of the material and type localities, the printed symbol for
"worker" was misprinted as "female"; in fact, K. Arnoldi did not described queens of this
form and the holotype is a worker.
G. Wheeler and ]. Wheeler (1953) described larvae of M. dshungarica (noted by Bolton
1995). However, Wheeler and Wheeler wrote that they studied "six larvae from Siberia"
(loc. cit., p. 123). If this locality is correct, then those larvae could not beM. dshungarica
but could be one of any Siberian species from the lobicornis-group.
Ruzsky (1905) recorded for Dzhungarsky Alatau M. smythiesii. It is now known that
this species is restricted to the Himalayan region and almost certainly Ruzsky's old record
is a misidentification. Kuznetsov-ugamsky (1927: 192, 193) made a study of M. dshunga-
rica and he was also of the opinion that Ruzsky's record was a misidentification. Addi-
tionally, he described four infrasubspecific forms of M. dshungarica: var. dentata (m), var.
subacuta (m), var. infuscata (w), and var. brevispina (w); ali this material seems lost.
Tarbinsky (1976) and Radchenko (1994e) referred this material to M. dshungarica, but
according to the fourth Edition of the International Code of Zoological Nomenclature
(1999) these names are unavailable (quadrinomens).

Map 29. Distribution of M. dshungarica.

126
 Ecology. M. dshungarica predominately inhabits the mountain fir-forest belt at altitudes
1600-3000 m a.s.l., or even up to 3600 m a.s.l. (in Alaj and Pamir), where it is quite
common. It seems to be an hygrophilous species, living mainly in quite wet or even damp
places near rivers and streams, and often penetrating along river valleys at lower altitudes
below the fir-forest vegetation belt. In subalpine meadows, where environmental conditions
are generally more humid than in fir forests, it lives far from water, nesting in soil under
stones. Alate gynes and males emerged in July-August, nuptial flight - in August-
September.

Myrmica eidmanni Menozzi, 1930

(Figs 75-77)

Myrmica eidmanni Menozzi, 1930: 331, w, Russia (Buryatia); Radchenko 1994f: 91, 2005: 139; Pfeif­
fer et al. 2007: 4.
Myrmica scabrinodis subsp. eidmanni: Weber 1950: 204; Bolton 1995: 278.
Myrmica jessensis: Ruzsky 1925: 46; Kupyanskaya 1979: 128, 1986 : 85, w, q, m, 1990: 110; Rad­
chenko 1994b: 43, 1994c: 142, 1994f: 88 (misprinted as iessensis); 1994g: 109; Bolton 1995: 279
{part., material from continental Asia); Wu and Wang 1995: 94; Kim B.-J. 1996: 180 (part.?); Wei
С et al. 2001: 561 (part.); Elmes et al. 2001: 110, misidentifications.
Myrmica lobicornissubsp.littoralis Weber, 1948: 287 (first available use oiMyrmica scabrinodis subsp.
lobicornis var. littoralis Kuznetsov-ugamsky, 1928: 33, w, q, m, Russian Far East). Synonymy by
Kupyanskaya 1986a: 85; Radchenko 1994f: 88; Imai et al. 2003: 183 (as synonym of M. jessensis);
Radchenko 2005: 139 (as synonym of M. eidmanni); the latter confirmed here.

Type localities and type specimens. M. eidmanni: "Parecchie operane di Verkneudynsk (Siberia)".
Myrmica lobicornis subsp. littoralis: "Океанская; мыс Басаргин; Тигровая; ст. Уссури" [Okeanskaya;
cape Basargin; Tigrovaya; station Ussuri].
Material examined. Lectotype of M. eidmanni, w (designated by Radchenko 2005) , "Verkhneu-
dinsk, Sibirien" [now- Ulan-Ude, Buryatia, Russia], "Typus", "Myrmica eidmanni Menozzi, typus"
(BOLOGNA); non-type material: > 100 w, > 30 q and m from South Siberia, Mongolia, Russian Far
East, NE China and Korean Peninsula.
Distribution (Map 30). South Siberia (from Altai Mt. in the west), Mongolia, Russian
Far East, Korean Peninsula, NE China.
Etymology: M. eidmanni: dedicated to Mennozzi's friend, the entomologist and expe­
ditionist Prof. Hermann A. Eidmann (1897-1949) of the University of Göttingen.
M. littoralis: from the Latin word littoralis (f.) = of the sea shore; one of the type localities
(cape Basargin) is near to the ocean.
Notes. M. eidmanni was an enigmatic taxon for more than 60 years after its descrip­
tion. Nobody seemed to have examined the type material in BOLOGNA and only Weber
(1950) and Bolton (1995) recorded this name in catalogues (as a subspecies of M. scabri­
nodis). Radchenko (1994f) considered M. eidmanni as incertae sedis in the lobicornis
species group, supposing that it could be a junior synonym of M. angulinodis. Based on the
investigation of the type specimens of this species, we concluded that M. eidmanni is a good
species distributed from Altai Mts. to the Pacific Ocean that previously was determined by
most Russian entomologists as M. jessensis (see Radchenko 2005). It is closely related to
M. jessensis, which is mainly distributed in Japan, but clearly differs from the latter by its
wider frons and less extended frontal lobes (see Notes to M. jessensis).

ill
 Map 30. Distribution of M eidmanni.

Kuznetsov-ugamsky (1928) described var. littoralis based on workers, queens and males
from the southern part of the Russian Far East. We can not find the types of this form and
they most probably are lost. However, based on the original description, it is quite obvious
that this name is a synonym of M. eidmanni. Kuznetsov-ugamsky {loc. cit., p. 35) also
noted: "described form, most probably, closely related to M. se. lobicornis var. jessensis".
As the first available use of this name is Myrmica lobicornis subsp. littoralis Weber, 1948,
the name M. eidmanni has priority.
Ecology. Despite M. eidmanni is widely distributed in the Taiga Zone, it avoids dense
forests and is more common at relatively low altitudes in anthropogenic environments
(the highest altitude recorded is 2100 m a.s.l.). Its preferred habitats are grasslands, forest
clearings, roadsides, riverbanks, sandy lakeshores and peat bogs but it lives also in sparse
forests (pine, larch) and their edges and glades, where it builds nests in the soil, often under
stones, and in soil and moss mounds (Dmitrienko, Petrenko 1976; Kupyanskaya 1990).
Colonies are moderately polygynous (ranging from 1-5 queens) and contain on average 500
workers. Sexual forms emerge in July-August, nuptial flight - in August-September.

Myrmica emeryi R a d c h e n k o et Elmes, 2 0 0 9

(Fig. 78)

Myrmica emeryi Radchenko et Elmes, 2009c: 3, w, Indonesia.

Myrmica ritae: Emery 1895b: 451 (misidentification).

Type localities and type specimens. "Pulo Laut", for details see material examined and Notes.
Material examined. Holotype, w, "Pulo Laut" (middle specimen on the pin with 3 workers)
(GENOA); paratypes: 2 w, on the pin with holotype; 1 w, "Pulo Laut" (GENOA).
Distribution (Map 31). It is known only from the type locality, Isl. Pulo Laut, SE of
Borneo - the only known location for a Myrmica species from the Southern Hemisphere.

128
 Etymology. M. emery: dedicated to the great myrmecologist Prof. Carlo Emery (1848-
1925) (see biography in Chapter 1.6), in whose collection we found this species.
Notes. M. emeryi clearly belongs to the ritae-complex of the ritae species group (see
Radchenko and Elmes 1998, 2001a, b; Radchenko et al. 2001, 2006), it most resembles
M. margaritae, M. pulchella, and M. sinensis, which are characterised by a very coarse ru-
gosity of the head (frons between frontal carinae level with the eyes with only four coarse
longitudinal rugae). It is hot so surprising that Emery attributed it to M. ritae because until
relatively recently the ritae-group was poorly understood due to the shortage of material
in collections. M. emeryi has abundant long standing hairs on the alitrunk dorsum and
occipital margin of the head, a feature that well differentiates it from M. margaritae, which
has only a few standing hairs on these areas. On the other hand, it differs from M. pulchella
and M. sinensis by the much less abundant standing hairs on the lateral margins of the head
(less than five versus more than 20). Additionally, M. pulchella, unlike M. emeryi, has no
reticulation on the head dorsum and alitrunk dorsum. The absence of reticulation on the
petiolar and postpetiolar dorsum of M. emeryi clearly distinguishes it from M. sinensis,
which has coarse reticulation on these surfaces.
The type location is fascinating. The pins with the holotype and paratype specimens
both have labels that read "Pulo Laut" ("pula" means "island" in Malaysian), but give no

Map 31. Distribution of M. emeryi.

129
other collection information. When we described this species we wrote " We ... found
several localities with the same or similar spelling. All of them are islands situated near
Malaysia or Borneo. If any of these localities is correct, then M. emeryi is the most geo-
graphically isolated and has the most southern distribution of all known Myrmica species"
(Radchenko and Elmes 2009c). Since then we discovered that Emery (1895b: 451) wrote
"Le Myrmica che a primo aspetto, potevano sembrare un elemento nordico non hanno
questo significato, la Ritae essendo stata ritrovata a Borneo (Pulo Laut) da Doherty" (The
Myrmica [of Burma] that at first sight might seem to be a Nordic element [in that fauna]
does not have this interpretation because the ritae have been found again in Borneo (Pulo
Laut) by Doherty).
William Doherty (1857-1901) was an American zoologist and collector active in the
late 19th century collecting birds and invertebrates (especially butterflies) in the "East In-
dies". Numerous ant species were collected by him and most were described by Emery, who
named three species after him. The type locality for many of Doherty's collections is given
as Pulo Laut (off the southeast of Borneo). Based on material from other groups it seems
most probable that the M. emeryi specimens were taken in June 1891 by Doherty some-
where on Pulo Laut, near Borneo. These specimens show that r/iae-group species can live
south of the equator. Quite possibly, when the ant fauna of isolated high mountains of Bor-
neo and elsewhere in Malaysia and Indonesia are better known, more ritae-group species
and possibly other unusual forms of Myrmica will be found.
Queens, males and ecology are unknown.

Myrmica ereptrix Bolton, 1988

(Fig. 79)

Myrmica ereptrix Bolton, 1988: 2, q, India; 1995: 279; Radchenko and Elmes 2001b: 250, 2003a: 236.

Type localities and type specimens. "Holotype female, India: Kashmir, Gulmarg, 20.vii.1986, 2800
m, Picea forest (P. H. Williams)".
Material examined. Holotype, q, "Kashmir, Gulmarg, 20.vii. 1986, 2800 m, leg. Williams" (LON-
DON).
Distribution (Map 32). The species is known until now only from the single queen, col-
lected in Kashmir.
Etymology. M. ereptrix: from the feminine form of the Latin word ereptor = thief, to
indicate that this species is almost certainly a social parasite.
Notes. M. ereptrix is a socially-parasitic species and at the time of writing, is known only
from the single holotype queen. It has very conspicuous morphology, particularly the ex-
tremely wide petiole and postpetiole, both with well-developed, large ventral lobes. Bolton
(1988) gave M. rugosa as the host species, but we identified the host as
M. aimonissabaudiae (Radchenko and Elmes 2001b).
M. ereptrix is quite distinctive from all other known Palaearctic Myrmica social parasites.
This is not surprising if it follows Emery's rule, because the Himalayan Myrmica fauna dif-
fers from all other Palaearctic Myrmica with almost the entire fauna being endemic to the
region (Radchenko and Elmes 2001b, 2003a).

130
 Map 32. Distribution of M. ereptrix.

Myrmica excelsa Kupyanskaya, 1990

(Figs 80-82)

Myrmica excelsa Kupyanskaya, 1990: 115, w, q, m, Russia; Radchenko 1994b: 43, 1994c: 144, 1994f:
90; Bolton 1995: 279; Elmes et al. 2002: 167; Radchenko 2005: 140; Yamane 2008: 29.
Myrmica cadusa Kim B.-J., Park et Kim J.-H., 1997: 425, w, South Korea. Synonymy by Radchenko
2005: 140; confirmed here.
Myrmica sinica Wu and Wang, 1995: 94, 197, w, q, China; Wei С et al. 2001: 561. Synonymy by
Radchenko et al. 2008: 783; confirmed here.
Myrmica kasczenkoi: Collingwood 1976: 302, misidentification.
Myrmica sp. 6: Imai et al. 2003: 181.

Type localities and type specimens. M. excelsa: "Голотип - рабочий, Приморский край, Барабаш-
Лсвада, 9.V 1972 (Купянская); иаратипы: 36 рабочих с такой же этикеткой; Кедровая Падь, 30.VIII 1974,
14.IX 1973, 9.1.1974, 16.VI 1975, 9.VII 1979 (Купянская), 94 рабочих, 5 самок и 8 самцов; Анисимовка,
20.VIII 1979 (Купянская), 32 рабочих; Евсссвка, 22, 23.VI 1978 (Купянская), 23 рабочих; Лазовский
заповедник, 22.IX 1974 (Купянская), 6 рабочих, 1 самка; Комиссаровка, 7.V 1972 (Купянская), 20 рабочих;
Рудная Пристань, 7.1.1972, 90 рабочих; хр. Чандолаз, 26.1 1974 (Сторожсва), 12 самок" [Holotype- w,
Primorsky Krai, Barabash-Levada, 9.V.1972, leg. Kupyanskaya; paratypes: 36 w with the same label;
Kedrovaya Pad', 30.vui.1974, 14.ix.1973, 9.ix.1974, 16.vi.1975, 9.VÜ.1979, leg. Kupyanskaya, 94 w, 5
q', 8 m; Anisimovka, 20.viii.1979, leg. Kupyanskaya, 32 w; Evsecvka, 22, 23.vi. 1978, leg. Kupyanskaya,
23 w; Lazovsky Natural Reserve, 22.ix.1974, leg. Kupyanskaya, 6 w, 1 q; Komissarovka, 7.V.1972, leg.
Kupyanskaya, 20 w; Rudnaya Pristan', 7.ix.1972, 90 w; Chandolaz Range, 26.ix.1974, leg. Storozheva,
12 q]. M. cadusa: "Holotype: worker, Mt. Moak, Chonbuk, 24 V 1995 (Kim JH and SJ Park). Paratype:
20 workers, Mt. Moak, Chonbuk, 24 V 1995 (Kim JH and SJ Park)". M. sinica: "Holotype: worker,
Yantai City of Shandong Province, Huo Yulin leg.; paratypes: 27 workers, 2 queens in the same nest
as the holotype".
Material examined. Holotype of M. excelsa, w, "Прим. край, Барабаш-Лсвада, 9.V.72, Купянская"
[Primorsky Krai, Barabash-Levada, 9.V.1972, leg. Kupyanskaya], " H ol oty pu s My rm/CÛ excelsa Kupyan-
skaya" (MOSCOW); paratypes: 6 w, 3 q, 5 m, "Прим. край, Кедровая Падь, 9.IX.74, Купянская"

131
[Primorsky Krai, Kedrovaya Pad', 9.ix.74, log. Kupyanskaya]; 9 w, "Прим. край, Анисимовка, 20.VI1I
1979 (Купянская)" [Primorsky Krai, Anisimovka, 20.viii.79, leg. Kupyanskaya] (MOSCOW, VLADI­
VOSTOK, KIEV); holotype of M. cadusa, w, "Korea, Mt. Moak, Ch nbuk, 24 V 1995, Kim and Park"
(IKSAN); paratypes: 2 wwith the same label (IKSAN); paratype of M. sinica, w, "Yantai City, Shan­
dong Province, China, date unknown, leg. Yulin Huo", "Myrmica sinica Wu et Wang (Chinese Acad.
Forestry)" (BEIJING); non-type material: several tens of w, about 20 q and m from Russian Far East,
Korean Peninsula and Japan.
Distribution (Map 33). Southern part of Russian Far East, NE China, Korean Peninsula,
Japan.
Etymology. M. excelsa: from the Latin word excelsa = elevated or tall, to describe the
elevated part of the clypeus in front of the antennal insertions (see Notes below). M. ca­
dusa: from the ancient Latin word cadus = barrel, to describe the "jar-like" antennal pit,
resulting from the elevated clypeal margin. M. sinica: Sinica means Chinese, derived from
Sina, an old Persian word for China.
Notes. M. excelsa is a distinctive species belonging to the lobicornis-group. Its most
characteristic feature is the raised lateral portions of clypeus that form a sharp ridge in front
of the antennal insertions, so that antennal sockets are distinctly separated from the clypeal
surface (as in species of Tetramorium). This led Kim et al. (1997) to describe the antennal
pits as "jar-like". Only two more Myrmica species are known to possess this feature:
M. transsibirica and M. tamarae, both are lobicornis-group species. M. excelsa well differs
from both by the shape of its antennal scape (for details see Kupyanskaya 1990; Radchenko
2005, and Key, Chapter 5.3), but due to the Tetramorium-Uke clypeus we place them to­
gether in the excelsa-complex.
When we compared the type specimens of M. cadusa and M. sinica with those of
M. excelsa we had no doubts that they were conspecific. We also had an opportunity to in­
vestigate specimens of this species from Japan, kindly sent us by Prof. K. Onoyama; they
were originally determined as Myrmica sp. 6 by Imai et al. (2003).

Map 33. Distribution of M. excelsa.

132
 Collingwood (1976) misidentified specimens of M. excelsa from North Korea as
M. kasczenkoi; this error was repeated by B.-J. Kim (1996).
Ecology. M. excelsa is a rather thermophilous species. In the Russian Far East it inhabits
fir and mixed forests of the southern type, that support a considerable number of rare and
relict taxa. In Korea, where the climate is definitely warmer and drier than in the Far East
of Russia, M. excelsa was found at a low and middle altitudes (100-650 m a . s i ) , preferring
deciduous and mixed forests (oak, beech, sweet chestnut, pine) and shrubby areas but usu­
ally avoiding open grasslands. In these forests workers appeared to forage close to the soil
surface under litter and dead leaves and were very cryptic, even for lobicornis-group species
(personal observations). In Japan M. excelsa is rather a rare species living in grasslands at
altitudes between 500 and 1500 m a.s.l. In Japan and Korea nests are usually built in the
soil, often under stones, but in the Russian Far East nests are mostly in dead wood. Nup­
tial flight - in August-September (see also Kupyanskaya 1990; Imai et al. 2003; Radchenko
2005).

Myrmica ferganensis Finzi, 1926

(Fig. 83)

Myrmica laevinodis var. ferganensis Finzi, 1926: 84 (first available use of Myrmica rubra subsp. laevi­
nodis var. ferganensis Karawajew, 1916: 503, w, Kyrgyzstan; Kuznetsov-ugamsky 1927: 193).
Myrmica ferganensis: Arnoldi 1976: 550; Radchenko 1994b: 41, 1994c: 137, 1994e: 124; Bolton 1995:
279.
Myrmica tenuispina Ruzsky, 1905: 670 (part, material from Tash-Kurgan and Kugart; see Notes to
M. tenuispina, below).
Junior synonym of M. rubra var. ruginodolaevinodis: Sadil 1952: 241; not confirmed, see Notes to
M. ruginodis, below.
, Junior synonym of M. rubra: Tarbinsky 1976: 40; not confirmed, see Notes to M. rubra, below.

Type localities and type specimens. "P. Киргиз-ата, около 9000 фут., Ферганской обл., лето 1914 г.,
Б. Сварчсвский. Под камнем у берега реки. Много рабочих" [Riv. Kirgiz-ata, about 9000 feet, Fergana
Prov., summer 1914, leg. B. Svarchevsky. Under stone on river bank. Many workers].
Material examined. Syntypes, 6 w, "Киргиз-ата, 9000', Ферг. обл., Б. Сварчсвский" [Kirgiz-ata, 9000',
Fergana Prov., В. Svarchevsky], "5262 Coll. Karawajewi", "Myrmica rubra laevinodis var. ferganen­
sis Karaw." (KIEV); 2 w with the same labels (MOSCOW); non-type material: 18 w from Kazakhstan,
Kyrgyzstan and Tajikistan.
Distribution (Map 34). Tien-Shan, Alai, Pamir.
Etymology. M. ferganensis: from the name Fergana with the Latin suffix ensis = place
of origin, to indicate that the types were taken in Fergana province of Kyrgyzstan.
Notes. Radchenko (1994a) placed M. ferganensis to the rubra species group, but now
we believe it belongs to the related, but separated dshungarica-group. It is similar to
M. juglandeti, but differs from it by the weakly developed striae surrounding antennal sock­
ets (often they are completely absent).
For a long time M. ferganensis was confounded with M. tenuispina because when Forel
designated the type series of M. tenuispina besides the "true" M. tenuispina specimens he
erroneously included in it two series of M. ferganensis: 13 w, Tash-Kurgan, Darvaz, E
Bukhara, 22.viii.97, leg. Kaznakov (MOSCOW, PETERSBURG) and 6 w, Kugart, 6-8000',
Fergana obi., leg. Korzhinsky, 5.viii.95 (PETERSBURG). These specimens well differs from

133
the lectotype and paralectotypes of M. tenuispina by their less extended frontal lobes (for
more details see Notes to M. tenuispina, below).
Ecology is poorly known. M. ferganensis mainly inhabits mountain meadows, river
banks at altitudes over 2000 m a.s.l., where it builds nests in the soil, often under stones.

Map 34. Distribution of M. ferganensis.

Myrmica forcipata Karawajew, 1931

(Figs 84-86)

Myrmica forcipata Karawajew, 1931 b: 105, w, Russia; Weber 1950: 211; Pisarski 1969a: 228, 1969b:
295; Pisarski and Krzysztofiak 1981: 155; Radchenko 1994b: 43, 1994c: 142, 1994f: 88, q, m, 1994g:
107; Bolton 1995: 279; Pfeiffer et al. 2007: 5, nec Kupyanskaya 1990: 107, misidentification.

Type localities and type specimens. "Fluss Tshona, Distr. Viljuisk, 27.VII. bis 2.VIII. 1914, V. Dro-
bov, 2 w. - Tyilyminskij Nasleg, 2.VII.1925, A. Ivanov, 2 w".
Material examined. Syntypes, 2 w, "Тыылыминск. наслег, Якут, окр., 25.VII.25, Иванов", "Якутск.
Экспсд. АН" ["Tyylyminsky nasleg, Yakutsky okrug, 25.vii.25, leg. Ivanov", "Yakutskaya expedition of
Ac. Sci."], "Myrmica forcipata Kar. V. Karawajew det.", "Syntypus Myrmica forcipata Karaw."
(KIEV); non-type material: > 200 w, several tens of q and m from the whole area.
Distribution (Map 35). South and East Siberia (to the west until Sayany Mts.), Mon­
golia, absent in the Russian Far East.
Etymology. M. forcipata: the name derived from the Latin word forceps = pincher or
tongs, to describe its forceps-like propodeal spines.
Notes. M. forcipata belongs to the lobicornis species group. Specimens are characterized
by the blunt, thick and inwardly curved propodeal spines, and by a sharply angled petiolar
node. By these features, it is similar to M. angulinodis, but differs from the latter by the
distinct, though not large, vertical lobe at the scape bend.

134
 Map 35. Distribution of M. forcipata.

Kupyanskaya's (1990) record of this species for the Russian Far East in fact belongs to
M. angulinodis.
Ecology. In the ELMES collection there are series from > 80 nests of M. forcipata col-
lected by Michal Woyciechowski from northern Mongolia at about 1000 m a.s.l. (1999-
2001, see also ecological data for M. angulinodis). About 48% of nest series were taken in
herb-rich and wet grassy meadows near rivers, where most nests were built in grass clumps
with some directly in the soil or under pieces of rotten wood; 22% were found in the other
wettish habitats, shrubland and riparian woodland, where nests were most likely to be found
in grass and moss tussocks. A further 28% of series came from drier meadow-steppe, where
nests were mostly built directly in the soil. Only 2 nests of M. forcipata were found in moun-
tain-forest and mountain dry steppe (for description of biotopes see Muehlenberg et al.
2000). Sexuais were found in nests from July to August and mating swarms were observed
on a hilltop (1040 m. a.s.l.) on 12th and 15th August 1999, mixed with M. angulinodis.
Despite M. forcipata and M. angulinodis being morphologically quite similar (see above)
and often living sympatrically and sharing mating swarms, this pair of species appear to be
quite well-separated ecologically. In Mongolia M. angulinodis were clearly associated
(> 80% of series) with mountain-forest and its ecotones, with montain-steppe and herb-
meadows, less than 5% of M. forcipata were taken in these habitats. It appears to be a much
more hygrophilous species, preferring the grassy habitats near rivers.

Myrmica foreliana Radchenko et Elmes, 2001

(Fig. 87)

Myrmica foreliana Radchenko et Elmes, 2001b: 247 (replacement name for Myrmica smythiesii
r. carbonaria Forel, 1902: 227).
Myrmica smythiesi r. carbonaria Forel, 1902: 227, w, India [junior primary homonym of Myrmica
carbonaria Smith F., 1858: 127 (transferred to Monomorium Mayr)]; 1903: 697; Bingham 1903: 269;

135
 Ruzsky 1905: 661; Emery 1921: 41; Weber 1947: 458; Chapman and Capco 1951: 130; Bolton 1995:
278.

Type localities and type specimens. "Un seul exemplaire de Pachmarhi (Schurr)".
Material examined. Holotype, w, "Himalaya, Pachmarhi, Shurr, 30.vii.93 (Rothney)" (GENEVA).
Distribution (Map 36). It is known only from the type locality in Central Himalaya.
Etymology. M. foreliana: dedicated to Auguste Forel, the famous Swiss myrmecologist
(see biography in Chapter 1.6), to indicate this species was originally described using
a homonym, by Auguste Forel. M. carbonaria: from the Latin word carbo = carbon, to de-
scribe its dark charcoal-like colour.
Notes. M. foreliana belongs to the rugosa species group and is most similar to M. ru-
pestris. This species was described only from a single worker, we found no more examples
in any of the collections examined. The holotype specimen clearly differs from M. rupestris
by its smaller size (HW 0.77, AL 1.46 mm, vs. HW > 0.78, AL > 1.80 mm), somewhat
longer head (CI 1.39 vs. CI < 1.35) and by the surfaces between rugae on its head and
alitrunk being densely punctated and appearing more or less dull. However, additional
material is required before a clearer treatment of this species can be made. Queens, males
and the ecology are unknown.

Map 36. Distribution of M. foreliana.

Myrmica fortior Forel, 1904

(Figs 88-90)

Myrmica smylhìesii var. fortior Forel, 1904b: 22 {part., nec material from Russian Far East, misiden-
tification); Ruzsky 1905: 660 (part., nec material from Russian Far East, misidentification);'Forel
1906: 87; Emery 1908a: 168, 1921: 41 (part., nec material from Russian Far East, misidentifica-
tion); Menozzi 1939: 295; Weber 1947: 460; Chapman and Capco 1951: 130; Bolton 1995: 279.
Myrmica fortior: Radchenko and Elmes 2001b: 254, 2002: 39, q, m.

136
 Type localities and type specimens. "Sind Valley, 1981™, Cachemire (Wroughton); Ussuri méri-
dional, Sibérie orientale (Musée de S'-Pétersbourg) [sec Notes below]; Deota, 914m, Himalaya N.O.
(Smythies)".
Material examined. Lectotype, w (designated by Radchenko and Elmes 2001b), "India, Sind Val-
ley, 6500 ft, Kashmir (Wroughton), var. fortior Forel" (GENEVA); paralectotypes: 2 w with the same
labels as lectotype (GENEVA); 1 w, "Smythiesi var. fortior Sind Valley" (GENOA); non-type material:
> 50 w, 6 q, 4 m from India (Kashmir).
Distribution (Map 37). Until now it is known only from Kashmir State in India.
Etymology. M. fortior: from the Latin word fortior = stronger, to describe its greater ro-
bustness compared to M. smythiesii.
Notes. We place M. fortior in the smythiesii species group. It is probably a Kashmir
endemic very close to M. smythiesii, from which it differs by a suite of minor characters
including the shape and sculpture of the petiole and postpetiole, slightly longer propodeal
spines and somewhat larger body size (about 5% larger).
M. fortior workers differ from those of гаЬгя-group species by having a distinctly more
prominent and anteriorly pointed clypeus. However, by other characters they are superfi­
cially similar to those of several oriental species, related to M. ruginodis, which could ex­
plain the mistake made by Forel (1904b) when he referred material from the Russian Far
East to M. fortior. We are sure that M. fortior is restricted to the southern Himalaya and that
Forel's specimens must be M. ruginodis or related species. This error was repeated by
Ruzsky (1905) and Emery (1908a, 1921). Also, despite having quite long antennae, the
male scape is relatively shorter than that of males of oriental rabra-group species (SI2 0.83-
1.0 versus 1.23-1.36).
Ecologyis poorly known. It has been found at altitudes between 2000 and 2700 m a.s.l.,
where it nested in the soil and under stones, in open pine wood and open rocky pasture, at
the edge of spruce-fir groves (Phil Ward, pers. comm.).

Map 37. Distribution of M. fortior.

137
 Myrmica gallienii Bondroit, 1920
(Figs 91-93)

Myrmica gallienii Bondroit, 1920a: 150, w, France (also described as a new by Bondroit, 1920b: 302);
Finzi 1926: 90 (misspelled as gallieni).
Junior synonym of M. sulcinodis: Bernard 1967: 121.
Revived from synonymy: Collingwood 1978: 67; 1979: 48, q, m; Agosti and Collingwood 1987a: 53,
1987b: 268; Seifert 1988: 9; Atanassov, Dlussky 1992: 91; Radchenko 1994b: 42, 1994c: 143,
1994d: 76; Seifert 1994: 13; Bolton 1995: 279; Seifert 1996: 228; Radchenko et al. 1997: 487; Della
Santa 2000: 171; Czcchowski et al. 2002: 24; Radchenko et al. 2002: 418; Radchenko, Czechowska
andCzechowski2004:47; Seifert 2007: 203, nec Wei С et al. 2001: 561; Chang and He 2001: 26,
misidentifications.
Myrmica rugulosa limanica Arnoldi, 1934: 162, w, q, m, Ukraine; Karawajew 1934: 75; Weber 1948:
307. Synonymy by Collingwood 1979: 48; Seifert 1988: 9; Radchenko 1994d: 76; Bolton 1995:
280; confirmed here.
Myrmica limanica: Arnoldi 1968a: 1170; 1970: 1840.
Myrmica jacobsoni Kutter, 1963: 133, w, q, m, Latvia, Estonia; Kutter 1977: 65. Synonymy by Colling­
wood 1979: 48; Seifert 1988: 9; Radchenko 1994d: 76; Bolton 1995: 280; Della Santa 2000: 171;
confirmed here.
Myrmica limanica subsp. jacobsoni: Arnoldi 1970: 1840; Pisarski 1975: 11.
Myrmica limanica subsp. obensis Arnoldi, 1970: 1840, w, m, Russia; Bolton 1995: 281. Synonymy by
Radchenko 1994d: 76; confirmed here.
Myrmica rolandi: Jacobson 1940: 145, misidentification.
Myrmica bergi: Sadil 1952: 244, misidentification.

Type localities and type specimens. M. gallienii: "Haute-Marne: Latrecy (Le François)". M. lima-
nica: "Ost Ukraine, Districkt Charkov, Mittellauf von Donctz". M. jacobsoni: "3 w, 4 q und 2 defekte
m. Ein weiteres m, 1 q und 3 w. Umgebung des Kanjersees am Meerbusen bei Riga, Strandwiesen und
feuchte Triften auf der Insel Oesel, ferner weitere nordcstländische Inseln, die Nordküste Estlands und
die Umgebung von Päinurme. Die Art scheint also im nördlichen Balticum weit verbreitet zu sein".
Material examined. Lectotype of M. gallienii, w (designated here), "Latrecy Ht. Marne", "Coll. R.
I. Sc. N. B. France", "type", "R. I. Sc. N. B. I. G. 21.400", "Myrmica gallienii Bondr. Type", "TYPE"
(BRUSSELS); paralectotype, w with the same labels as lectotype (BRUSSELS); lectotype of
M. limanica, w (upper specimen on the pin with 3 w, designated here), "1503, Zmiev, Kharkov, IX.26,
Бишкин [Bishkin], Arnoldi", "M. rugulosa limanica Arn., Arnoldi det., Holotyp." (see Notes below)
(MOSCOW); paralectotypes: 44 w, 10 q, 17 m, "Змиев, Харьк., бор, IX.29, (Арн.)" [Zmiev, Kharkov
Prov., pine forest, ix.29 (Arn.)], No. 4437, 4456, 4458, 4461, 4505, 4506, 4509, 5322, 5323, 5324,
5325 (MOSCOW); 5 w, 2 m, "Лиман, Змисвск. р. Арнольди" [Liman, Zmiev Distr. Arnoldi], "5468
Coll.'Karawajewi", "Myrmica rugulosa subsp. limanica Arnoldi typ.", "Paratypus Myrmica rugulosa
subsp. limanica Arnoldi"; 2 w, 1 m, "5323 Змиев, Лиман, IX. 1931, К. Арнольди" [Zmiev, Liman,
ix.1931, К. Arnoldi], "5625 Coll. Karawajewi", "Myrmica rugulosa subsp. limanica Arnoldi typ.",
"Paratypus Myrmica rugulosa subsp. limanica Arnoldi" (KIEV); syntypes of M. jacobsoni, 3 w, 1 q,
1 m, "Littland, 18.8.36, H. Jacobson" (MOSCOW); non-type material: > 200 w, > 50 q and m from
whole area.
Distribution (Map 38). Central and East Europe, southern Finland, Caucasus, West
Siberia (but not China, see Notes below).
Etymology. M. gallienii: almost certainly named for the controversial World War I
French General Joseph Gallieni (1849-1916) who won an early victory over the Germans
"that saved Paris". To some people, probably including Bondroit, who may have served un­
der him, Gallieni was the "Hero of the Marne" (type locality of the species). M. limanica:

l38
almost certainly this name derives from one of the type localities, the village of Liman, nr
Zmiev in the Kharkiv distr. of East Ukraine. The Ukrainian and Russian word liman (trans­
ferred by the Turks from the original Greek) means an estuarine lake (often freshwater)
and Liman village is situated close to a series of large lakes. Thus the name might be ap-
proprate to the ecological preference of this species. M. jacobsoni: dedicated to the collec­
tor, the famous Russian entomologist and coleopterist Georgiy Georgievich Jacobson
(1871-1926) of St-Petersburg, who worked in the Zoological Museum (now Zoological In­
stitute of the Russian Academy of Science). M. obensis: named for the Ob' river that flows
north through the West Siberian plain of Russia, combined with the Latin suffix ensis =
place of origin. M. chersonensis (unavailable name, see Notes below) : from the name of
Cherson (Kherson) with the Latin suffix ensis = place of origin, to indicate that the type
locality was the Kherson region of southern Ukraine. M. strandi (unavailable name, see
Notes below) : named for Prof. E. Strand - Arnoldi (1934: 165) wrote: "Diese Ameise ist zur
Ehre des hochverdienten Herrn Prof. Dr. E. Strand benannt".
Notes. Direct comparison of the types of all described species confirms earlier estab­
lished synonymy (see above). There have been two records of this species for China, but
these are misidentifications: we examined specimens identified as "M. gallienii" by Wei С
et al. (2001) and Chang and He (2001), and are certain that they are M. kurokii.
M. gallienii belongs to the bergi'-complex of the scabrinodis species group (see Rad­
chenko and Elmes 2004). Its antennal scape is smoothly bent at the base in an almost ideal
curve with no trace of a ridge or carina. It resembles sympatric M. bergi, female castes dif­
fering from the latter by their distinctly longer propodeal spines, lower postpetiole and
lighter body colour, while males well differ by their shorter scape. M. gallienii workers could
also be confused with those of M. rugulosa, but the scape of the latter is somewhat more
angularly curved at the base, and the head dorsum has much more developed reticulation.
Additionally, the second funicular segment of the males of M. gallienii is more than 1.5
times longer than the third one, while in M. rugulosa it is shorter, less than 1.5 longer than
the third one (see also Key, Chapter 5.1, 5.2).

Map 38. Distribution of M. gallienii.

139
 Arnoldi (1934) did not designate the holotype of M. rugulosa limanica; as a result, all
type specimens should be considered as syntypes. However, we found in MOSCOW the
specimen, labelled by Arnoldi as "holotype", and we designate this specimen as the lecto-
type. Also, Arnoldi (1970) referred to M. limanica the unavailable names M. rugulosa
limanica nat. chersonensis Arnoldi, 1934 and M. rugulosa limanica nat. strandi Arnoldi,
1934, both forms were described from the southern Ukraine. Radchenko (1994d) referred
them to M. gallienii; we examined "syntypes" of these forms, preserved in MOSCOW and
KIEV and confirm that undoubtedly these specimens are M. gallienii.
Bernard (1967) erroneously considered M. gallienii as a junior synonym of M. sul-
cinodis.
Ecology. M. gallienii is widespread in Central Europe, where it is associated with wet
meadows, lake and pond shores. In some habitats, particularly in Poland and probably Be-
larus, it can be the dominant species, but throughout most of its range colonies are not
common or at best they are restricted to very small areas of habitat. Colonies of M. gallienii
can easily be mistaken for those of M rubra in the field because in warm wet grassland both
have quite populous, polygynous colonies and often construct quite large earth solaria. Fur-
thermore, although these two ants are easily separated using a microscope, it is not so easy
for inexperienced myrmecologists armed only with a hand lens in the field. One of the best
"tips" is to observe the way in which workers walk: M. gallienii tend to walk slightly slower,
hold their antenna more at right-angles to their head and have a distinctly darker and square
head than M. rubra, which have more shiny heads and tend to hold their scape and antenna
more forward. Unfortunately, ecological data for these two species were sometimes con-
founded in the excellent ecological studies by Joanna Petal and co-workers (e.g. Petal 1967).
The distribution of M. gallienii in Poland and Germany was characterized by Czechowska
and Czechowski (1998) and by Munch and Engels (1994), who described how colonies
of this species live on floating reedbeds, this indicates that they might we well adapted to
flooding in much the same way as the related M. bergi.
Elmes and Pçtal (1990) gave colony population data for M. gallienii (under the syno-
nymised name M. limanica jacobsoni) from the Strzeleckie Meadows (wet abandoned
meadow) near Warsaw, Poland, sampled for 6 years. There were on average fewer workers
(640) than in a typical M. rubra colony and queens varied from year to year, annual means
ranging from > 5 to about 2 per colony, which is less than in a typical M. rubra nest
(see ecological notes on that species). Most interestingly the data indicated that during
the course of a year (the active season May-September) the number of queens in the
total population fell by 60% being replaced by newly recruited queens. Seppä (1996)
working on a different population showed that co-existing queens had very low relatedness
to each other, which is consistent with high mortality and general recruitment within
a population. However, over the 6 year period that the Strzeleckie Meadows were studied,
recruitment never quite matched queen loss, so that the average number of queens per
colony gradually declined. This was interpreted as indicating the gradual decreasing
suitability of the meadow to support the population as it gradually scrubbed-over, be-
coming increasingly more suitable for M. rubra, which also lived on the site. There have
been few laboratory studies of this species other than a brief report of its chemical secretions
(Jackson et al. 1989).
Sexuais are produced in early summer and nuptial flights and subsequent queen re-
cruitment takes place in August. It is interesting to note that in several Polish populations
we observed that a considerable number of gynandromorphs were present in many nests

.140
before nuptial flights. Usually these were mosaics with half, three-quarters or more of the
body being male. The only other species, where we have found such a high proportion of
such intercastes, is M. sulcinodis.

Myrmica georgica Seifert, 1987

(Fig. 94)

Myrmica georgica Seifert, 1987: 183, Georgia (w, nec m; only material from Georgia; see Notes below).
Junior synonym of M. turcica: Seifert 1988: 18.
Junior synonym of M. scabrinodis: Radchenko 1994: 79.
Revived from synonymy: Radchenko and Elmes 2004: 230.

Type localities and type specimens. "Workers: Holotype: 21.7.1985, leg. Seifert, east of Tiflis/Geor-
gia; 44°E, 41°43'N; preserved in Staaltlisches Museum für Naturkunde, Görlitz (SMNG). Paratypes: al-
together 96 specimens; 57 specimens from the same nest as holotype and 36 specimens from the same
site and date as holotype, but from several other nests, all leg. Seifert and preserved in SMNG" (see
also Notes below).
Material examined. Paratypes, 8 w (ELMES, KIEV) from the same nest series as the holotype
(which is preserved in GÖRLITZ).
Distribution (Map 39). Georgia.
Etymology. M. georgica: from the name Georgia with the adjective suffix for nouns ica
(from the Greek mo) = belonging-to or from, to indicate that the series with holotype were
found in Georgia.
Notes. Seifert (1987) described M. georgica based on several tens of workers that he
collected in Georgia, and on 3 workers and 2 males, collected in Northern Kazakhstan; one
year later he synonymised his species with M. turcica (Seifert 1988).

Map 39. Distribution of M. georgica.

141
 In 2004 we studied 8 paratype workers of M. georgica from the holotype nest series
that were kindly gifted to us by B. Seifert, and the 3 workers and 3 males paratypes from
Kazakhstan (MOSCOW- Щучинск, Кокчетавская обл., 18.viii.1966, № 40, Малоземова
[Shchuchinsk, Kokchetavskaya Prov., 18.viii.1966, No. 40, leg. Malozemova]). We were
able to compare them with the type specimens of M. turcica and the abundant samples that
had been recently collected by us in Turkey (several hundred workers and several tens of
males and queens of M. turcica). We concluded that Seifert's type series of M. georgica
included two species: the Georgian material belongs to one species and the Kazakhstan
specimens (MOSCOW) represented a second (undescribed) species that is distributed from
southern Ukraine to Kazakhstan. We suggested earlier (Radchenko and Elmes 2004) that
the Georgian material remain synonymised with M. turcica, while the Kazakhstan material
might be considered as M. georgica, and so revived the name M. georgica from synonymy.
However, Seifert pointed out that the holotype series must be considered to be M. georgica
and that the name must remain a junior synonym of M. turcica (Bernhard Seifert, pers.
comm.).
In this monograph therefore, we exclude the Kazakhstan "paratype" specimens
(MOSCOW) from the type series of M. georgica, and group them with several series of the
previously undescribed species from Ukraine, which we describe as M. kozakorum (see
below). This leaves the type series of M. georgica comprising the several nest series (in­
cluding the holotype series) that Seifert himself collected from Georgia. However based on
the paratypes from the holotype series in our collections, we can not agree with their
synonymy with M. turcica (Seifert 1988).
We found that the workers of M. georgica more resembles those of M. sabuleti, especially
by the size and shape of the lobe at the base of antennal scape, and by a relatively narrow
frons and more extended frontal lobes. In contrast, the antennal lobe of workers of
M. turcica is generally relatively small and is more similar to that of M. scabrinodis or even
M. specioides (see also Notes for M. turcica). However, we had the benefit of examination
of many specimens of M. turcica, whereas when Seifert (1988) established the synonymy,
he had only the five syntype workers and one queen of M. turcica on which to base his
opinion of variability in that species. Furthermore, Santschi's (1931b) original description
of M . turcica was misleading because he said ".... the higher edge [of the scape bend] almost
as lobate as that of sabuleti"; this indeed also fits M. georgica.
If M. georgica is not a synonym of M. turcica then for now we treat it as a good species.
In our opinion the workers are quite similar to M. tulinae. However, when Seifert collected
the type series of M. georgica he also collected specimens from other Myrmica species
from Georgia. He very generously gave Graham Elmes samples (including a few
males) from more than 20 nest series: we identified 13 samples as M. tulinae (mostly
identified at that time by both Seifert and Elmes as M. sabuleti), 8 - as M. scabrinodis,
1 each of M. sabuleti and M. specioides. Seifert clearly recognised M. georgica workers
to be different from these series including M. tulinae (thought to be M. sabuleti at the
time) and other than the paratypes from the holotype nest series there were no M. georgica
among this material. We can not exclude the possibility that it may be conspecific
with M. tulinae (in which case the name M. georgica would have priority), but unfortu­
nately Seifert had no males from the type series from Georgia that might have helped
clarify the situation. The status of M. georgica might only be resolved after obtaining
additional material, including males, from the Transcaucasus. Queens and males are
unknown.

142
 Ecology is poorly known. Seifert (1987) noted that M. georgica seems to be a lowland
species, found at altitudes 500-620 m a.s.l., where it inhabits "north-exposed steppe slope
with a dense field layer of Saloia and Eryngium species-containing phytoassociations. This
suggests that Myrmica georgica will avoid at lower geographical latitudes the most
xerothermous and oligotrophic places" (loc. cit., p. 187).

Myrmica gigantea (Collingwood, 1962)

(Fig. 95)

Aphaenogaster (Nystalomyrma) gigantea Collingwood, 1962: 225, w, q, Burma; Bolton 1995: 279.
Myrmica gigantea: Radchenko and Elmes 1998: 13.
Junior synonym of M. ritae: Collingwood 1970: 374; not confirmed, see Notes to M. ritae, below.

Type localities and type specimens. "1 q, 1 w, Kombaiti Mts., 2000 m, 24/6/34, leg. R. Malaise".
Material examined. Holotype, w, "NE Burma, Kombaiti, 15.iv.1934, leg. R. Malaise" (STOCK-
HOLM); paratype, 1 q with same collection data as the holotype (STOCKHOLM); non-type material:
1 w, "NE Burma, Kombaiti, 15.iv.1934, leg. R. Malaise" (STOCKHOLM).
Distribution (Map 40). The species is known only from 2 workers and 1 queen from
Burma (Myanmar).
Etymology. M. gigantea: from the Latin word gigas = giant, to indicate its exceptionally
large size compared with similar Aphaenogaster Mayr species (see Notes below).
Notes. M. gigantea was originally placed to the genus Aphaenogaster. Later Colling-
wood (1970) removed it to Myrmica, but erroneously synonymised it with M. ritae. It be-
longs to the ritae-complex of the ritae species group and most resembles M. urbanii by the
sculpture of head dorsum but well differs from the latter by its greater size, shorter scape,
petiole and propodeal spines. When examining a miscellaneous collection of myrmycines

Map 40. Distribution of M. gigantea.

143
loaned by STOCKHOLM in 1999, we found a second worker with the same collection data
as the holotype that without doubt belongs to the type series, but formally it can not be
considered as a paratype, because Collingwood described this species based only on the
holotype worker and single paratype queen. Males are unknown.
Ecology is unknown.

Myrmica hecate Weber, 1947

(Figs 96-98)

Myrmica smythiesi subsp. hecate Weber, 1947: 461, w. India; Bolton 1995: 279.
Myrmica hecate: Radchenko and Elmes 2001b: 250, 2002: 42, q, m.

Type localities and type specimens. "Described from two workers with the Indian Museum labels
"sweepings in grass and low herbage, Brunetti, Darjiling, 6,000 feet, 24.IX.08, No. 8607-19" and
"Botanical gardens, Darjiling, 6,900 feet, 7.viii.09, С Pavia, No. 8609-19".
Material examined. Lectotype, w (designated by Radchenko and Elmes 2001b), "Himalaya, Dar-
jeeling, Botanical gardens, 6900 ft, 7.viii.09, No. 8609-19, Pavia" (HARVARD); paralectotype,
w (damaged), "Darjeeling, 6000 ft, 24.ix.09, No. 8607-19, Brunetti (HARVARD); non-type material:
about 50 w, 2 q, 2 m from India and Nepal (for details see Radchenko and Elmes 2001b).
Distribution (Map 41): Himalaya.
Etymoiogy. M. hecate: named for Hecate {'Екатц), the Greek goddess of sorcery and
witchcraft.
Notes. M. hecate belongs to the rugosa species group and most resembles M. aimonis­
sabaudiae, differing from the latter by its multidentate mandibles that have more than 10
small teeth.
Ecology is unknown except that it lives at altitudes between 1300 and 3700 m a.s.l.

Map 41. Distribution of M hecate.

144
 Myrmica hellenica Finzi, 1926
(Figs 99-101)

Myrmica rugulosa var. hellenica Finzi, 1926: 93, w, q, m (first available use of Myrmica scabrinodis
r. rugulosa var. hellenica Forel, 1913: 431, w, q, Greece); Weber 1948: 306.
Myrmica hellenica: Agosti and Collingwood 1987b: 267; Seifert 1988: 13; Radchenko 1994b: 42;
Assing 1994: 298; Seifert 1994: 10; Bolton 1995: 279. Seifert 1996: 224; Radchenko et al. 1997:
488; Czechowski et al. 1998: 103, 2002: 25; Radchenko, Czechowska and Czechowski 2004: 48;
Seifert 2007: 196; Radchenko 2009: 70 (part).
Myrmica {Myrmica) rugulosa Nyl. var. rugulososcabrinodis Karawajew, 1929: 205, w, q, m, Georgia
and Armenia (nec Myrmica rubra var. rugulososcabrinodis Forel, 1874: 226, nomen nudum);
Weber 1948: 308. Synonymy by Seifert et al. 2009: 68; confirmed here.
Myrmica rugulososcabrinodis: Radchenko and Elmes 2004: 321; Radchenko and Antonova 2004: 19.
Myrmica rugulosa subsp. caucasica Arnoldi, 1934: 165 (not necessary proposed replacement name
for Myrmica rugulosa Nyl. var. rugulososcabrinodis Karawajew, 1929: 205); Weber 1948: 306.
Synonymy by Seifert et al. 2009: 68; confirmed here.
Myrmica sancta subsp. caucasica: Arnoldi 1970: 1841. Raised to species: Seifert 1988: 19.
Junior synonym of M. bessarabica: Dlussky et al. 1990: 182; Atanassov, Dlussky 1992: 93; Radchenko
1994d: 77; Bolton 1995: 278; not confirmed, see Notes to M. bessarabica in Chapter 3.5.
Myrmica rugulosa var. minuta Karawajew, 1929: 204, w, m, Ukraine (junior primary homonym of
Myrmica minuta Say, 1836: 294 and Myrmica laevinodis var. minuta Ruzsky, 1905: 670).
Synonymy by Radchenko 2009: 70; not confirmed, see Notes to M. constricta.
Myrmica rugulosa var. constricta Karawajew, 1934: 74 (replacement name for Myrmica rugulosa var.
minuta Karawajew, 1929: 204). Synonymy by Arnoldi 1970: 1840 (as synonym of M. rugulosa);
Radchenko 2009: 70 (as synonym of M. hellenica); not confirmed, see Notes to M. constricta.
Myrmica rugulosa var. slobodensis Karawajew, 1936: 274 (first available use of Myrmica rugulosa
rugulosa var. slobodensis Arnoldi, 1934: 162) (not necessary proposed replacement name for Myr-
mica rugulosa var. constricta Karawajew, 1934: 74). Synonymy by Arnoldi 1970: 1840; Radchenko
1994d: 75 (as synonym of M. rugulosa); Radchenko 2009: 70 (as synonym of M. hellenica); not
confirmed, see Notes to M. constricta.
Myrmica rugulosoides var. striata Finzi, 1926: 96, w, q, m, Italy. Synonymy by Collingwood 1979: 56;
Bolton 1995: 283; Gusten et al. 2006: 29 (as synonym of M. specioides); Seifert et al. 2009: 68 (as
synonym of M. hellenica); the latter not confirmed, see Notes to M. specioides.

Type localities and type specimens. M. hellenica: "Patras et Corfou (Grèce)". M. rugulososcabri-
nodis: "Fluss Mzymta, Tschernomorskij Kreis, Kaukasus, 10.IX.1910 (Nr. 3980), Satunin, ww und
1 geflüg. q. - Georgijevsko-Osetinskij Aul, Kuban-Gebiet, 28.VII.1903 (Nr. 3982), Kaznakov und Die-
terichs, ww und 2 m . - Jailatsh-Chaman-Tshaj, 6000', Gouv. Jelisavetpol, 27.VI.1909 (Nr. 3983),
A. Shelkovnikov. - Betsho, Tshaba, Svanetien, 23.VII. (Nr. 3981), derselbe, ww. - Shusha, Gouv.
Jelisavetpol, VIII.1906 (Nr. 3985), Zaizev, ww. - Wahrscheinlich auch die Übrigen, in meiner oben
genannten Arbeit angegeben Orte, von denen die entsprechenden Exemplare im meiner Sammlung
nicht mehr besitze. - Ausserdem - Sotschi, Tschernomorkij Kreis, VII.1917 (Nr. 3984), Karawajew,
1 w. - Insel Sevan auf dem See Goktscha, Armenien (Nr. 3997), Zaizev, zahlreiche ww und 6 m".
Material examined. Syntypes of M. hellenica, 1 w, "Corfu", "J. Sahib.", "584", "Myrmica scabrin-
odis Nyl. r. rugulosa Nyl. var. hellenica For. W type"; 1 w, "Patras", "U. Sahib." (HELSINKI); lecto-
type of M. rugulososcabrinodis (designated by Seifert et al. 2009), w (top specimen on the pin with 3
workers), "Шуша, Елизавет, [польской] [-[губернии], VIII. 1906, Зайцев" [Shusha, Elizavetpolskaya Gu-
bernia, viii.1906, Zaitsev], "3985 Coli. Karawajewi", "Myrmica arnoldii Kar. Karawajew d et.", "Lec-
totype (top specimen) Myrmica rugulososcabrinodis Karawajew, 1929, design. Seifert, 2005" (KIEV);
paralectotypes: 2 w on the same pin; "Бсчо, г. Чусби, Сванстия, 23.VII, Шслковников" [Becho, Mt.
Chubsi, Svanetia, 23.vii, Shelkovnikov], "3981 Coli. Karawajewi"; 2 w, 2 q, "р. Мзымта, Черномор, ок.,

145
10.IX.1910, Сатунин" [riv. Mzymta, Chernomorski okrug, 10.ix.1910, Satunin], "3980 Coll. Karawa­
jewi", "Myrmica arnoldii [Siel] Kar. Karawajew det.", "Syntypus Myrmica rugulosa var. ruguloso­
scabrinodis Karaw."; 2 w, 2 m, "Георг.[исвско]-Осст.[инский] аул, Кубан.[скнй] ок.[руг], 28.VII.1907
[Sic\], Казнаков и Дагсрикс" [Georgievsko-Osetinski aul, Kubansky okrug, 28.vii.1907, Kaznakov and
Dageriks], "3982 Coll. Karawajewi", "Myrmica arnoldii [Siel] Kar. Karawajew det", "Syntypus Myr­
mica rugulosa var. rugulososcabrinodis Karaw."; 1 w, "Сочи, Черном.[ский] ок.[руг], VII.1917, В.
Караваев" [Sochi, Chernoraorsky okrug, vii.1917, W. Karawajew], "3984 Coll. Karawajewi", "Myrmica
arnoldii [Siel] Kar. Karawajew det", "Syntypus Myrmica rugulosa var. rugulososcabrinodis Karaw.";
20 w, б m, "Севан, на Гокчс, 8.VIII.1924, Зайцев" [Sevan, on Goktcha, 8.VÍÜ.1924, Zaitsev], "3997
Coll. Karawajewi", "Myrmica arnoldii [Siel] Kar. Karawajew det", "Syntypus Myrmica rugulosa var.
rugulososcabrinodis Karaw." (KIEV); non-type material: > 300 w, 25 q and 15 m from Caucasus,
Turkey, Bulgaria and Ukraine.
Distribution (Map 42). Caucasus, Asia Minor, Ukraine (Crimea), Bulgaria, Greece,
Italy, Croatia, Slovenia and Serbia.
Etymology. M. hellenica: derived from the Greek word ШАтка (Hellenica), which re­
ferred to writings on Greece, but now means anything Greek, to indicate that it was de­
scribed from Greece. M. rugulososcabrinodis: a combination of the names M. rugulosa and
M. scabrinodis, to indicate that Karawajew considered that this species had morphological
elements of both these species and was intermediate.
Notes. M. hellenica belongs to the rugulosa-complex of the scabrinodis species group
(see Radchenko and Elmes 2004) and might be confused with M. specioides and
M. constricta. It differs from M. specioides by its less curved frontal carinae and wider frons.
The Caucasian and Turkish populations of this species have mean FI 0.43 and FLI 1.15, and
the Crimean populations have mean FI 0.41 and FLI 1.20, while mean FI in M. specioides
is 0.38, and FLI 1.32. Although the scape of M. hellenica is clearly angled at its base, it has
at most a narrow longitudinal ridge on the bend rather than the distinct lobe of
M. specioides.
M. hellenica separates from M. constricta first by the sculpture of the head, its frons is
completely longitudinally rugulose whereas the upper (rear) third of that of M. constricta
has distinct reticulation, and secondly its propodeal spines are distinctly longer than those
of M. constricta. Queens of M. hellenica are distinctly larger than those of M. constricta
(and M. rugulosa), which are quite small (HW > 1 mm vs. HW < 1 mm respectively). Males
of M. constricta well differ from those of M. hellenica by much shorter standing hairs on the
mid and hind tibiae and tarsi.
The taxonomic history of the name M. rugulososcabrinodis is somewhat tangled.
Karawajew (1929) described Myrmica rugulosa var. rugulososcabrinodis from Georgia and
Armenia based on all three castes. Arnoldi (1934) considered that this name was a primary
homonym of M. rubra var. rugulososcabrinodis Forel, 1874 and proposed a replacement
name - M. rugulosa caucasica that was generally accepted in subsequent myrmecological
literature. However, the name M. rubra var. rugulososcabrinodis Forel, 1874 is nomen
nudum, hence Arnoldi's caucasica is an unnecessary proposed replacement name and the
name M. rugulosa var. rugulososcabrinodis Karawajew is valid. Recently we believed that
we had evidence to show M. rugulososcabrinodis to be a good species and we revived
the name from synonymy (Radchenko and Elmes 2004), our study was based on a rich
material collected by us from Turkey, the Caucasus, Crimea and Bulgaria, and a detailed re­
investigation of the types of Myrmica rugulosa var. rugulososcabrinodis (KIEV).
Arnoldi's "caucasica" was at different times considered to be a subspecies of M. sancta
(Arnoldi 1970), a good species (Seifert 1988) or a synonym of M. bessarabica (Dlussky et

146
al. 1990; Atanassov, Dlussky 1992; Radchenko 1994d). However, Seifert (2002) proposed
that M. bessarabica should be considered as incertae sedis in the genus Myrmica (see notes
to M. specioides, below). This complicated taxonomic situation caused Bolton (1995) and
Bolton et al. (2006) to erroneously treat M. caucasica and M. rugulososcabrinodis as dif-
ferent species. Most recently Seifert et al. (2009) synonymised M. rugulososcabrinodis with
M. hellenica, considering M. constricta as a separate species that is closely related to
M. hellenica. We accept this latest opinion.
Ecology. The following is a combination of our (Radchenko and Elmes 2004) observa-
tions for M. rugulososcabrinodis in Turkey and those of Seifert et al. (2009) for M. hellenica.
Over its geographical distribution it occupies a very wide range of both anthropogenic and
semi-natural biotopes but we found it to be most abundant in hay meadows on the north
facing slopes (higher rainfall areas) at 400-1700 m a.s.l. in the northeast Turkish mountains.
Here it builds nests in soil, often under stones and colonies have one to several queens and
contain several hundred of workers. Seifert et al. (2009) write that "It is in its abundance
and competitive power the absolutely dominant Myrmica species at the Caucasian coast of
the Black Sea at altitudes below 600 m a.s.l.". In the Caucasus it can be found in sunny po-
sitions as high as 1600 m a.s.l. and in Turkey Seifert found it at 2300 m a.s.l., which accords
with our observations.
Seifert et al. (2009) use climate data to suggest that the species is ecologically very plas-
tic with air temperatures at recorded sites for M. hellenica ranging from 7°C in May to 24°C
in August with a huge amplitude around the mean (TAS 17.7 ± 4.0°C) in contrast to the
related M. constricta (TAS 16.3 ± 1.8°C; 14-23°C from May - August); they suggest that
something other than low winter and summer temperatures limit the northern expansion of
M. hellenica. We suggest that probably M. hellenica has adapted behaviourally and physi-
ologically to make use of southern mountain habitats that are cold in winter and hot in
summer with a relatively short "growing season" (see Chapter 1.3 for an overview of such
differences).
In Turkey we found that the microhabitats used by M. hellenica are usually quite cool
in spring, either they are wet (sometimes even flooded) or they live under the long grass in
hay meadows. In high summer theses microhabitats become typically xerothermous, the
wet meadows are grazed and dry out, the hay is cut and the grass becomes dormant. It
seems to us that were it not for the cool shady conditions in spring the sites where
M. hellenica is abundant would be dominated by ants of other genera better adapted to
withstand the high summer soil temperatures - Tapinoma Foerster, Tetramorium, Lasius
and Formica species, the cool spring conditions prevent such species from successfully com-
pleting their annual life cycle. Therefore the challenge for M. hellenica is to complete its life
cycle before the habitat becomes too hot and dry for normal foraging. In Turkish popula-
tions sexuais are in the nest by early August and we suspected that nuptial flights usually
take place by mid-August. Colonies then more or less aestivate foraging for very short pe-
riods at the start and end of daylight.
We suggest that M. hellenica might have a high metabolism more similar to that of
northern species (e.g. M. ruginodis, see Chapter 1.3.5) that enables it to rear larvae in the
cool spring conditions when there could be abundant insect prey. On the other hand, Seifert
et al.'s (2009) climatic data for the related M. constricta indicates that a lower metabolism
associated with more southern species (e.g. M. sabuleti) might be more appropriate to
its habitat. If the ants can achieve significantly longer foraging time in spring and autumn
they tend to "grow brood" more slowly than species adapted to short growing seasons

147
(see Chapter 1.3.5). So it might be that M. hellenica originally derived from M. constricta
in order to exploit southern mountain habitats, then due to anthropogenic influences, such
as the creation of hay meadows, M. hellenica has become more common and widespread
than it would be without man's influence. It would be interesting to test this idea by a com-
parative physiological and behavioural study of the two species.

Map 42. DistributionofM . hellenica.

Myrmica heterorhytida R a d c h e n k o et Elmes, 2 0 0 9

(Fig. 102)

Myrmica heterorhytida Radchenko et Elmes, 2009a: 83, China.

Type localities and type specimens. China, Yunnan Prov., for details see material examined.
Material examined. Holotype, w, China, N Yunnan Prov., Zhongdian, 3300 m, 6-8.vii.2002, leg.
S. Murzin and I. Shokhin (PETERSBURG); non-type material: 2 w, China, Yunnan, Zhongdian Mts.,
Shangri-La, 3800-4300 m, 26.V.2006, leg. I. Shokhin (PETERSBURG, KIEV).
Distribution (Map 43). The species is known only from Yunnan Province of China.
Etymology. M. heterorhytida: from the Greek words heteros (èrepoç) = different or
various, and rhytis (poriç) = wrinkle or ruga, to reflect the various kinds of sculpture on the
alitrunk.
Notes. M. heterorhytida belongs to the pachei species group. It resembles
M. yunnanensis, but differs from the latter by the curved outwards frontal carinae, which
merge with the rugae that surround antennal sockets, by the narrower frons, and by the
coarsely and densely punctated, dull surface of the head dorsum between rugae. Queens and
males are unknown.

148
 Ecology is unknown except that the workers were found on mountain meadow at the
altitude 3300-4300 m a.s.l.

Map 43. Distribution of M. heterorhytida.

Myrmica hirsuta Elmes, 1978

(Figs 103-105)

Myrmica hirsuta Elmes, 1978a: 131, q, m, England; Collingwood 1979: 51; Assing 1987: 78; Bolton
1988: 4; Seifert 1988: 42; Vepsäläinen and Pisarski 1982: 328; Elmes 1994: 439, w; Bolton 1995:
278; Saaristo 1995: 155; Seifert 1994: 15, 1996: 234; Czcchowska and Radchenko 1997: 53; Rad-
chenko et al. 1997: 491; Bregant 1998: 3; Czechowski et al. 2002: 31; Radchenko and Elmes 2003a:
228; Müller H. 2003: 11; Boer and Nordijk 2004: 25; Radchenko, Czechowska and Czechowski
. 2004:48; Seifert 2007: 201.

Type localities and type specimens. "Material examined. 145 queens were discovered living in
eight colonies of Myrmica sabuleti from Durlstone Country Park, Purbeck, Dorset (50°36'N,
1°58'W)... Forty-seven of these queens were mounted and placed in my collection, one of these
has been designated as holotype while others form paratypes... Twenty-eight Myrmica hirsuta males
were collected from three of these colonies in 1974. These also form paratypes. The holotype,
two female paratypes and three male have been deposited in the British Museum (Natural His-
tory)".
Material examined. Holotype, q, "Durlstone Country Park, Purbeck, Dorset ., 1973, Leg. GW
Elmes" (LONDON); paratypes: 34 q, 19 m, Durleston Country Park, Purbeck, Dorset, UK, under
stone, limestone grassland, leg. G. W. Elmes, 1973 (in 9 different infested colonies of M. sabuleti)
(LONDON, ELMES, KIEV); non-type material: > 250 q, 75 m, 3 w, 1 gynandromorph from England,
Denmark, Finland, Austria, Germany, Hungary and Poland.
Distribution (Map 44). The species is known from southern England, Denmark, Swe-
den, southern Finland, Netherlands, Germany, Austria, Hungary, southern Poland, Czech
and Slovak Republics, Serbia and European part of Russia.

149
 Etymology. M. hirsuta: from the Latin word hirsuta .= hairy, to describe its especially
hairy appearance (all castes) compared to most Myrmica species.
Notes. M. hirsuta belongs to the sabuleti-compiex of the scabrinodis species group (Rad-
chenko and Elmes 2004). When Elmes (1978a) described M. hirsuta from southern England,
he considered it as a workerless social parasite of M. sabuleti. Later, in infested nests from
Denmark he found two workers (Elmes 1994), both are pseudogyne, having minute ocelli and
at least a trace of scutum (see also Radchenko and Elmes 2003a). In Finland, this species was
also found in the nests of M. lonae.
M. hirsuta is a very hairy species and generally similar to M. bibikoffi. Its queens are
smaller than those of M. bibikoffi and also differ by their body sculpture and wider frons.
Males have a relatively long scape, similar to that of M. sabuleti, but well differ from the lat-
ter by the presence of long standing hairs on the head margins.
Ecology. In Central Europe M. hirsuta is clearly an obligatory social parasite of M. sa-
buleti. In North Europe, M. hirsuta usually lives in M. lonae colonies. This could indicate
a degree of host transference but equally might indicate that M. lonae is only an "ecologi-
cal race" of M. sabuleti (see Notes to M. lonae).
The worker caste is produced only very rarely: three entire infested host colonies of
M. sabuleti from Denmark were collected and each worker examined, this yielding only 3
M. hirsuta workers (2 from one colony and 1 from another). Although 8 entire infested
colonies were collected at the type locality (Elmes 1978) only queens and males were
examined individually, so it was possible these colonies also contained a few very worker-
like pseudogynes. We suggest that if host colonies infested by other related social parasites,
such as M. laurae, were exhaustively searched in the same way, a few workers of these
species might be found. Elmes (1983) showed that in the laboratory M. hirsuta queens pro-
duced two types of offspring: larger larvae develop slowly, overwinter and eclose to become
new fully reproductive M. hirsuta gynes the following spring, whereas smaller larvae either
develop quickly and eclose the same summer, becoming infertile (no spermatheca) inter-
castes (winged workers), or they overwinter and eclose to become infertile (or sub-fertile)
small M. hirsuta queens. M. hirsuta queens can suppress the sexual development of their
host's larvae (queen effect, see Chapter 1.3.3), but their own larvae appear to be "immune"
to queen effect (Elmes 1983).
Pitfall trapping showed M. hirsuta queens were present in traps from August to
October, numbers peaking in early September, at four sites in Southern England where
the host is common (Elmes 1982). Other similar studies (unpublished) confirm this trend
so that one might state with reasonable confidence that on sites where M. sabuleti is the
dominant species of Myrmica, one might expect to find some colonies infested with
M. hirsuta. Even so, the levels of infestation is not high on average only about 1 in 50-
100 host colonies are infested (Elmes 1983), thus one has to examine a large number of
host colonies before finding M. hirsuta; generally this is easier when the host lives under
stones. However, infested colonies tend to be clumped as was the case at the type
locality and in Denmark, while Seifert (1988) found that up to 50% of colonies were in-
fested on one small site. The low density of overall infestation combined with the
apparent availability of young queens searching for host colonies in autumn, suggests
that penetration of new host colonies might be quite difficult. Elmes (1983) had results
that suggested that once insinuated into a host nest, a M. hirsuta queen might delay or
suppress the onset of oviposition of the host queen by several weeks. If this is correct
then it might be a way in which the relatively low number of eggs laid by the parasite

150
might avoid direct competition with the much larger number of eggs laid by the host (see
Chapter 1.3.4).
It was suggested (Elmes 1978b) that there might be a trend for extreme polygyny to
"degenerate" into social parasitism via forms such as microgyne M. rubra (see Chapter
1.3.6). Certainly, the closest relative of M. hirsuta genetically is M. sabuleti (Savolainen
and Vepsäläinen 2003). On the other hand, its closest relative among the social parasites,
M. bibikoffi (see above), has been recorded as free living and in mixed colonies with M. sa-
buleti and arguably this could illustrate a trend from temporary social parasitism to facul-
tative social parasitism (with some workers) in certain ecological conditions, to obligate
social parasitism with just a few workers being produced in some conditions.

Map 44. Distribution of M. hirsuta.

Myrmica hlavaci Radchenko et Elmes, 2009

(Fig. 106)

Myrmica hlavaci Radchenko et Elmes, 2009a: 76, w, China.

Type localities and type specimens. China, Sichuan Prov., for details see material examined.
Material examined. Holotype, w, China, Sichuan Prov., Li Xian Miyaluo Nat. Res., Danzhamu val-
ley, 3250 m, 31°23,902'N, 103°14,827'E, 18-22.V.2006, leg. Sehnal and Tryzne (KIEV); paratypes: 2
workers with the same label (KIEV, HLAVAC).
Distribution (Map 45). It is known from the type locality in Sichuan Prov., China.
Etymology. M. hlavaci: this species was dedicated to the amateur entomologist Peter
Hlavac of Kosice, Slovakia, who provided to us material for investigation.
Notes. M. hlavaci belongs to the pachei'-group and most resembles M. phalacra. It dif-
fers from the latter species by the densely rugulose head dorsum, sculpture of propodeal
dorsum, shorter scape, etc. (see also Key, Chapter 5.7). Queens and males are unknown.
Ecology. Unknown.

151
 Map 45. Distribution of M. hlavaci.

Myrmica indica Weber, 1950

(Figs 107-109)

Myrmica ritae subsp. indica Weber, 1950: 221, w, India.

Myrmica indica: Radchenko 1994b: 44; Bolton 1995: 279; Radchenko and Elmes 1998: 11, w (part),
q, m, 2001b: 241 (part.); Elmes and Radchenko 2009: 117.
Myrmica ritae: Collingwood 1970: 374, misidentification.

Type localities and type specimens. "Described from two workers from Tonglu, Darjiling distr.,
E. Himalayas, 10,000 feet, 22.IV.10 (С. W. Beebe) in Dr. W. M. Wheeler's collection, with the Indian
Museum labels and numbers 8612-19 and 8614-19".
Material examined. Lectotype, w (designated by Radchenko and Elmes 1998), "Tonglu, E. Hi­
malayas, 10,000 ft., 22.04.10 (C. W. Beebe)" (HARVARD); paralectotype: worker, with same label
(HARVARD); non-type material: about 100 w, 1 q, 2 m from India, Nepal and Bhutan.
Distribution (Map 46). India (Darjeeling), Nepal and Bhutan.
Etymology. M. indica: from the Latin word indica = pertaining to India.
Notes. M. indica belongs to the rziae-complex of the ritae species group. Recently we
showed that "M. indica" as known in earlier publications (see Radchenko and Elmes 1998,
2001b) was in fact a complex of several related species (Elmes and Radchenko 2009). Two
new species were recognised and morphological differences confirmed by morphometric
analysis; M. alperti had a more robust body and shorter appendages, a differently shaped
petiole and more rounded apically propodeal lobes (see above and Key, Chapter 5.7);
M. weberi was distinctly smaller and more "glossy" with frontal carinae that curve outwards
to merge with the ruga that surround the antennal socket (see below and Key, Chapter 5.7).
All three forms resemble M. serica by the sculpture on the head dorsum, but differ from the
latter by the absence of standing hairs on the genae.
The remaining material was grouped under the name M. indica. However, the type
specimens are from Darjiling district, an area under represented in the material, and they
differ somewhat from the material collected in Bhutan. We would not be surprised when

152
more material is available if it is eventually shown that M. indica actually includes at least
one more species. The new species are clearly closely related to M. indica and probably all
forms are local endemics derived from a common ancestor isolated on different Himalayan
mountain systems.
Material from Nepal that was determined as M. ritae by Collingwood (1970) is in fact
M. indica (examined).
Ecology is poorly known. It lives in open forests at about 2500 m a.s.l.

Map 46. Distribution of M. índica.

Myrmica inezae Forel, 1902

(Figs 110, 111)

Myrmica inezae Forel, 1902: 226, w, India; 1903: 696; Bingham 1903: 268; Emery 1921: 37; Weber
1947: 467; Chapman and Capco 1951: 126; Radchenko 1994b: 42; Bolton 1995: 279; Radchenko
and Elmes 2001b: 255, 2002: 37, q, nec Wei Z. et al. 1999: 60; Wéi С. et al. 2001: 561, misidenti-
fications.

Type localities and type specimens. "Une seule w de Pachmarchi (Schurr)".

Material examined. Holotype, w, "Himalaya, Pachmarchi (Schurr)" (GENEVA); non-type mate­
rial: 20 w, 1 q form India (Himahal Pradesh).
Distribution (Map 47). India (Himahal Pradesh).
Etymology. M. inezae: Forel's eldest daughter was called Inez and although he did not
specifically say so, this species was most probably dedicated to her; the name is derived
from Greek and means chaste. Forel also gave this name to species in two other genera.
Notes. M. inezae is a very conspicuous species, belonging to the inezae species group.
Even though this species was described only from a single worker, it so well differs from any
other known Myrmica species that we had no trouble attributing the non-type material to
this species. Its main features are the coarse transversa] rugosity on the alitrunk dorsum

153
combined with long propodeal spines and a long petiole having a very long and thin
peduncle.
Radchenko (1994b) mistakenly placed M. inezae to the rugosa species group. The record
of M inezae for China (Wei Z. et al. 1999; Wei С. et al. 2001) should be attributed to M. wei.
Males are unknown.
Ecology is unknown.

Map 47. Distribution of M. inezae.

Myrmica inucta Radchenko et Elmes, 2006

(Fig. 112)

Myrmica inucta Radchenko et Elmes, 2006: 511, w, Kazakhstan (in: Radchenko, Elmes and Alicata 2006).

Type localities and type specimens. Kazakhstan, for details see material examined.
Material examined. Holotype, w, "Казахстан, Наурзумский заповедник, No 625, Т. Брагина" [Kaza­
khstan, Naurzumsky Nature Reserve, No 625, leg. T. Bragina] (KIEV); paratypes: 3 workers from the
nest of the holotype (KIEV, ELMES).
Distribution (Map 48). It is known only from the type locality in Kazakhstan.
Etymology. M. inucta: from the Latin word inuctus = oily sheen, to indicate the black,
shiny appearance of the body surface.
Notes. M. inucta belongs to the schencki species group and well differ from other species
of this group by its strongly reduced body sculpture and entirely black body. Generally,
reduced sculpture of the body is very rare in the genus Myrmica: prior to the discovery of
this species, M. nitida from the Himalaya was the only Old World species known to have
a reduced sculpture. Queens and males are unknown.

154
 Ecology. When we described this species we speculated that M. inucta might be a relict
species of the salted steppes that at one time were quite common in northwest Kazakhstan.
The type specimens were found in a salted depression on tall-grass halophytous meadow-
steppe, in a nature reserve famous for its salted marshes. Such depressions become very
salted with salt deposited on the soil surface, due to evaporation after being filled with
water. The margins of these salted ponds or lakes form distinct habitats for many species
of plants and animals, including some Myrmica species (see account of M. bergi).

Map 48. Distribution of M. inucta.

Myrmica jennyae Elmes, Radchenko et Aktaç, 2002

(Figs 113-115)

Myrmica jennyae Elmes, Radchenko et Aktac, 2002: 163, w, q, m, Turkey, Russia.

Type localities and type specimens. Turkey and Russia (Dagestan), for details see material
examined.
Material examined. Holotype, w, NE Turkey, Trabzon Region, nr. UzungòT, 40°35'05"N,
40°17'47"E, alt. 2257 m, 16.viii.1999, TR-43 (leg J: E. and G. W. Elmes) (EDIRNE); paratypes: 20 w,
13 q, 4 m from nest of holotype; 4 w, same locality! date and collectors, TR-45; 12 w, 1 q, NE Turkey,
Trabzon Region, Súmela Yayla, 40°38'44"N, 39°40'40"E, alt. 1570 m, 19.viii.1999, TR-55 (leg J. E.
and G. W. Elmes); 11 w, NE Turkey, Trabzon region, Hamsiköy - Balahor deresi, 46 km. SE Trabzon,
alt. 1600 m, 08.vi.1975, T-94 (leg. N. Aktac); 5 w, NE Turkey, Artvin Region, Kafkasör mt., above
and just west of Artvin town, alt. 1750 m, 14.viii.2000, TR-113 (leg. G. W Elmes); 6 w, NE Turkey,
Artvin region, Kafkasör yaylasi, alt. 1730 m, 14.viii.2000, No. 00/107 (leg. K. Kiran); 3 w, NE Turkey,
Rize Region, Ovitdagi Geçidi, 50 km SE Rize, alt. 2600 m, 30.vi.1993, No. 1190 (leg. A. Schulz); 9 w,

155
Russia, Dagestan, high mountains regions, 1972 (leg. Exp. of Dagestan State University) (EDIRNE,
KIEV, WARSAW, ELMES).
Distribution (Map 49). The species is known from the NE Turkey and Dagestan.
Etymology. M. jennyae: this species was dedicated to Jennifer (Jenny) Elmes, wife of
Graham Elmes, who followed foraging workers to find the nest that contained the holotype
series.
Notes. The taxonomic position of this peculiar species is unclear. The workers and
queens of M. jennyae clearly differ from all other known Myrmica species by their extremely
wide frons (FI > 0.50). They superficially resemble scabrinodis-gvoup species, such as
M. rugulosa, M. gallienii, M. hellenica and M. constricta, by the combination of a wide
frons, a feebly curved frontal carinae and a gradually, while quite strongly curved antennal
scape. On the other hand, males of M. jennyae have a relatively long scape that is similar
to members of the lobicornis-group and is much longer than in any species of the scabri-
nodis-group. When compared to the lobicomis-group, the strongly curved antennal scape
of the female castes of M. jennyae, that is never angular and with no trace of a carina, den-
ticle or lobe, resembles some members of that group (e.g. M. sulcinodis, M. kasczenkoi,
M. displicentia, etc.). However, even apart from its extremely wide frons, M. jennyae also
well differs from each of these on at least one other character (e.g. size, colour, body sculp-
ture, shape of petiole, etc.). Furthermore, the anterior clypeal margin of M. jennyae is promi-
nent and narrowly rounded medially, while in other members of lobicornis-group it is
slightly convex and with at least a shallow notch medially.
Ecology. This species builds nests in the soil, sometimes under stones, in high mountain
meadows, cleared fir forests and forest edges. The colony containing the holotype was found
nesting in soil at a forest edge. The nest was found by following foragers, the entrance was
just a 3-4 mm hole with no visible above ground structure. The foraging workers were hard
to see, moving relatively slowly and close to the surface of the ground.

Map 49. Distribution of M. jennyae .

156
 Myrmica jessensis Forel, 1901
(Figs 116-118)

Myrmica lobicornis var. jessensis Forel, 1901a: 371, w, Japan; Emery 1908a: 180, 1921: 38; Weber
1948:286.
Myrmica scabrinodis lobicornis var. jessensis: Ruzsky 1905: 701 (unavailable name).
Myrmica jessensis: Collingwood 1976: 302 (part.); ¡Kupyanskaya 1979: 28, 1986a: 85, w, q, m, 1990:
110 (part.); Radchenko 1994b: 43, 1994c: 142, 1994Í: 88 (misprinted as iessensis), 1994g: 109
(part.); Bolton 1995 b: 279 (part.); Kim B.-J. 1996: 180 (misprinted as yessensis) (part.); Elmes et
al. 2001: 110 (pari.); Imai et al. 2003: 183; Radchenko 2005: 141, nec Ruzsky 1925: 46; Wu and
Wang 1995: 94; Wei С et al. 2001: 561, misidentifications.
Myrmica hyungokae Elmes, Radchenko et Kim, 2001: 109, w, South Korea. Synonymy by Radchenko
2005: 141; confirmed here.

Type localities and type specimens. M. jessensis: "Sapporo (ile de Jesso)". M. hyungokae: Korean
Peninsula, for detail see material examined.
Material examined. Lectotype of M. jessensis, w (designated by Radchenko 2005), "Sapporo, Mat­
sumura" (GENEVA); paralectotypes: 1 w with the same label as lectotype (GENEVA); 1 w,
"M. lobicornis jessensis, Sapporo" (GENOA); holotype of M. hyungokae, w, S. Korea, Mt. Jiri, nr.
Gure, Jeonnan prov, Nogodan area -high mountains, 1972, in pitfall trap, leg. B.-J. Kim [K-101 (2)]
(LONDON); paratypes: 3 w, same series as holotype; 4 w, [K-8 (1-3)], S. Korea, Mt. Jiri, nr. Gure, 7-
8-1972, in pitfall trap, leg. B.-J. Kim; 4 workers [K-9 (1-4)], S. Korea, Mt. Jiri, nr. Gure, 19-8-1972,
in pitfall trap, leg. B.-J. Kim; 3 workers [K-120 (2-4)], S. Korea, nr. Tokyusan 1200 m., 21.viii.1994,
leg. B.-J. Kim (ELMES, LONDON, KIEV, IKSAN); non-type material: > 50 w, 10 q and 10 m from
Japan and Korean Peninsula .
Distribution (Map 50). Japan, Korean Peninsula.
Etymology. M. jessensis: from name Jesso combined with the Latin suffix ensis = place
of origin, to indicate that the type region is Jesso (modern Hokkaido, Japan). In the 16th

Map 50. Distribution of M. jessensis.

157
century Jesso was the northern border areas of Japan, including Hokkaido, Sakhalin and
Kurilen islands inhabited by the Ainu peoples. M. hyungokae: this species was dedicated to
Hyung-Ok Kim, wife of our friend and colleague, the Korean myrmecologist Prof. Byung Jin
Kim of Wonk Wang University.
Notes. M. jessensis belongs to the lobicornis species group and it most resembles M. eid­
manni, from which it differs principally by its narrower frons and shallower metanotal
groove. Recently we described a new species, M. hyungokae from South Korea (for details
see Elmes et al. 2001). Unfortunately, this is without any doubt a synonym of M. jessensis
(Radchenko 2005). We made this mistake because all previous authors determined such
material from the continental part of East Asia as M. jessensis, even though the majority
in fact, belonged to M. eidmanni. At the time we described M. hyungokae the types of
M. eidmanni were not known (see notes to this species) and our mistake was not to refer
back to the types of M. jessensis, instead we simply accepted the quite large amount of ma­
terial from the Russian Far East as being representative of "M. jessensis" when in fact it was
M. eidmanni.
Ecology. In Japan M. jessensis is primarily a grassland species (Ichinose 1990) and we
found that it was quite common on mown roadside verges on Hokkaido island. More
generally we can say that it is a species of open areas at altitudes up to 1700 m a.s.l. and
sometimes it is quite abundant on riverbanks (Kondoh 1994). It builds nests in soil, often
under stones (Imai et al. 2003; Radchenko 2005). A nuptial flight was observed in
September (Ichinose 1990).

Myrmica juglandeti Arnoldi, 1976

(Figs 119, 120)

Myrmica juglandeti Arnoldi, 1976: 549, w, m, Kyrgyzstan; Tarbinsky 1976: 22, w, q, m; Radchenko
1994b: 4 1 , 1994c: 137, 1994c: 74; Bolton 1995: 180; Schultz et al. 2006: 206.
Myrmica tianshanica Arnoldi, 1976: 550, w, Kyrgyzstan. Synonymy by Radchenko 1994e: 74; con­
firmed here.
Myrmica tianshanica subsp. alajensis Arnoldi, 1976: 550, w, Kyrgyzstan. Synonymy by Radchenko
1994e: 74; confirmed here.
Myrmica tenuispina: Arnoldi 1948: 847, 1949: 304, misidentifications.

Type localities and type specimens. M. juglandeti: "Ферганский, Чаткальский (Арнольди,

Тарбинский) хребты, обычен преимущественно в поясе ореховых лесов. Голотип (w), аллотип (m),
паратипы - оз. Сары-Чслск (Арнольди, 1945)" [Fergansky and Chatkalsky Ranges (leg. Arnoldi, Tarbin­
sky), common mainly in the nut forests. Holotype (w), allotype (m), paratypes - Lake Sary-Chelek
(Arnoldi, 1945)]. M. tianshanica: "Тянь-Шань. Голотип (w) и 8 паратипов. Киргизский Алатау, Аламсдин
Карабалта (Арнольди, 1943)" [Tien-Shan. Holotype (w) and 8 paratypes. Kirgizsky Alatau, Alamedin
Karabalta (leg. Arnoldi, 1943)]. M. tianshanica subsp. alajensis: "Алайский хребет, голотип (w) и 3
паратипа - под перевалом Талдык (Тарбинский, Длусский - 1963); просмотрено много ww" [Alaisky
Range, holotype (w) and 3 paratypes - near Taldyk Pass (leg. Tarbinsky, Dlussky - 1963); checked
many ww].
Material examined. Paratypes of M. juglandeti: 2 w, 1 m, "Сары-Чслск, Н[ижний] Карангитун,
орешник, 13.VIII.61, Тарбинский" [Sary-Chelek, L[ower] Karangitun, nut forest, 13.VÜ.61, leg. Tarbin-
sky]; 1 w, "Киргизия, Сары-Чслск, 12.VIII.63, Тарбинский" [Kirgizia, Sary-Chelek, 12.viii.63, leg.
Tarbinsky]; 1 w, 1 m, "Киргизия, Сары-Чслск" [Kirgizia, Sary-Chelek]; (MOSCOW); holotype of

158
M. tianshanica, w, "Holotyp. Myrmica tianshanica К. Arnoldi det.", "No. 43-73" (outer specimen on
the upper triangular on the pin); paratypes: 5 w on the pin with holotype; 7 w, "Paratype. Myrmica tian­
shanica K. Arnoldi det.", "No. 43-73", "No. 43-20", "No. 45-35" (MOSCOW); holotype of
M. tianshanica subsp. alajensis, w, "Алайский xp., под пер. Талдык, Киргизия, 18.VI.63, IO. Тарбин-
ский" [Alaisky Range, near Taldyk Pass, Kirgizia, 18.vi.63, leg Yu. Tarbinsky], "Holotype Myrmica
tianshanica alajensis K. Arnoldi det." (upper specimen on the pin) (MOSCOW); paratypes: 2 w on
the pin with holotype; 3 w, "3000 м, пер. Талдык, Алай; 63-335, VI.1963, Г. Длусский" [3000 m, Taldyk
Pass, 63-335, vi.1963, leg. G. Dlussky], "Paratype Myrmica tianshanica alajensis K. Arnoldi det.";
1 w, "Алайский xp., пер. Талдык, Тарбинский, 18.VI.63" [Alaisky Range, Taldyk Pass, leg. Tarbinsky,
18.vi.63] (MOSCOW); non-type material: > 30 w, 4 q, 7 m from Tien-Shan.
Distribution (Map 51). Tien-Shan and Alai.
Etymology. M. juglandeti: named for the walnut trees (Juglandaceae) with which forests
the type specimens were associated. M. tianshanica: from the name Tian-Shan with the
adjectival suffix for nouns ica (from the Greek iko) = belonging-to or from, indicating that
it is a resident of the Tien-Shan (or Tian-Shan) mountain range that covers much of Kyr­
gyzstan and part of North-West China. M. alajensis: named from Alai (Alaj) combined with
the Latin suffix ensis = place of origin. The type locality, the Taldyk Pass, is in the Alai
Range.
Notes. M. juglandeti was placed to the "rubra-smithiesi" species group by Arnoldi
(1976), and to the rubra-group by Radchenko (1994b, e). Now we believe it belongs to the
dshungarica-group (see notes to M. dshungarica). It is similar to M. dshungarica and
M. ferganensis but differs from them by the shape of petiole, propodeal spines etc. (see also
Key, Chapter 5.4). It is a rather variable species with many isolated populations in the
mountain regions of Middle Asia, some of which were described as different species or sub­
species (see above). Based on the direct comparison of the types of M. tianshanica and
M. tianshanica subsp. alajensis with the types and non-type material of M. juglandeti, we
believed them to be local variants of M. juglandeti.

Map 51. Distribution of M. juglandeti.

159
 Ecology. Inhabits mainly walnut forests at altitudes between 1000 and 2500 m a.s.l.
where it builds its nests in a leaf litter, in soil, often under stones and in tree stumps. A nup-
tial flight was observed in July-August.

Myrmica kabylica (Cagniant, 1970)

(Figs 121, 122)

Sifolinia kabylica Cagniant, 1970: 40, q, m, Algeria; Kutter 1973: 256, 267 (misspelled as cabylica).
Myrmica kabylica: Bolton 1988: 4, 1995: 280; Radchenko and Elmes 2003: 236.
Symbiomyrma kabylica: Seifert 1994: 16.

Type localities and type specimens. "Syntypes: 6 femelles et 2 mâles. Le 10 juillet 1967, en élevage.
Provenance du nid: Plateau de PHeidztr, versant nord du Djurjura, Grande Kabylie, Algérie. Altitude
1800 m. Pâturage pseudo-alpin à Bupleuram spinosum (L.), Astragalus armatus ssp. numidicus (Coss.
et Dur.) (M.), Euphorbia bupleuroïdes ssp. luteola (Coss. et Dur.) M. et Juniperus communis var.
hemisphaerica (Presl.) Pari."
Material examined. 1 q and 1 m, "Tala Guilef, Algeria (Kabylia), 17.vii.1966, leg. Cagniant", "Syn-
types Sifolinia kabylica Cagniant" (see Notes below).
Distribution (Map 52). Algeria.
Etymology. M. kabylica: from the name Kabylie with the adjectival suffix for nouns ica
(from the Greek IKO) = belonging-to or from, to indicate that it is a resident of Kabylie,
a mountainous region of Algeria.
Notes. M. kabylica is a workerless socially-parasitic species. It was originally described
in the genus Sifolinia. Later Sifolinia (and its junior synonym Symbiomyrma) were
synonymised with Myrmica (Bolton 1988), while Seifert (1994) revived Symbiomyrma from
synonymy placing in it S. kabylica and some other species. Recently we again synonymised
Symbiomyrma with Myrmica, and placed it to the karavajevi species group (Radchenko and
Elmes 2003a).

Map 52. Distribution of M. kabylica.

1бО
 We examined a queen and male, kindly loaned to us by H. Cagniant and labelled by him
as syntypes of M. kabylica, but the locality label did not correspond with the data in the
original paper (see above) and so they can not formally be considered as types. However,
we are sure that these specimens had been compared with the types by H. Cagniant and
were indeed M. kabylica.
In its major features, M. kabylica is most similar to M. karavajevi and M. lemasnei, but
clearly differs by a combination of morphological features (for details see Radchenko and
:
Elmes 2003, and Key, Chapter 5.1).
Ecology. M. kabylica was taken from a nest piM. cagnianti living in subalpine meadow,
at altitude 1800 m (originally the host species was determined as M. aloba; see also
Espadaler 1996). It is quite possible that this species parasitizes all M. aloba-lïke species and
might well be found in nests of M. aloba on the Iberian Peninsula.

Myrmica kamtschatica Kupyanskaya, 1986

(Figs 123-125)

Myrmica kamtschatica Kupyanskaya, 1986a: 88, w, q, m, Russia; 1986b: 94, 1990: 111; Radchenko
1994b: 43, 1994c: 139, 1994Í: 86; Bolton 1995: 280; Radchenko 2005: 141; Pfeiffer et al. 2007: 5.
Myrmica aborigenica Zhigulskaya, 1991: 58, w, q, m, Russia; Zhigulskaya et al. 1992: 72; Bolton 1995:
277. Synonymy by Radchenko 1994f: 86; confirmed here.
Myrmica angulinodis: Collingwood 1976: 302; Kim B.-J. 1996: 179, misidentifications.

Type localities and type specimens. M. kamtschatica: "Материал. Голотип: Камчатская обл., Елизово,
склон сопки, 18.VII 1976, 1 рабочий (Купянская). Паратипы: более 1 тыс. рабочих, 13 самок и 10 самцов
из разных мест Камчатской, и Магаданской областей и Севера Хабаровского края. Голотип
и часть паратипов хранятся в Зоологическом музее Московского государственного университета, остальные
паратипы - Биолого-почвснном институте ДВНЦ АН СССР (Владивосток)" [Material. Holotype: Kam­
chatka Prov., Elisovo, slope of hill, 18.vii.1976, 1 worker (Kupyanskaya). Paratypes: more than 1000
of workers, 13 queens and 10 males from different localities of Kamchatka and Magadan Provinces
and northern part of Khabarovsky Krai. Holotype and1 part of the paratypes preserved in Zoological Mu­
seum of Moscow State University, rest of paratypes - in Soil and Biological Institute of FESC Ac. Sci.
of the USSR (Vladivostok)]. M. aborigenica: "Голотип - один рабочий, 30.VII 1984 г., гнездо
№ 278, Магаданская обл., Тснькинский р-н, юго-восточная оконечность хребта Большой Анначаг, северный
макросклон пика "Властный" (долина ручья Олень, притока р. Сибит-Тыэллах, впадающей слева в Колыму),
лиственничное кустарничко-сфагновос редколесье (Жигульская). Паратипы - 60 рабочих, 40 самок, 5
самцов из того же гнезда, что и голотип. Кроме того,1 имеются фаунистическис сборы из многих точек
бассейна верхней Колымы, а также рабочие (более 20 тыс.), самки (около 200) и самцы (около 100) из 50
полностью раскопанных гнезд в этом районе. Материал хранится в Институте биологических проблем
Севера ДВО АН СССР (Магадан)." [Holotype - one worker, 30.vii.1984, nest No 278, Magadan Prov.,
Ten'kinsky Distr., SE edge of the range Bol'shoi Anriachag, northern macroslope of the Mt. "Vlastnyi"
(valley of the brook Olen', afflux of the riv. Sibit-Tyellakh, left afflux of the riv. Kolyma), larch sparse
forest (Zhigulskaya). Additionally, there are faunistic collections from many localities of the basin of
upper reaches of Kolyma riv., and also workers (more than 20 thousands), queens (about 200) and
males (about 100) from 50 completely excavated nests from this region. Material preserved in Institute
of the biological problems of the North Regions of FEB of the As. Sci. of the USSR (Magadan)].
Material examined. Holotype of M. kamtschatica: w, "Камчатская обл., Елизово, склон сопки,
18.VII.1976, Купянская" [Kamchatka Prov., Elisovo, slope of hill, 18.vii.1976, leg. Kupyanskaya]
(MOSCOW); paratypes: 3 w from the nest of holotype; 10 w, 4 q, "Камчатка, Козыревск, поляна

lóí
в ельнике, 20.8.76, Купянская" [Kamchatka, Kozyrevsk, glade in fir forest, 20.viii.1976]; 3 w, 1 m,
"Камчатка, Ключи, листвен, лес на р. Бекеш, 23.8.76, Купянская" [Kamchatka, Klyuchi, larch forest, riv.
Bekesh, 23.viii.76, leg. Kupyanskaya]; 5 w, "Камчатка, окр. Усть-Болыпсрсцка, склон поймы, из бревна,
17.7.76, Купянская" [Kamchatka, vicinity of Ust'-Bolsheretsk, river valley, from log, 17.vii.76, leg.
Kupyanskaya]; 2 w, 1 m, "Камчатская обл., Ключи, 30 км С вулк. Шивелуч, 28.8.76, Купянская" [Kam­
chatka Prov., Klyuchi, 30 km N volcano Shivcluch, 28.viii.76, leg. Kupyanskaya] (MOSCOW, VLADI­
VOSTOK); paratypes of M. aborigenica: 20 w, "Верх. Колымы, Абориген, 88-13, Жигульская, 20.VII.88"
[Upper reaches of Kolyma riv., Aborigen, 88-13, Zhigulskaya, 20.vii.88]; 5 w, 5 q, 7 m, "Верх. Колымы,
Абориген, 10.7.83, Жигульская" [Upper reaches of Kolyma riv., Aborigen, 10.vii.83, Zhigulskaya]
(MOSCOW, PETERSBURG, KIEV); non-type material: > 100 w, about 20 q and 15 m from South and
East Siberia, Mongolia, Russian Far East and North Korea.
Distribution (Map 53). South and East Siberia (to the west until Altai Mts.), Mon­
golia, Russian Far East, North Korea.
Etymology. M. kamtschatica: from the name Kamchatka with the adjectival suffix for
nouns ica (from the Greek IKO) = belonging-to or from, to indicate that it is a resident of
Kamchatka Peninsula. M. aborigenica: from the name Aborigen with the adjectival suffix
for nouns ica (from the Greek IKO) = belonging-to or from, to indicate that it is a resident
of the Aborigen village in Magadan Province of Russia, where a Field Research Station of
the Institute of the biological problems of the North Regions is situated.
Notes. M. kamtschatica belongs to the lobicornis species group. It resembles
M. angulinodis by the lack of a vertical lobe on its strongly angled scape-base, but well dif­
fers by the shape of its petiole and propodeal spines (see Key, Chapter 5.3). Some popula­
tions of M. kamtschatica can have a weakly developed subvertical ridge on the bend at the
scape-base (somewhat like a "rudimental" vertical dent), a feature that led Zhigulskaya to
describe M. aborigenica. However, examination of the types of both taxa and comparison
with rich non-type material from over its whole range, shows that M. kamtschatica is rather
variable in this respect, even within populations. We therefore confirm the earlier synonymy.
Part of the material from North Korea, determined by Collingwood (1976) as
M. angulinodis, in fact is M. kamtschatica.

Map 53. Distribution of M. kamtschatica.

1Ó2
 Ecology. M. kamtschatica is a boreal species that is widespread in humid places in
several biotopes within the Taiga Zone. In the Kamchatka region it is most common in
sparse forests but also lives on meadows and peat bogs, nests are built in soil, moss mounds
and rotten wood (see Kupyanskaya 1990). In the upper Kolyma region (see Berman et al.
2010, p. 71-74) colonies contain 200-600 workers and are reported to be strictly mono-
gynous; small nests are built in mossy tussocks or "pillows" that overlay the permafrost
and are insulated by snow in winter. M. kamtschatica was used in studies of cold-hardiness
(Berman et al. 2010). Nuptial flight occurs in July-August.

Myrmica karavajevi {Arn oidi, 1930)

(Figs 126, 127)

Symbiomyrma karavajevi Arnoldi, 1930: 269, q, m, Ukraine (also described as new by Arnoldi, 1933:
41); Seifert 1994: 15; Saaristo 1995: 158; Seifert 1996: 236.
Sifolinia karavajevi: Samsinak 1964: 156; Kutter 1973: 258 (misspelled as karawajewi); Pisarski
1975: 16; Bolton and Collingwood 1975: 18; Dowes 1977: 147; Arnoldi, Dlussky 1978: 535;
Collingwood 1979: 58; Maavara 1981: 6; Blinov 1985: 7; Saaristo 1986: 97; Agosti and Collingwood
1987a: 53.
Myrmica karavajevi: Bolton 1988: 4, 1995: 280; Czechowski et al. 2002: 33; Radchenko and Elmes
2003: 231; Radchenko, Czechowska and Czechowski 2004: 44; Espadaler et al. 2004: 82.
Myrmica (subgenus Symbiomyrma) karavajevi: Seifert 2007: 210.
Sifolinia pechi Samsinak, 1957: 167, q, Czechia; Pisarski 1962: 367, m. Synonymy by Samsinak 1964:
156; Kutter 1973: 262; Seifert 1994: 15;-Bolton 1995: 281 (misspelled as pechei); Radchenko and
Elmes 2003: 231; confirmed here.
Myrmica faniensis van Boven, 1970: 127, q, Belgium; jKutter 1973: 257; van Boven 1977: 114; Bolton
1988: 4, 2005: 279. Synonymy by Seifert 1994: 15; Radchenko and Elmes 2003: 231; confirmed here.
Sifolinia winterae Kutter, 1973: 263, q, m, Switzerland; 1977: 73; Bolton 1988: 7, 1995: 284. Syno-
nymy by Seifert 1994: 15; Radchenko and Elmes 2003: 231; confirmed here.
Myrmica winterae: Bolton 1988: 4, 1995: 284.
Sifolinia laurae: Yarrow 1968: 236, q, m, misidentification.

Type localities and type specimens. M. karavajevi: "Im Sommer 1926 habe ich die Ameisenge-
meinschaften der Ostukraine auf der Biologischen Donetz-Station in der Nähe der Stadt Zmiev des
Charkowschen Kreises untersucht. Geflügelte Ç und 3 am 1. VIII. 1926 (Nr. 1197, Koll. Arnoldi) in
einem trockenen Eichenwalde auf einer Wasserscheide, in einem normalen Nest von Myrmica scabri-
nodis, welches unter dem faulenden Laub und zwischen den Baumwurzeln in der Erde gebaut war. Im
September 1929 in demselben Walde ein Nest von Myrmica scabrinodis mit vielen flügellosen $ von
Symbiomyrma (Nr. 4467). 20. IX. 1929 ein Nest derselben Ameise mit einigen flügellosen § von Sym-
biomyrma (Nr. 4480) in einer ausgehauenen Waldblöße auf einer Wasserscheide zwischen zwei trok-
kenen kleinen Schluchten in der Nähe der früheren Fundorte". S. pechi: "Typus: ein einziges, voll
ebtvwickeltes aber schon flügelloses Weibchen in meiner Sammlung. Fundort: Böhmen, Eibsand-
steingebirge, Janov in der Nähe von H ensko (Herrehskretschen), 20.VIII.1955". M. faniensis: "Le
type unique de cette nouvelle espèce est une reine sans ailes. Eli fut trouvée dans le nid de Myrmica
scabrinodis Nyl. dans les Hautes-Fagnes (Baraque Michel) le 31 mai 1951 par A. Raignier et J. van
Boven". S. winterae: "Material: 1flügellosesWiebchcn (Holotypus) im Museum Basle. Fundort: Bei
1400 m zwischen Schwyz und dem Grossen Mythen auf Baumstrunk, leg. Fräulien Winter am 17. Sep-
tember 1970. 1 gefl. ' (Allotypus) im Museum Basle. Nax, Wallis, bei Myrmica ruginodis Nyl., leg. Bu-
schinger 19.VII.1971". i

163
 Material examined. Syntypes of M. karavajevi: 2 q, 2 m, "Zmiev Kharkovsk., 1.viii.26 No 1198";
1 q, 1 m "Zmiev, prov. Kharkov, 29.ix.1929, К. Arnoldi" (MOSCOW, LONDON); paratype of S. win­
terae: 1 m, "Nax, V5, 19.vii.1971, leg. A. Buschinger", "Typus"; 1 q, "Pont de la Morge, Wallis bei
Nax, 21.vii.1971, leg. A. Buschinger", "Sifolinia winterae q, M. ruginodis w" (GENEVA) (see also
Notes below); holotype of M. faniensis: q, "Hautes Fagnes, Baraque Michel, 31-5-195, v. B.-R.", "Myr­
mica faniensis nov. spec, holotype, dct. Van Boven, 1969" (MAASTRICHT); non-type material:
several tens of q and m from Ukraine, Russia, Belarus, Moldova, Poland, Czechia, England, France,
Spain, Germany, Finland and Sweden.
Distribution (Map 54). Ukraine, European part of Russia, Belarus, Moldova, Estonia,
Finland, Sweden, Norway, Poland, Czechia, Germany, Belgium, Austria, Switzerland, Eng­
land, Italy, France, Spain.
Etymology. M. karavajevi: this species was dedicated to the famous Ukrainian myrme­
cologist Prof. Vladimir Afanasievich Karawajew (also spelled as Karavajev, see biography
in Chapter 1.6). M. pechi: this species was dedicated to the Czechoslovakian natural-
scientist Pavel Pech. M. faniensis: named for its type locality, the Fagnes region of Belgium,
being the medieval name Fanias combined with the Latin suffix ensis = place of ori­
gin. M. winterae: this species was named for the collector, the German myrmecologist
Dr. Ursula Winter of Bremen University.
Notes. M. karavajevi belongs to the karavajevi species group and is related to
M. lemasnei and M. cagnianti (for details see Radchenko and Elmes 2003, and Key, Chap­
ter 5.1, 5.2).
Arnoldi (1930) described the species (and new genus) Symbiomyrma karavajevi from
eastern Ukraine (Kharkov Province), and repeated his description in the Russian language
in 1933. S. karavajevi was considered to be a workerless socially parasitic species with its
primary host being M. scabrinodis.
Samsinak (1957) described a new species (Sifolinia pechi), which he presumed to be
a social parasite from Czechia (Bohemia), and placed it in the hitherto monotypic genus
Sifolinia; his description was based on a single queen found on the ground while observing

Map 54. Distribution of M. karavajevi.

164
a nuptial flight of M. rubra. Pisarski (1962) found this species in southern Poland and de-
scribed the males. However, Samsinak (1964) considered S. pechi as a junior synonym of
Symbiomyrma karavajevi, and at the same time synonymised Symbiomyrma with genus
Sifolinia. We have not studied the holotype of S. pechi, only Pisarski's material from Poland.
Without any doubt, the males and queens fromj Poland are the same species as S. karava-
jevi. Given that Pisarski saw the holotype of Sifolinia pechi and identified his material as
this species, we are confident in accepting the Samsiñák's (1964) synonymy.
We have seen specimens of "Sifolinia laurae" sensu Yarrow (1968) (queens and males
from Southern England, Dorset) (LONDON) and entirely agree with Kutter (1973) and
Bolton and Collingwood (1975), that these specimens are M. karavajevi.
When van Boven (1970) described Myrmicalfaniensis (from Belgium, based on a single
dealate queen) he compared it with all other known socially parasitic Myrmica species, from
which it well differs, but he did not compare it with known species of Sifolinia. Kutter
(1973) supposed that Myrmica faniensis should be a junior synonym of S. karavajevi, but
it was Seifert (1994) who formally synonymised these species (without making any com-
ments). Our examination of the holotype of M. faniensis showed that it lies within the range
of variation seen for S. karavajevi, thus we confirm the synonymy.
Finally, Kutter (1973) described S. winterae based on the holotype queen, found in
Switzerland by Miss Winter 17.ix.1970 near Schwyz road. He also described, as a paratype,
the male, collected by A. Buschinger 19.vi.1971 in Valais, near Nax, but for some reason
Kutter failed to include in the type series several queens, collected by Buschinger together
with the male mentioned above. We have studied the paratype male and one of the non-type
queens collected by Buschinger (GENEVA). The holotype queen is missing: we obtained
a queen (BASLE) with labels fully corresponding to Kutter's (1973) data for the holotype
of S. winterae (i.e. "leg. Winter, 17.ix.1970, 1400 m, Schwyz, CH", "Sifolinia winterae",
"Typus"), but this specimen is a queen of M. hecate Weber (a distinctive Himalayan species,
queens and males of which were described by Radchenko and Elmes 2002). We presume
that somebody had remounted this specimen in error after several specimens had fallen
from their pins. Obviously, Kutter's original description and drawings were based on
another specimen very similar to the queen, collected by Buschinger (GENEVA). The main
features distinguishing S. winterae from S. karavajevi, according to Kutter (1973), are
a slightly different shape of propodeal spines and the pilosity of the body. However, both of
these features are very variable in the studied nest series of M. karavajevi from its whole
range; using Kutter's characters, we would identify some specimens (from the same series)
as M. karavajevi and others - as S. winterae. Therefore, we confirm this synonymy.
Ecology and biology. M. karavajevi is probably the most widespread of the workerless
Myrmica social parasites. Yet, nowhere is it common, usually myrmecologists can find one
or two infested colonies and no more, despite ¡repeated searching. On a few sites, it has
been found in the same colonies of host ants in different years (e.g. in Finland by Savolainen
and in France by Elmes) and the M. sabuleti nest containing the specimens taken by Yarrow
(1968) continued to support the parasite for several years (unpublished observations).
It is more catholic in its choice of host species compared to many of the Myrmica social
parasites. Arnoldi (1930, 1933) found it with M. scabrinodis; in England its host was
M. sabuleti; Elmes found it with M. scabrinodis in France; in Poland the host species was
M. gallienii (Pisarski 1962 noted as host speci'es M. rugulosa, misidentification, material
examined); in Finland it hosts appear to be M. [scabrinodis, and in Sweden - M. rugulosa,
while we found M. karavajevi near St.-Petersburg in a nest of M. lonae. Similarly, the host

1651
worker on the same pin as a S. winterae queen (GENEVA) was M. gallienii (although
erroneously determined as M. ruginodis by Kutter). The host data indicates that it is either
a generalist specialising on a range species from the scabrinodis-gvoup of Myrmica, or in
some way comprises cryptic lineages each specialising on a particular species. One com-
mon feature is that all its hosts live in warm but relatively wet places in marshes, meadows
and forest glades. The host colony of M. sabuleti from England lived in rather cool damp
conditions, which begs the question whether a cryptic ecomorph of M. sabuleti, with
biology similar to M. lonae, exists in England (Elmes, unpublished).
Jansen et al. (2010) based on a molecular phylogenetic study showed that M. karavajevi
belongs to the same clade (scabrinodis-group) as its recorded hosts, but it appears to have
had a common ancestor with its host species a long time ago (ca. 17 Ma; see Fig. 332 in
Chapter 4). Thus it obeys Emery's rule in a loose sort of way: its multiple host use could be
explained either by a sudden switch to more generalist behaviour after the extinction of
its first host species or by its tracking of daughter species evolved from the original host
species (Savolainen and Vepsäläinen 2003). The great age of M. karavajevi and its rela-
tionship to its host species, illustrated by Jansen et al. (2010), suggests that the latter is more
probable.

Myrmica kasczenkoi Ruzsky, 1905

(Figs 128-130)

Myrmica scabrinodis kasczenkoi Ruzsky, 1905: 702, w, nec m, Russia; Emery 1908a: 180, 1921: 41;
Ruzsky 1936: 94; Weber 1950: 205.
Myrmica kasczenkoi: Pisarski 1969a: 228 (part.), 1969b: 296 (part.); Dlussky, Pisarski 1970: 85;
Dmitrienko, Petrenko 1976: 17; Pisarski, Krzysztofiak 1981: 156 (part.); Radchenko 1994b: 43,
1994c: 140, 1994f: 82, q, m, 1994g: 109; Bolton 1995: 280; Pfeiffer et al. 2007: 5, nec Kupyan-
skaya 1979: 128, 1986b: 94, 1990: 113; Collingwood 1976: 302; Kim B.-J. 1996: 179, misidentifi-
cations.

Type localities and type specimens. "Мсстонахожд.: Енисейская губ., Ачинский у., VIII.99 (Кащенко).
Забайкальская обл., Троицкосавск. окр., Ямаровския минеральн. воды, под камнем в соснов. бору, среди
кустарников на лужайке, 3.VII.1900 (Михно)" [Localities: Eniseiskaya Gub., Achinsky Uezd, viii.99
(leg. Kaszenko). Zabaikalskaya Obi., Troitskosvsky Uezd, vicinity of Yamarovsk mineral sources, under
stone in pine forest, in shrubs on glade, 3.vii. 1900 (leg. Mikhno)] (now - Krasnoyarsky Krai and Chita
Province of Russia).
Material examined. Neotype, w (designated by Radchenko 1994f), "Читинская обл., Даурский
заповедник, Соловьевский участок, No 105-90, 18.07.90 (А. Радчснко)" [Chita Province, Daurian
Natural Reserve, Solovievsky distr., No 105-90, 18.vii.90, leg. A. Radchenko] (KIEV); non-type
material: > 50 w, 5 q, 5 m from South Siberia and Mongolia.
Distribution (Map 55). Altai Mts., South Siberia, Mongolia.
Etymology. M. kasczenkoi: this species was dedicated to the collector, the well-known
Ukrainian zoologist, one of the founders of the Zoological Museum of the Ukrainian
Academy of Sciences (Kiev), Dr. of medicine and Dr. of zoology, Prof. Nikolai Feofanovich
Kasczenko (1855-1935).
Notes. M. kasczenkoi belongs to the lobicornis species group. It is most similar to M. pi­
sarskii by the shape and sculpture of its alitrunk and waist, but differs by the base of the
scape having a less angled curve without a vertical lobe or dent.

166
 Map 55. Distribution! of M. kasczenkoi.

Radchenko (1994f) believed that workers and males described by Ruzsky (1905) under
the name kaszcenkoi most probably belonged to different species: the male had a short
scape and probably was a specimen of M. divergens. Radchenko {loc. cii.) believed that
most probably Ruzsky's types of M. kasczenkoi are lost, therefore he designated a neotype
and described queens and males. We have not found the original types of this species in
any of the many collections examined since then.
The specimens of "M. kasczenkoi" discussed by Kupyanskaya (1989b, 1990) are in fact
M. angulinodis, and Collingwood's (1976) record of this species from North Korea belongs
to M. excelsa (corresponding material examined).
The unavailable name M. scabrinodis kasczenkoi var. subaltaica Ruzsky, 1925 was
referred to M. kasczenkoi (Radchenko 1994f).j
Ecology. M. kasczenkoi is an hemixeropliilous species, inhabiting dry and light pine
forests and steppes where it nests in the soil, often under stones. Nuptial flights are from
August-September.

Myrmica kirghisorum Arnoldi, 1976

(Figs 131-133)

Myrmica lobicornis subsp. kirghisorum Arnoldi, 1976: 553, w, m, Kyrgyzstan; Tarbinsky 1976: 33, w,
q, m . ' I
Myrmica kirghisorum: Radchenko 1994b: 43 (here and hereinafter-in 1994 misspelled as kirgisorum),
1994c: 142, 1994f: 87; Bolton 1995: 280; Schultz et al. 2006: 206.
Myrmica lobicornis subsp. foreliella Arnoldi, 1976:! 553, w, Tajikistan; Bolton 1995: 279. Synonymy
by Radchenko 1994f: 87; confirmed here.
Myrmica lobicornis: Kuznetsov-ugamsky 1927: 196, misidentification.,

167
 Type localities and type specimens. M. kirghisorum: "Голотип (w), аллотип (ó*), 12 паратнпов:
Ферганский хребет, Кара-Алма (Арнольди - 1945). Чаткальский хребет (Тарбинский), Алайскнй хребет
(Романов - 1939; Тарбинский - 1971)" [Holotype (w), allotype (m), 12 paratypes: Fergansky Range,
Kara-Alma (leg. Arnoldi - 1945), Chatkalsky Range (leg. Tarbinsky). Alaisky Range (leg. Romanov -
1939; Tarbinsky - 1971)]. M. lobicornis subsp. foreliella: "Хорог, Гунт-Шахдара (Казнаков)" [Khorog,
Gunt-Shakhdara (leg. Kaznakov)].
Material examined. Paratypes of M. kirghisorum: 6 w, 1 m (without head), "Чаткальский хр., дол.
p. Качансай, Тарбинский, 7.VII.968" [Chatkalsky Range, valley of riv. Kachansai, leg. Tarbinsky,
7.vii.968]; 2 W, "Тянь-Шань, Чаткальский хр., ур. Ак-Куль, Тарбинский, 7.VII.968" [Tien-Shan,
Chatkalsky Range, ur. Ak-КиГ, leg. Tarbinsky, 7.VÜ.968] (MOSCOW, PETERSBURG); lectotype of
M. lobicornis subsp. foreliella (designated here), w, "Харог, Гунт+Шахдара, Шунган, Казнак., 11.VIII.97"
[Khorog, Gunt + Sakhdara, Shungan, leg. Kaznakov, 11.viii.97], "M. rubra r. lobicornis Nyl. det.
Forel", "Lectotype M. lobicornis foreliella Arn." (MOSCOW); paralectotypes: 2 w (MOSCOW) and
3 w (PETERSBURG) with the same labels as lectotype; non-type material: > 30 w, 2 q, 2 m from
Tien-Shan.
Distribution (Map 56). Tien-Shan, Alai, Pamir.
Etymology. M. kirghisorum: named from the generative form of Kirghis, to indicate that
the species originates in Kirghisia (Kyrgyzstan). M. foreliella: from Latin ella diminutive of
the name Forel, named for Auguste Forel (see biography in Chapter 1.6).
Notes. M. kirghisorum belongs to the lobicornis species group. It well differs from
M. lobicornis by its distinctly wider frons and generally much smaller, often dentiform lobe
at the base of its scape; additionally, males have a scape that is gradually curved at the base,
not strongly curved as in M. lobicornis. It differs from other lobicornis-group species that
have a developed vertical dent on the scape (except for M. forcipata) by its subtriangular
petiolar node (seen in profile), and it well differs from M. forcipata by its straight (not
curved inwards) propodeal spines.
Arnoldi (1976) did not definitely give the locality (or any other data) for the holotype
of M. kirghisorum, although we can infer that it was probably from the same series as the
12 paratypes collected by him from the Fergansky Range, Kara-Alma. However, we can not

Map 56. Distribution of M. kirghisorum.

168
locate any specimen in the collection of MOSCOW and PETERSBURG labelled as the holo­
type. On the other hand, Arnoldi did not designate a holotype for subsp. foreliella, there­
fore we have designated a lectotype specimen of this form. The main character that Arnoldi
(1976) used to distinguish M. kirghisorum from subsp. foreliella was the distinctly longer
head of the latter (CI 1.18-1.23, according Arnoldi). However, when we studied the type
specimens of subsp. foreliella we found that the head was relatively shorter than suggested
by Arnoldi (CI 1.14-1.18) and fully overlapped the data for the M. kirghisorum types
studied, in fact the heads of some specimens of the latter were longer (max CI 1.21). We
could not find any consistent difference using other features (e.g. shape and sculpture of
alitrunk and waist, body colour, etc.), which are somewhat variable in both forms, therefore
we conclude that subsp. foreliella can not be discriminated from M. kirghisorum and we
confirm the synonymy.
Ecology. M. kirghisorum is a mountain species, living at altitudes 1500-3000 m a.s.l.
where it inhabits mountain steppes and subalpine meadows, nesting in soil, often under
stones. Nothing else is known of its ecology. ;

Myrmica koreana Elmes, R a d c h e n k o et Kim, 2 0 0 1

(Figs 134-136)

Myrmica koreana Elmes, Radchenko et Kim B.-J., 2001: 108, w, Korean Peninsula; Seifert 2003: 153;
Radchenko 2005: 141; Radchenko, Elmes and Alicata 2006: 521, q, m; Pfeiffer et al. 2007: 5.
Myrmica schencki: Gee 1924: 102; Ruzsky 1936: 94; Kupyanskaya 1986b: 87, 1990: 109 {part., only
material from East Siberia and Russian Far East), misidentificatioris.

Type localities and type specimens. Korean Peninsula, for details see material examined.
Material examined. Holotype, w, [K-100 (9)], SJ Korea, unknown location, in pitfall traps, leg. B -
J. Kim (LONDON); paratypes: 13 w, [K-100 (1-8, 10-14)] same series as holotype; 1 w, [K-99 (1)]
S. Korea, Gechon-ri, nr. Pyongchang, Pyong chang-gun. In pitfall traps с.1970, leg. B.-J. Kim; 4 w, [K-
115 (1, 4, 5, 7)] S. Korea, mountains, locality not known, leg. B.-J. Kim; 1 w, [K-122 (1)], S. Korea,
mountains, locality not known, leg. B.-J. Kim; 7 w, [¡K-118 (2, 5, 7-11)], S. Korea, Sunchon City, 300
m. in soil. 2.vii.1982, leg. B.-J. Kim; 4 w, N. Korea, Pyongyang, 21.vii.1959, leg. B. Pisarski and
J. Prószyñski, Inst. Zool. P.A.N. Warszawa 58/59 (LONDON, IKSAN, ELMES, KIEV, WARSAW);
non-type material: > 100 w, 20 q and 19 m from South Siberia, Mongolia, Russian Far East and North
Korea. !
Distribution (Map 57). South-eastern part of West Siberia, northeastern Kazakhstan
(to the south until Saur Range), South Siberia, Mongolia, Russian Far East, northeast of
China and Korea. j
Etymology. M. koreana: named for Korea where we believed it to be endemic when we
first described it, however since then it has been shown to be widespread in Siberia and the
Far East. j
Notes. M. koreana belongs to the schencki species group and most resembles
M. schencki. Female castes differ from the latter by their distinctly wider frons and much
less extended frontal lobes; queens are considerably smaller than those of all other known
queens of the schencki-group species. Males' differ from those of M. schencki by their
shorter and much less curved scape.
Records of M. schencki (Ruzsky 1936; Kupyanskaya 1986b, 1990) from East Siberia
and the,Russian Far East actually belong to M. koreana (corresponding material examined).

169
 Ecology. M. koreana inhabits mainly steppes and steppe-like habitats both on planes
and mountains up to 1700 m a.s.l., only rarely it has been found living in light deciduous
and mixed forests. Nests are normally built in soil, with no obvious indication of the nest
entrance other than small woven entrance funnels, these are very similar to, but perhaps
somewhat shorter than those of M. schencki. Nuptial flights are in August-September.

Map 57. Distribution of M. koreana.

Myrmica kotokui Forel, 1911

(Figs 137-139)

Myrmica rubra subsp. kotokui Forel, 1911a: 267, w, q, m, Japan and Russia; Weber 1947: 449.
Myrmica ruginodis subsp. kotokui: Terayama et al. 1992: 25.
Myrmica kotokui: Collingwood 1976: 300; Radchenko 2005: 142, nec Imai et al. 2003: 182; Kikuchi
et al. 1999: 252 (part), misidentifications (see also Notes below).
Myrmica ruginodis subsp. orientalis Karawajew, 1926a: 65, w, Russia; Kuznetsov-ugamsky 1928: 37,
q, m. Synonymy by Radchenko 1994e: 73 (as synonym of M. ruginodis); Radchenko 2005: 142
(as synonym of M. kotokui); the latter confirmed here.
Myrmica orientalis: Kupyanskaya 1990: 101 (part.); Bolton 1995: 281.
Myrmica laevinodis: Ruzsky, 1920: 76, misidentification.
Myrmica rubra: Onoyama 1989: 131; Imai et al. 2003: 182, misidentifications.

Type localities and type specimens. M. kotokui: "Nördliches Japan (Dr. Haberer)". M. ruginodis
orientalis: "30 km nordwärts von Wladivistok, W. Fedynsky, 1 w".
Material examined. Lectotype of M. kotokui, w (designated by Radchenko 2005), "Typus", "Nöral
(? Nösal), Japan, D. Haberer", "M. rubra L. r. kotokui typ. Forel", "Coli. A. Forel" (BASLE); para-
lectotype, w, "Typus", "Iturup, Kurilen, D. Haberer", "M. rubra L. r. kotokui Forel, typ.", "Coll.
A. Forel" (BASLE); holotype of M. ruginodis subsp. orientalis, w, "30 вер. к сев. от Владивостока,
Фсдинский" [30 km N Vladivostok, leg. Fedinsky], "3044 Coll. Karawajewi", "M. (Myrmica) rubra (L.)
ssp. orientalis Kar. Karawajew det.", "Holotypus Myrmica ruginodis orientalis Karaw." (KIEV); non-
type material: > 100 w, several tens q and m from Russian Far East, Korean Peninsula and Japan.

17O
 Distribution (Map 58). Russian Far East, Korean Peninsula, NE China and Japan.
Etymology. M: kotokui: Forel does not say to whom he dedicated this species. Most
probably he had in mind Shusui Kotoku, a well known Japanese socialist, journalist and au-
thor, who was hanged six months earlier for treason, on 24th January 1911, and was con-
sidered by many to be a martyr for peace and,¡freedom. M. orientalis: name from the Latin
orientalis = eastern, to indicate that it is the eastern form of M. ruginodis.
Notes. Collingwood (1976) raised this form to species rank. However, specimens
from Sam-zi-yan (North Korea) determined by him as M. kotokui are in fact M. kurokii
(material is in BUDAPEST, examined).
M. kotokui belongs to the rubra species group and is very difficult to distinguish from
M. ruginodis. Female castes differ from M. ruginodis mainly by the shape and sculpture of
the petiole: the petiolar node dorsum (seen ¡n profile) is slightly more rounded without
a distinct dorsal plate, and the sides of the petiolar node are smoothly sculptured whereas
M. ruginodis has coarse longitudinal rugosity (in these respects it more resembles M. rubra).
In males of M. kotokui the clypeus is distinctly longitudinally striated while in males of
M. ruginodis it is smooth.
Examination of the holotype worker of Myrmica ruginodis subsp. orientalis confirmed
that this form is synonym of M. kotokui arid is not the oriental form of M. ruginodis.
Examination of specimens of M. ruginodis from across its range indicates the existence of
an east-west cline in morphological variation (see notes to that species). Kupyanskaya's
(1990) M. orientalis in fact included both "oriental" M. ruginodis and M. kotokui. In the
monograph on Japanese ants (Imai et al. 2003) M. ruginodis is identified as "M. kotokui"
(the name M. ruginodis being absent from that book) while M. kotokui is considered under
the name "M. rubra". We are sure that M. rubra is absent from the orientai region (apart
from a possibility of accidental introductions).
We think that the cline in features of M. ruginodis and some similarities between
M. kotokui and M. rubra has led to these names becoming confounded with each other in
Japan. Similarities in the ecology of M. kotokui and M. rubra combined with a similar dif-
ference in petiole shape compared to M. ruginodis has led authors to consider some popu-
lations of M. kotokui as M. rubra. For example, specimens from Japan determined by
Onoyama (1989) as M. rubra are M. kotokui (material kindly sent us by K. Onoyama,
examined). A further confusion has developed over the last few decades from reports of the
microgyne form of M. ruginodis from Japan (see notes to M. ruginodis), probably most re-
ports of polygynous populations of "M. ruginodis" (inferred to be race microgyna) are in fact
M. kotokui. For example, in Kikuchi et al. (1999) the material from the polygynous colonies
belongs to M. kotokui and the other samples are M. ruginodis (material examined).
Ecology. M. kotokui is one of the commonest Myrmica species in East Asia, living in
various habitats. It prefers different kinds of forests (pine, oak, maple, birch, lime, spruce,
fir, larch), it occurs also in shrublands and meadows, but avoids dry places. In Japan on
Honshu Island we found it living at altitudes of 1200-2000 m. a.s.l., further north on
Hokkaido Island it lived at lower altitudes from sea level to about 1000 m. It builds nests
predominantly in the soil, very often under stones, but also constructs soil mounds or nests
in decayed wood. In Japan nests were usually found under "Bamboo grass" or in open
deciduous forest. In contrast to the closely related M. ruginodis, colonies of M. kotokui are
quite polygynous, with population of the nests reaching several thousand workers (Kikuchi
et al. 1999). Nuptial flight occurs in August-September.

171
 Map 58. Distribution of M. kotokui.

Myrmica kozakorum, sp. nov.

(Figs 140-142)

Myrmica georgica Seifert, 1987: 183 (part, only w and m from Kazakhstan; see also Notes to
M. georgica, above).

Material examined. Holotype, w, Ukraine, Kherson Prov., Black Sea Natural Reserve, Ivano-
Rybalchansky Distr., 54-04, l.ix.2004, leg. A. Radchenko (KIEV); paratypes: 15 w, 5 m from the nest
of holotype; 5 w, same locality and collector, 21-81, 17.vi.1981; 5 w, 1 q, same locality and collector,
26-81, 17.vi.1981; 5 w, same locality and collector, 65-81, 22.vi.1981; 3 w, same locality and collec-
tor, 35-82, 6.V.1982; 55 w, same locality and collector, 44-82 (= UA-13), ll.v.1982; 5 w, same
locality and collector, 100-82, 9.VÌ.1982; 3 w, same locality and collector, 112a-82, 12.V.1982; 1 q,
same locality and collector, 262-82, 24.viii.1982; 15 w, same locality and collector, 17-83, 13.iv. 1983;
5 w, same locality and collector, 28-83, 23.iv.1983; 15 w, same locality and collector, 21-83, 15.iv.1983;
3 w, same locality and collector, 23-83, 18.iv.1983; 3 w, same locality and collector, 24-83, 3.V.1983;
15 w, same locality and collector, 33-83, 25.V.1983; 10 w, 1 m, same locality and collector, 11.ix.1997;
22 w, 8 m, lq, same locality, UA-86, 10.ix.1997, leg. G. W. Elmes; 13 w, 1 m, 11 q, same locality, date
and collector, UA-87; 33 w, 7 m, lq, Ukraine, Kherson Prov., Black Sea Natural Reserve, Solenoozerny
Distr., UA-81, 11.ix.1997, leg. G. W. Elmes; 29 w, lm, same locality, data and collector, UA-82; 15 w,
same locality, 53-04, 1.ix.2004, leg. A. Radchenko; 5 w, Ukraine, Kherson Prov., Black Sea Natural Re-
serve, Solenoozerny Distr., 49-99, 28.vi.1999, leg. A. Radchenko; 6 w, Ukraine, Kherson Prov, Black
Sea Natural Reserve, UA-123, 7.v.2001, leg. L. Rusina; 10 w, 5 m, 5 q, Ukraine, Kherson Prov., Tsu-
rupinsky Distr., Radensky Forest, 312-88, 24.viii.1988, leg. A. Radchenko; 10 w, Ukraine, Donetsk
Prov., Natural Reserve Kammenyc Mogily, 216-83, 14.vi.1983, leg. A. Radchenko; 0 w, same locality
and collector, 234-83, 15.vi.1983; 5 w, Ukraine, Donetsk Prov., Natural Reserve Khomutovskay Step',
5.3, 4.VÌ.1976, leg. V. Vakarenko; 5 w, same locality, 263-83, 18.vi.983, leg. A. Radchenko; 25 w,
Ukraine, Lugansk Prov., Natural Reserve Streltsovskay Step', 363-83 (= UA-17); 20 w, 27.vi.1983,
leg. A. Radchenko; Ukraine, Odessa Prov., shore of Khadhzybeisky Liman, 435-83 (= UA-14),
3.viii.1983, leg. A. Radchenko; 1 w, Russia, Kalmykia, Svetlyj Erik, 21.v.2001, leg. ]. Savranskaya;
1 w, Russia, 20 km N Orenbugr, 3.vi.985, leg. A. Petrenko; 5 w, 1 q, Kazakhstan, Borovoie, 28.viii.1967,

172
leg. Malozemova; 1 w, 1 m, Kazakhstan, Borovoie1, Mt. Sinjukha, 68-50, 15.viii.1968, leg. Maloze-
mova; 1 q, 1 m, Kazakhstan, Borovoie, 68-56, 27.viii.1968, leg. Malozemova; 3 w, [Kazakhstan],
Borovoie, 18.v.1985; 4 w, same locality, 2.v. 1965; leg. K. Arnoldi; 1 w, [Kazakhstan], Aktiubinsk
Prov., riv. Aulya, 6.VÌ.1985, leg. V. Ermolenko; 3 w,|Kazakhstan, Saur, Usgan Obo, 47°02'N, 84°54'E,
1650 m, 2001, leg. R. Schutz; 3 m, [Kazakhstan], Kokchetav Prov., Schuchinsk, 40, 18.viii.1966, leg.
Malozemova ("paratypes" of M. georgica) (KIEV, MOSCOW, ELMES).
Description of workers (Fig. 140). |
Head somewhat longer than broad, with slightly convex sides and occipital margin, and
widely rounded occipital corners. Anterior clypeal margin rounded, not prominent and not-
notched medially. Frontal carinae moderately curved, frons relatively narrow (narrower
than in M. scabrinodis); antennal sockets surrounded by fine concentric striation; frontal
lobes moderately extended. Scape shorter than head-width, sharply curved at the base (ap-
proximately a right angle), with well developed, quite large horizontal lobe (somewhat
similar to that of M. sabuleti). Promesonotal dorsum convex, promesonotal suture indistinct
(seen from above). Metanotal groove distinct, though not very deep. Propodeal spines
relatively long, straight, sharp, widened at the base, directed backward at an angle about 45°.
Petiole with distinct peduncle, its anterior surface strongly concave, dorsum of node with
inclined posteriorly dorsal plate. Postpetiole with convex dorsum, higher than length. Spurs
on middle and hind tibiae often reduced to different extents.
Frons with slightly sinuous longitudinal rugae, numbering < 15 between frontal carinae
level with the eyes; occiput and temples with! reticulation, surface between rugae at most
with very fine superficial microsculpture, appearing shiny; central part of clypeus with re-
duced rugosity, smooth and shiny; frontal triangle smooth and shiny; mandibles longitudi-
nally rugose. Alitrunk dorsum with sinuoùs rugulosity, pronotal dorsum also with
reticulation, sides of alitrunk with longitudinal, slightly sinuous rugae. Surface between
rugae on alitrunk smooth and shiny. Petiolar dorsum with short longitudinal sinuous rugae,
postpetiole with longitudinally-concentric rugosity, surface between rugae at most with very
fine superficial microsculpture, appearing shiny. Gaster smooth and shiny.
Head margins with quite long subdecumbent to suberect hairs. Alitrunk with erect hairs,
petiole and postpetiole with not numerous longer hairs. Scape with not-numerous subde-
cumbent to suberect hairs, tibiae with subdecumbent ones. Body colour yellowish-brown,
mandibles and appendages somewhat lighter.
Description of queens (Fig. 141).
Generally like worker by all diagnostic features, having somewhat wider frons and less
sinuous rugosity on the alitrunk.
Description of males (Fig. 142).
Head with convex sides and occipital margin, and very widely and gradually rounded oc-
cipital corners; its length approximatey equal to width, usually slightly longer, but some-
times shorter than width; anterior clypeal margin widely rounded, not prominent and
not-notched medially. Antennal scape quite short (similar to that of M. specioides), anten-
nae 13-segmented, with 5-segmented club; second funicular segment approximately 1.3
times longer then third one. Alitrunk relatively long, scutum convex, scutellum does not
project dorsally above scutum when seen in profile. Propodeum with blunt, thick subtrian-
gular teeth. Petiole with distinct peduncle, distinctly longer than height, its anterior sur-
face almost straight; postpetiole higher than long, with weakly convex dorsum.
Head dorsum densely punctated, clypeus mostly smooth, appearing shiny. Central part
of scutum in front of Mayrian furrows smooth and shiny, the rest of scutum, scutellum and

173
propodeal dorsum longitudinally rugulose. Sides of pronotum and posterior part of meso-
pleura longitudinally rugulose. Petiolar node with a few longitudinal rugulae, postpetiole
appearing smooth, surface of both petiole and postpetiole smooth and shiny.
Head margins and mandibles with numerous, relatively long, curved suberect hairs.
Alitrunk, petiole and gaster with sparser, but longer hairs. Tibiae and tarsi with relatively
short subdecumbent to suberect hairs (like M. specioides), the longest hairs on tibiae shorter
than the maximal tibial width, those on basitarsus subequal or slightly longer than its
maximal width. Scape and first 7 funicular segments with relatively long suberect hairs,
segments of antennal club with very short subdecumbent hairs. Body colour dark brown to
black, appendages somewhat lighter.
Distribution (Map 59). Steppes from southern Ukraine to West Siberia and Kazakhstan.
Etymology. M. kozakorum: from the generative of the name Kozak (Cossack) to indi-
cate that it is from the Kozak steppes of Ukraine and southern Russia.
Notes. Seifert (1987) described M. georgica based on several tens of workers collected
in Georgia, and on 3 workers and 2 males, collected in northern Kazakhstan. We be-
lieve now that the type series of Seifert's M. georgica include two species (see Notes to
M. georgica, and Radchenko and Elmes 2004): "true" M. georgica (workers from Georgia,
including the holotype) and another species (the "paratype" workers and males from
Kazakhstan). We include the latter to the species described here, M. kozakorum.
We place M. kozakorum to the speciodes-complex of the scabrinodis-group based on the
combination of the features of female castes and males. The workers have a mixture of mor-
phological features associated with different species in the scabrinodis-group: by the shape
of the lobe on the base of their scape, they most resemble M. sabuleti (perhaps being some-
what intermediate with M. scabrinodis), by the width of their frons they are more similar
to M. scabrinodis, the shape of their propodeal spines and petiole more resembles that of
M. specioides, while the spurs on the middle and hind tibiae are often reduced, similar to

Map 59. Distribution of M. kozakorum.

174
that of M. tuliпае. This would make a single wjorker very hard to identify with high confi­
dence. Fortunately the males of M. kozakorum well differ from those of M sabuleti by their
much shorter scape, and differ from M. scabrinodis and M. tulinae by the much shorter
hairs on their tibiae and tarsi; however, they are hardly distinguishable from the males of
M. specioides (see also Key, Chapter 5.2).
Ecology. Although M. kozakorum is a species of the Steppe Zone, within that biome it
is normally found in intrazonal, relatively wet arid shaded-places (small woods, meadow-like
associations around lakes, etc.). It appears to be fairly tolerant of high soil salinity often
being found in scrub on the margins of salt lakes, however it does not appear to have evolved
a highly adapted behaviour to such habitats as for example is seen in M. bergi (see Notes to
that species). We suspect that it forages in the patches of shaded vegetation and does not
compete with M. bergi in the fully exposed open conditions. It is most common in the small
relatively open oak woods that develop in shallow depressions in the steppe, and on the
edges of larger lakes, these become flooded in periods of high rainfall and the shade helps
retain the soil moisture. Here colonies are quite small, at most a few hundred workers, and
nests are usually built in the soil under small rotten branches, pieces of bark or even leaves.
The nuptial flight is in August-September.

Myrmica kozlovi] Ruzsky, 1915

(Fig. 143)

Myrmica kozlovi Ruzsky, 1915: 435, w, Tibet; Emery 1921: 37; Menozzi 1939: 294; Weber 1947: 470;
Collingwood 1970: 374; Radchenko 1994b: 42; Bolton 1995: 280; Radchenko and Elmes 2001b:
259. |
Myrmica kozlovi subsp. mekongi Ruzsky, 1915: 437, w, Tibet; Emery 1921: 37; Menozzi 1939: 294;
Weber 1947: 471; Bolton 1995: 281, syn. nov. j
Myrmica kozlovi subsp. subbrevispinosa Ruzsky, 1915: 437, w, Tibet; Emery 1921: 37; Menozzi 1939:
294; Weber 1947: 472; Bolton 1995: 284, syn. nov.
Myrmica kozlovi subsp. subalpina Ruzsky, 1915: 438, w, Tibet; Emery 1921: 37; Menozzi 1939: 294;
Bolton 1995: 283, syn. nov.
Myrmica kozlovi subsp. ruzskyi Weber 1947: 471 (unresolved junior primary homonym of Myrmica
kozlovi subsp. ruzskyi Kiseleva, 1925: 76 and!not necessary proposed replacement name for
Myrmica kozlovi var. subalpina Ruzsky, 1915: 438; see also Notes io Myrmica kurokii). Synonymy
by Bolton 1995: 282 (as synonym of M. kozlovi var. subalpina); synonym of M. kozlovi: syn. nov.
(see Notes below).
Myrmica specularis Donisthorpe, 1929: 446, w, Tibet; Menozzi 1939: 293; Weber 1950: 224; Chapman
and Capco 1951: 130. Synonymy by Radchenko and Elmes 2001b: 259; confirmed here.

Type localities and type specimens. M. kozlovi: "Мсстонахожд.: 1) прит. р. Дза-чю, Кам, басе. р.
Голубой, 12-12000 и 13000', нач. V. 1901 (ww); 2) дол. р. Голубой, Кам, восточн. Тибет, III. 1901 (ww),
(Козлов)" [Localities: 1) tributary of riv. Dza-chju, Kam, basin of riv. Yangtze, 12-12500 and 13000
feet, beginning of v.1901 (ww); 2) valley of riv. Yangtze, Kam, eastern Tibet, iii.1901 (ww), leg. Ko­
zlovi. M. kozlovi subsp. mekongi: "Найдена в восточном Тибете, в верховьях р. Меконга. Мсстонахожд.:
р. Бар-чю, басе. р. Меконга, Кам, 12.000', кон. IX. ÍJ900 (рабочие), (Козлов)" [Found in the Eastern
Tibet, in upper reaches of riv. Mekong. Localities: riv. Bar-chju, basin of riv. Mekong, Kam, 12,1000',
end of ix.1900 (workers), leg. Kozlov]. M. kozlovi subsp. subbrevispinosa: "В коллекции П. К. Козлова
нашелся один экземпляр раб. из восточнаго Тибета. Мсстонахожд.: дол. р. Голубой, Кам, вост. Тибет, III.
1901 (рабочий), (Козлов)" [In the collection of Р. К. Kozlov was found one specimen, worker from

175
Eastern Tibet. Locality: valley of riv. Yangtze, Eastern Tibet, iii. 1901 (worker), leg. Kozlov]. M. kozlovi
subsp. subalpina: "Мсстонахожд.: дол. р. Голубой, Кам, вост. Тибет, III. 1901 (рабочий) (Козлов)" [Lo­
cality: valley of riv. Yangtze, Kam, Eastern Tibet, iii.1901 (worker), leg. Kozlov]. M. speculans: "De­
scribed from eight workers, Tibet, Gautsa, at a height of 13,000 ft., on April 5 th , 1924 (Everest
Expedition). There are also ten specimens in the British Museum Collection taken at Khamba Jong,
Sikkim, 15-30.vii.03, at a height of 15,000-16,000 ft. (Tibet Expedition, 1903), which agree quite well
with the above species with the exception of being not quite so dark. Type and paratypes in the British
Museum Collection".
Material examined. Lectotype of M. kozlovi, w (designated here), "пр. р. Дза-чю, Кам, Голубая, 12-
13000', Козлов, нач. III.01" [tributary of riv. Dza-chju, Kam, riv. Yangtze, 12-13000', leg. Kozlov, be­
ginning of iii.01] (PETERSBURG); paralectotypes: 34 workers with same labels as lectotype; 27 w,
"дол. Голубой р., Кам, в. Тибет, Козлов, III.01" [valley of riv. Yangtze, Kam, Tibet, leg. Kozlov, iii.01];
1 w, "p. Дза-чю, 11000', Кам, бас. Голубой, Козлов, сер. IV.01" [riv. Dza-chju, 11000', Kam, basin of riv.
Yangtze, leg. Kozlov, middle of iv.01]; 10 w, "p. Дза-чю, Голубая, 12-13000', Козлов, нач. V.Ol" [riv.
Dza-chju, riv. Yangtze, 12-13000', leg. Kozlov, beginning of v.Ol] (PETERSBURG, MOSCOW, KIEV);
lectotype of M. kozlovi subsp. mekongi (designated here), w, "р. Бар-Чю, 12000', бас. Меконга, Кам,
Козлов, кон. IX.00" [riv. Bar-chju, basin of riv. Mekong, Kam, end of ix.1900, leg. Kozlov], "Myrm. ko­
zlovi sub. mekongi n. sub. M. Ruzsky" (MOSCOW); paralectotypes: 15 w with the same labels; 1 w,
"речка Ба-чю (Sic\), 12 т' Кам, б. Голубой р. Козлов 2-3.viii.00" [riv. Ba-chju (Siel), 12,000' Kam, basin
of riv. Yangtze, leg. Kozlov, 2-3. viii. 00] (MOSCOW, PETERSBURG, KIEV); holotype of M. kozlovi
subsp. subbrevispinosa "дол. Голуб. Р., Кам, в. Тибет, Козлов, III.01" [valley of riv. Yangtze, Kam,
Eastern Tibet, leg. Kozlov, iii.01], "M. kozlovi v. subbrevispinosa n. var." (MOSCOW) (see also Notes
below); holotype of M. specularis, w, "Tibetan side of the Mt. Everest: Tibet, Gautsa, 13000 ft, 5.ÌV.1924
(Hingston)" (LONDON); paratypes: 4 workers with same labels (LONDON); non-type material:
2 w, SE Tibet, Dzogang, 9-14,000 ft, l-21.ix.1936 (Tibet expedition, 1903); 7 w, India, Sikkim,
Khamba Jong, 15-16,000 ft, 15-30.vii.[19]03 (LONDON).
Distribution (Map 60). Tibet, India (Sikkim), Nepal.
Etymology. M. kozlovi: dedicated to Colonel Peter Kuzmich Kozlov (1863-1935), the
outstanding Russian geographer and explorer, head of many expeditions to Central Asia
(Mongolia, China, Tibet). M. mekongi: named after its type locality in Tibet, the upper
catchment basin of the great river Mekong, which flows from Tibet to reach the sea in Viet-
nam. M. subbrevispinosa: from the Latin brevis = short and spinosa (F) = thorny with pre-
fix sub - under, to presumably to indicate fairly short (but not extremely short) propodeal
spines. It has no connection with the American species M. brevispinosa Wheeler, 1917.
M. subalpina: from the Latin prefix sub = under or beneath or close and alpina = of the
mountains, to indicate that it is a species of the sub-alpine habitat zone. M. specularis:
Donisthorpe gave no indication as to why he chose this name which could have several
roots: it could be derived from the Latin specula = "high place for observing" which might
be related to the mountain habitat; or from the Latin specus = chasm, because Donisthorpe
indicated the postpetiole "with somewhat deep longitudinal pits"; but most probably it is
from the Latin specularis = "of a mirror" because Donisthorpe wrote that the surface be-
tween the propodeal spines is "smooth and shining" and he emphasised this feature in his
comparison of the species with M. ruginodis.
Notes. Originally, Radchenko (1994b) placed M. kozlovi in the rugosa species group
but now we believe it belongs to the kurokii-group (Radchenko and Elmes 2001b).
Ruzsky (1915) described M. kozlovi from Eastern Tibet based on a reasonably large
sample of workers, at the same time he described subspecies mekongi from a series of
workers, while basing his descriptions of subspecies subbrevispinosa and subalpina on
a single worker of each. We first compared the type specimens of M. kozlovi with those of

176
subsp. mekongi and found that workers of subsp. mekongi slightly differ from M. kozlovi
by a more finely rugulose petiole and postpetiole, by a shorter scape (mean Slj 0.76 and SI2
0.86 vs. 0.80 and 0.90), and by a somewhat lighter colour. Although Ruzsky noted only one
specimen of subsp. subbrevispinosa (see above), we located 2 additional workers in the
MOSCOW collection. These have the same labels as the holotype and are practically indis-
tinguishable from it, and. could be treated as páratypes even if formally they do not belong
to the type series. All these 3 specimens of subsp. subbrevispinosa have shorter propodeal
spines than M. kozlovi (ESLI 0.23-0.26 vs. 0.30-0.34), but at the same time these data
overlapped with subsp. mekongi (ESLI 0.25-0.3:1). We could not find the holotype of subsp.
subalpina and it is probably lost. Based on the original description of the single worker it
differs from M. kozlovi by the shape of its propodeal spines that are widened at the base and
curved downwards (see Ruzsky 1915, fig. 23). We also examined the holotype and paratypes
of M. specularis that differ from the types and non-type material of M. kozlovi only by
a somewhat darker head, a slightly more massive petiolar node and by slightly coarser and
more regular rugosity of the petiole and postpetiole.
From these studies we conclude that M. kozlovi is probably widespread in the Himalayan
mountain system and isolated populations predictably can show morphological variations.
Generally such minor differences between workers are insufficient to separate species; far
more material, including sexual castes, would need to be collected and studied to confirm
whether any of these forms can be considered as good subspecies or species. For now, we
consider it best that all the above forms remain! synonymised with M. kozlovi.
Weber (1947) considered the name subalpina Ruzsky, 1915 as a junior primary homo-
nym of Myrmica rubra subsp. brevinodis var. subalpina Wheeler W. M., 1907 and proposed
the replacement name - M. kozlovi subsp. ruzskyi (that was itself a junior primary homo-
nym of M. kozlovi subsp. ruzskyi Kiseleva, 1925 - see notes to M. kurokii). However,
Wheeler's (1907) name "subalpina" is unavailable (quadrinomen), and the first available
use of this name is Myrmica brevinodis var. subalpina Wheeler W. M., 1917, which post-
dates Ruzsky 1915. Hence, it was not necessary for Weber to proposed a replacement name

Map 60. Distribution of M. kozlovi.

177
for subsp. subalpina Ruzsky, since according to the last edition of the International Code
of Zoological Nomenclature (1999) M. brevinodis var. subalpina Wheeler, 1917 is pre­
occupied name (see also Bolton 1995).
.-Ecology is poorly known. M. kozlovi has been found at the highest altitudes recorded for
any Myrmica species worldwide (4800 m) which probably accounts for it being the only
Myrmica species that has crossed the Himalayan barrier to live on both the western and
eastern slopes. Donisthorpe (1929) wrote that "According to Major Higston [the collector
of M. specularis] the ants found at the higher elevations [in the Himalaya] are exceedingly
lethargic and sluggish in their movements.".

Myrmica kryzhanovskii Arnoldi, 1976

(Fig. 144)

Myrmica kryzhanovskii Arnoldi, 1976: 551, w, Tajikistan; Radchenko 1994b: 41, 1994c: 137, 1994e:
74; Bolton 1995: 280; Radchenko and Elmes 2003b: 7.

Type localities and type specimens. "Хребты Таджикистана. Голотип ? (see Notes below) и 12
паратипов, Кулябская обл., Шорак (Валиахмсдов). Гнсеарекий хребет, Квак (Янушсв) [Mountain ranges
of Tajikistan. Holotype and 12 paratypes, Kulyab Prov., Shorak, leg. Valiakhmedov. Gissarsky range,
Kvak, leg. Yanushev].
Material examined. Holotype, w, "Таджикистан, Шорак, Муминабад. р-н, лес, 1.VI.65 Валиахмсдов"
[Tajikistan, Shorak, Muminabadsky Distr., forest, l.vi.65, leg. Valiakhmedov], "Holotype Myrmica
kryzhanovskii К. Arnoldi" (MOSCOW); paratypes: 1 w, same locality and collector as in holotype,
but date 26.V.65; 9 w, "Кондара, Квак, ореховый лес, пень, 19.7.69, № 128 Янушсв" [Rondara, Kvak, nut
forest, tree stump, 19.vii.69, No 128, leg. Yanushev], "Paratypus Myrmica kryzhanovskii K. Arnoldi"
(MOSCOW); non-type material: 6 w from Tajikistan and southern Kyrgyzstan.
Distribution (Map 61). Tajikistan, Kyrgyzstan.

Map 61. Distribution of M. kryzhanovskii.

178
 Etymology. M. kryzhanovskii: this species was named after Prof. Oleg Leonidovich
Kryzhanovski (1918-1997), a famous Russian entomologist and zoogeographer who
worked at the Zoological Institute, St.-Petersburg.
Notes. We placed M. kryzhanovskii to the dshungarica complex of the rubra species
group (Radchenko and Elmes 2001b). It differs from related species (M. juglandeti,
M. ferganensis) by much longer semierect hairs of the scape and legs.
This species is known only from workers. There was a printing error in Arnoldi's (1976)
paper: the holotype is marked by a symbol " 9 " indicating a queen, whereas in fact it is
a worker. Bolton (1995) also erroneously recorded that queens and males were described,
and wrongly gave its type locality as Russia. |
Ecology is poorly known. It was found in nut; forests and dry meadows at altitudes about
1500 m a.s.l.

Myrmica kurokii Forel, 1907

(Figs 145-147)

Myrmica rubra subsp. kurokii Forel, 1907: 18, w, Japan.

Myrmica kurokii: Emery 1908a: 166, 1921: 37; Weber 1947: 469; Collingwood 1976: 301, 1981: 26;
Kupyanskaya 1979: 128 {part), 1990: 102, q, m\ Radchenko 1994b: 42, 1994c: 137, 1994e: 74;
Bolton 1995: 280; Kim B.-J. 1996: 179, 2003: 2; linai et al. 2003: 183; Radchenko 2005: 143.
Myrmica chinensis Viehmeyer, 1922: 204, q, m, China; Wheeler 1930b: 61.
Myrmica tibetana subsp. chinensis: Weber 1947: 4661
Revived status as species: Radchenko 1994 b: 41; Bolton 1995: 278; syn. nov. (provisionai).
Myrmica helleri Viehmeyer, 1922: 204, w, China; Wheeler 1930b: 61; Eidmann 1941: 13.
Myrmica kurokii subsp. helleri: Weber 1947: 469.
Revived status as species: Radchenko 1994 b: 42; Bolton 1995: 279; syn. nov. (provisionai).
Myrmica kozlovi subsp. ruzskyi Kiseleva, 1925: 76, wj Russia; Bolton 1995: 282, nec Myrmica kozlovi
subsp. ruzskyi Weber, 1947: 471 (see Notes to M. kozlovi, above). Synonymy by Radchenko 1994e:
74; confirmed here.
Myrmica smythiesi var. fortior: Ruzsky 1905: 660, misidentification.
Myrmica kotokui: Collingwood 1976: 300; Kim B.-J. 1996: 179, misidentifications.
Myrmica silvestrii: Collingwood 1981: 26; Kim B.-J. 1996: 180, misidentification.
Myrmica gallienii: Wei С et al. 2001: 561; Chang and He 2001: 26, misidentifications.

Type localities and type specimens. M. kurokii: "Japan (ex coll. Fruhstorfer)". M. chinensis:
"China, Prov. Sztschwan: Sungpanting; Exped. Stötzner 1914". M. helleri: „China, Prov. Sztschwah:
Kwansien; Expect. Stötzner 1914". M. kozlovi subsp.[ruzskyi: "Были найдены только рабочие особи на
каменной сопке, 20 ноября 19 г." [Only workers were found, on stony hill, 20 November 1919] [Ussuri
Region, no precise locality given]. ¡
Material examined. Lectotype of M. kurokii (designated by Radchenko 2005), w, "Typus", "H.
Fruhstorfer vend. 15.1.1902", "Japan, Ex. Coll. Frughstorfer", "M. rubra L. r. kurokii Forel, w type",
"sp. M. Kurokii Forel", "Coll. Forel", "Lectotype Myrmica kurokii Forel. Design. Radchenko et Elmes"
(BASLE); paralectotype, w, "Japan Coll. Fruhstorfer", "Paralcctotypus M. kurokii des. Radch."
(GENOA); non-type material: about 100 w, > 20 q, 15 m from Russian Far East, China, Korean Penin-
sula and Japan.
Distribution (Map 62). Russian Far East, China, Korean Peninsula, Japan.
Etymology. M. kurokii: this species was obviously dedicated to a Mr. Kuroki but Forel
did not say who he was. He might possibly have been a Japanese collaborator of the

179
German collector and lepidopterist Hans Fruhstorfer who owned the material, or more
probably he was the Japanese General Kuroki Tamemoto who in 1904-1905 had become fa-
mous for his part in the Russo-Japanese war. M. chinensis: from a combination of the name
China and the adjectival suffix ensis = place of origin or habitat, to indicate this is a Chi-
nese species. M. helleri: this species was almost certainly dedicated to the Austrian
coleopterologist Karl Borromaeus Maria Josef Heller (1864-1945) who was a Professor and
Section leader in the Staatliches Museum für Tierkunde Dresden where his collection is
maintained. M. ruzskyi: dedicated to the great Russian myrmecologist Michail Dmitrievich
Ruzsky (see biography in Chapter 1.6).
Notes. Radchenko (1994b) placed M. kurokii to the rugosa species group, but now we
believe it belongs to a separate kurokii-group. This large and robust species is superficially
similar to some rubra-group species (e .g. M. ruginodis, M. arisana, etc.), but well differs
from them by having frontal carinae that do not curved outwards and do not merge with the
rugae surrounding antennal socket, a short petiole with weakly developed peduncle, much
denser rugulosity on the head dorsum and by having a densely punctated surface between
rugulae.
Kiseleva (1925) described from the Russian Far East ("Ussury Krai") Myrmica kozlovi
subsp. ruzskyi (see notes to M. kozlovi above). We can not find types of this form and al-
most certainly they are lost. However, M. kozlovi appears to be a Himalayan species with
mainly a Tibetan distribution and there are no other records from the Far East of Russia,
furthermore the original description and figure suggests that M. kozlovi subsp. ruzskyi is
a synonym of M. kurokii (Radchenko 1994e).
Ruzsky (1905) also recorded the Himalayan species M. fortior from the Russian Far
East. M. kurokii is only species from this region that might be confused with M. fortior and
although we could not find any corresponding material in the collections studied, we are
sure that Ruzsky's record should be referred to M. kurokii. Similarly, Collingwood (1976,
1981) determined several specimens of M. kurokii from North Korea as M. kotokui and

Map 62. Distribution of M. kurokii.

180
M. silvestrii, while records of M. gallienii from China (Wei С et al. 2001; Chang and He
2001) should be referred to M. kurokii (corresponding material examined).
We have not seen the type specimens ofj M. chinensis Viehmeyer and M. helleri
Viehmeyer, they were collected by the Walter Stotzner expedition of 1913-15 to China and
if they still exist, they are most probably among the expedition material held by the
Staatliches Museum für Tierkunde, Dresden (the Keeper was searching for these specimens
when the museum was disrupted by the floods of 2002 - we hope this material survived).
We provisionally consider these names as junior synonyms of M. kurokii, based on their dis-
tribution and descriptions and given the known variability of M. kurokii.
Ecology. Inhabits mainly mountain forests (fir, spruce, larch, birch), at altitudes
between 1000 and 2600 m a.s.l., mostly around 1600-2000 m, but also bushes (mainly
juniper), peat bogs, subalpine meadows and mountain tundra. In forests, it builds nests
almost exclusively in rotten wood, but in open habitats the ants construct moss mounds and
build nests in soil under stones or dead wood. Nuptial flight is in August.

Myrmica laurae (Emery, 1907)

(Figs 148 ; 149)

Sifolinia laurae Emery, 1907: 50, q, Italy; Emery 1908c: 550; 1916: 151; Bondroit 1918: 167; Arnoldi
1930: 270; Bernard 1967: 156; Baroni Urbani 197¡1: 75; Kutter 1973: 257, nec Yarrow 1968: 239.
Myrmica laurae: Bolton 1988: 4; Seifert 1994: 17; Bolton 1995: 280; Radchenko and Elmes 2003a: 224.
Myrmica samnitica Mei, 1987: 457, q, m, Italy; Seifert 1994: 17; Bolton 1995: 283. Synonymy by
Radchenko and Elmes 2003a: 224; confirmed here.

Type localities and type specimens. S. laurae: "Siena, un solo esemplare catturato a volo".
M. samnitica: "Materiale tipico. Holotypus: 1 q dealata, Abruzzo, Ovindoli (AQ), M. Magnola, 1400
m, 15/VIII/83, M. Mei leg in nido Myrmica sabuleti Mein, (nido A); l'antenna e la zampa anteriore de-
stra montate su cartellino distinto sullo stesso spollo, le zampe media e posteriore destra montate su
vetrino in Euparal, sigla M03. Paratypi: 17 qq dealate stessi dati dell'holotypus (nido A); 2 q alate, 1
m, stessi dati dell'holotypus (nido B); 14 qq alate, 3 mm, stessi dati dell'holotypus (nido C); 1 q dea-
lata, Abruzzo, Ovindoli (AQ), M. Magnola, 1400 m, 7/ÍVIII/82, M. Mei leg. (nido D). I genitali di 2 mm
(nido C) sono stati montati su vetrino in Euparal, sigla MOI, M02".
Material examined. Holotype of S. laurae, q (alate J without postpetiole and gaster), "Siena, Coggi",
"Typus", "Sifolinia Laurae Emery Typ.", "Holotypus Sifolinia laurae Emery" (GENOA); paratypes of
M. samnitica, 4 q, "Abruzzo, Ovindoli (A Q). M. Mangóla, 15.viii.1983, 1400 m., leg. M. Mei, nido C",
"Paratypes Myrmica samnitica det. M. Mei, 1984"; 1 q, 1 m, same labels, but "Nido B" (MEI, ESPA-
DALER); non-type material: 8 q, 1 male, [Italy] Lazio, Rieti Tarano, Vocabole Campana, 13.ix.1987,
leg. M. Mei (with w, q and m of host species, M. scabrinodis); 2 q, 4 m, [Italy], Abruzzo, Collelongo,
Prati di S. Elio, 13.x.1990, leg. M. Mei (with workers of M. scabrinodis).
Distribution (Map 63). Central Italy.
Etymology. M. laurae: this species was dedicated to Laura Coggi, the baby daughter of
Prof. Alessandro Coggi of Siena, who collected and gave the type specimen to Emery.
M. samnitica: from the name Samnia with the adjectival suffix for nouns ica (from the
Greek то) = belonging-to or from, to indicate this species was found in the region of cen­
tral Italy occupied by the pre-Roman tribe, the Samnites.
Notes. M. laurae differs from most other socially parasitic Myrmica species by a combi­
nation of the following features: it is generally a very hairy species; eyes in both sexes with

l8l
conspicuous hairs; cell 1 +2r on forewings closed and not partly separated by a short vein;
males with 12-jointed antennae, antennal scape long and slender, feebly curved at the base;
antennal scape of queen not angled, gradually curved at the base, but with a distinct nar-
row longitudinal ridge. This form of scape structure that is unique for any known Old World
Myrmica species and is quite different from that of M. hirsuta which M. laurae generally
superficially resembles.
Based on the features of the queen, M. laurae could be placed to the scabrinodis species
group, but we place it in the monotypic laurae-group because its males differs significantly
from the diagnostic characters for that group: they have a much longer scape (SIj > 0.75)
and 12-segmented antennae. However, based on a molecular phylogeny, Jansen et al. (2010)
placed M. laurae within the scabriiwdis-gvoup clade that appears to be one of the more
ancient species groups in Myrmica (estimated age > 20 Ma). M. laurae itself appears to be
quite an old species that diverged from the ancestral species of its host (M. scabrinodis and
/or M. sabuleti) about 10 Ma. If Jansen's results are borne out in subsequent analyses then
M. laurae could be considered as representing "transition" form in evolution of the female
scape in the scabrinodis-group. On the other hand, the long antennal scape of males com-
bined with the reduced number of funicular segments is the clearly an apomorphic charac-
ter compared to other scabrinodis-group species. We suppose that M. laurae evolved the
long male scape sometime during its long history probably as an adaptation finding mates
(see also Radchenko and Elmes 2001b).
Mei (1987) noted that M. samnitica is very similar to S. laurae but in the latter species
the spurs on middle and hind tibiae completely absent. However, examination of the holo-
type of M. laurae showed the middle and hind tibiae had distinct, but greatly reduced, short
spurs with no pectination. Examination of paratype queens of M. samnitica and other
specimens collected by Mei from Lazio and Abruzzo (see above), showed that individuals
varied greatly in tibial spur development, ranging from almost fully developed and
pectinate to short and non pectinate. Thus, we could find no consistent morphological
feature to separate M. samnitica queens from M. laurae, and considered these names as
synonyms.

Map 63. Distribution of M. laurae.

182
 Ecology and biology. M. laurae is a workerless social parasite. Mei (1987) identified the
host of M. samnitica as M. sabuleti. However, in the non-type series of Mei, collected from
Abruzzo and Lazio, the specimens mounted with them as hosts appeared to be M. scabri-
nodis or perheps, M. spinosior. The type series of M. samnitica was found with its host
colony at altitude 1400 m, living under a stone, Jin a shaded clearing in a Pinus nigra wood.

Myrmica lemasnei Bernard, 1967

(Figs 150 151)

Myrmica lemasnei Bernard, 1967: 123, q, France; Bolton 1988: 4, 1995: 280; Radchenko and Elmes
2003a: 232. !
Sifolinia lemasnei: Kutter 1973: 258, m; Espadaler 1981a: 179, 1981b: 121, q, m.
Symbiomyrma lemasnei: Seifert 1994: 17.

Type localities and type specimens. "Une ouvrière, prise sous une grosse pierre granitique, en avril
1939, dans un nid très peuplé de M. sabuleti, dans un bois de Châtaigniers au col de l'Ouillat. Ce col
est situé à 900 m, près du Perthus et de la frontière espagnole, dans le massif des Albères (Pyrénées-
Orientales... dédiée au myrmécologue G. Le Masne, en compagnie duquel j'ai récolté cette espèce)".
Material examined. 1 m, "Col. de l'Ouillat, leg. Le Masne, Nr. 2", "Sifolinia lemasnei, male Nr. 2",
"Cotypus"; 1 q, "Col de l'Ouillat, leg. Le Masne", "Sifolinia lemasnei, queen Nr. 2"; 1 q, "Cotypus",
"Col de l'Ouillat leg. Le Masne, det. Kutter", "Paratypus", "Sifolinia lemasnei Bernard det. С Baroni
Urbani" (BASLE, GENOA) (see also Notes below); 2 q, N. Spain, Jaca, Huesca, 28.x.1978, leg.
Espadaler (KIEV, ESPADALER, ELMES). J
Distribution (Map 64). France (Pyrénées-Orientales), northern Spain.
Etymology. M. lemasnei: this species is dedicated to the French myrmecologist Prof.
Georges Le Masne.
Notes. Based on the material that we examined, M. lemasnei is clearly a good species
belonging to the karavajevi species group, but its nomenclature is somewhat uncertain.
Bernard described M. lemasnei from a single dealate queen that he said he collected to-
gether with Le Masne in April 1939, from a nest of M. sabuleti living in forest on the Coll
de l'Ouillat (900 m..a.s.l.) in the French Pyrenees near the Spanish border. Quite possibly
the host species was M. spinosior, a species closely related to M. sabuleti (see notes to
M. spinosior below and Seifert 2005). Unfortunately, the unique holotype appears to be
lost; Bernard's collection is stored in the Museum National d'Histoir Naturelles, Paris and
specialists from that museum kindly checked the collection but could not locate the speci-
men. Therefore we have only Bernard's original description of this specimen to go on.
Kutter (1973: 258) also thought that the holotype of M. lemasnei "now seems to be
lost" when he made his review of the social parasites oîMyrmica, although gave no reason
why he believed this. Kutter had new material (8 queens and 3 males, collected by Prof. Le
Masne in July 1956, from the Col de l'Ouillet, the type locality for M. lemasnei) that he be-
lieved to be M. lemasnei. He then transferred M. lemasnei to the genus Sifolinia based on
numerical diagnosis of a combination of "parasitic features" tabulated at the start of his
paper: "darf whol mit Recht die neue Synonymie Myrmica lemasne Bernard = Sifolinea
lemasnei als genügend gesichert anerkannt warden". Then in the following section headed
"New Descriptions" Kutter (1973: 265) made a very clear description of the queens and
males of Sifolinia lemasnei, the description of the queen being treated as a redescription of
Myrmica lemasnei Bernard. We examined a male and a queens from this series and found

183
they totally agreed with Kutter's description, as did 2 queens collected from northern Spain
by X. Espadaler and identified by him as S. lemasnei (Espadaler 1981). Consequently, we
are certain that Kutter's S. lemasnei is a good species that well differs from any other known
Myrmica. These specimens are labelled as "cotypes" but formally they are not types of
M. lemasnei Bernard having been collected much later than the original holotype.
Unfortunately, Bernard's (1967) original description had many errors according to Kut-
ter (1973:258): apparently the holotype was collected by Le Masne in 1950 (not 1939), who
"handed it to Bernard to work on"; the description says nothing as to whether the tibial
spurs are present or have combs; "the size data in the text and under the accompanying
figure do not correspond (3.0 and 3.9 mm respectively)"; "In the text the epinotal spines are
described as blunt, yet in the photograph it is definitely sharply pointed"; "To begin with
a female is described, later a worker".
A further very important discrepancy is the statement by Bernard (1967: 124): "... base
du scape en angle presque droit, légèrement rebordé mais bien moin que selui de sabu-
leti..." ["...antennal scape curved at about right angle, with fine ridge, but it is much smaller
than that in M. sabuleti..."}. This is a key diagnostic feature in genus Myrmica and if cor-
rect M. lemasnei Bernard would belong to a different species group than S. lemasnei sensu
Kutter, which, like M. karavajevi and M. kabylica, has antennal scapes that are gradually
curved at the base and without any trace of a longitudinal ridge or lobe. Unfortunately, the
photograph of the holotype (Bernard 1967, Fig. 127) is of poor quality and does not help
to resolve this question. The pragmatic solution to this problem is dismiss this as another
example of Bernard's sloppy and inadequate description and to accept Kutter's later de-
scription based on Le Masne's new material.
Kutter (1973) infers that the new material taken in 1956 came from host nests close to
the original, if not the same nest, first sampled in 1950 (this date is probably correct because
surely Lemasne himself would have confirmed this with Kutter). It is not unreasonable that
Le Masne could find the original infested nest only six years later because we have noted
the persistence of M. hirsuta in individual M. sabuleti nests and microgynes in M. rubra

Map 64. Distribution of M. lemasnei.

184
nests for at least 4 or 5 years. When it is certain that Bernard's original holotype is lost,
a neotype from the later series could be designated. In the unlikely event that someone
should find the holotype and demonstrate that it is not "M. lemasnei" sensu Kutter, then
Bernard's holotype of M. lemasnei should be Redescribed and Kutter's material would be
a new species.
Ecology and biology. M. lemasnei is a workerless social parasite. In France, this species
was found in pine forest, at altitude 900 m a.s.l., under a granite stone in soil, in the nest of
M. sabuleti; in Spain it was found in oak forest, at altitude 800 m in a M. sabuleti nest
(Espadaler 1981a). Quite probably the host colonies at both sites were M. spinosior, a close
relative of M. sabuleti, on the other hand it is possible that M. lemasnei parasitizes colonies
of both species. |

Myrmica lobicornis Nylander, 1846

(Figs 152-154)

Myrmica lobicornis Nylander, 1846a: 932, w, q, Finland; 1846b: 1052, 1849: 31, m; Foerster 1850a:
68; Nylander 1856: 82; Smith F. 1858: 116; Mayr 1861: 63; Saunders 1880: 216; André 1883: 318;
Nasonov 1889: 35; Forel 1892: 315; Dalla Torre 1893: 111; Ruzsky 1895: 30, 1896: 73; Do-
nisthorpe 1915: 134; Forel 1915: 28; Emery 1916: 120; Bondroit 1918: 105; Müller G. 1923: 39;
Finzi 1926: 106; Karawajew 1926b: 95, 1927a: 283, 1927b: 259; Stärcke 1927: 76; Santschi 1931b:
347; Arnoldi 1934: 166; Karawajew 1934: 87; Ruzsky 1936: 94; Stitz 1939: 98; Weber 1948: 272;
Sadil 1952: 262; Collingwood 1958b: 73; Haustechek 1965: 325; Bernard 1967: 222 (part.); Kut-
ter 1970: 143;'Arnoldi 1970: 1842; Baroni Urbani 1971: 24; Dmitrienko, Petrenko 1976: 15; Kut-
ter 1977: 66; van-Boven 1977: 119; Arnoldi, Dlussky 1978: 534; Collingwood 1979: 51; Espadaler
1981b: 189; Agosti and Collingwood 1987a: 53,Jl987b: 268; Seifert 1988: 38; Atanassov, Dlussky
1992: 101; Radchenko 1994b: 43, 1994c: 141, 1994Í: 87, 1994g: 107; Seifert 1994: 15, 1996: 232;
Radchenko et al. 1997: 485; Czechowski et al. 2002: 22; Boer 2003: 69; Dekoninck et al. 2004: 32;
Radchenko, Czechowska and Czechowski 2004:!46; Seifert 2005: 6, 2007: 208; Radchenko 2007a:
29, nec Eidmann 1941: 14; Kupyanskaya 1985: ¡76; Wei С et al. 2001: 561 (misidentifications).
Myrmica rubra var. lobicornis: Forel 1874: 76; Emery and Forel 1879: 463; Emery 1895a: 315, nec
Wheeler W. M. 1906b: 316 (misidentification). |
Myrmica scabrinodis lobicornis: Mayr 1886b: 451; Ruzsky 1905: 693; Emery 1908a: 179.
Myrmica denticornis Curtis, 1854: 215, w, m, Great Britain (Scotland); Smith F. 1855: 121, q. Syno­
nymy by Mayr 1861: 63; Smith F. 1858: 116; Seifert 1988: 38; Radchenko 2004f: 87; Bolton 1995:
278; confirmed here.
Myrmica lobicornis var. arduennae Bondroit, 1911; 12, w, q, m, Belgium; Emery 1921: 38; Menozzi
1925: 24; Finzi 1926: 108; Weber 1948: 274; Sadil 1952: 263. Synonymy by Donisthorpe 1915: 134;
Bernard 1967: 122; van Boven 1977: 119; Seifert' 1988: 38, Radchenko 1994f: 87; Bolton 1995: 277;
Seifert 2005: 6; confirmed here.
Myrmica arduennae: Bondroit 1918: 105; Kutter 1970: 144.
Myrmica lobicornis subsp. angustifrons Stärcke, 1927: 81, w, q, Great Britain (England); Weber 1948:
274; Kutter 1970: 144. Synonymy by Seifert 1988: 38; Radchenko 1994f: 87; Bolton 1995: 277;
Seifert 2005: 6; confirmed here. I
Myrmica schencki var. starki Karawajew, 1929: 208, w, Russia; Weber 1948: 304, nec Karawajew
1931a:29. Synonymy by Arnoldi 1970: 1842; Seifert 1988: 38; Radchenko 1994f: 87; Bolton 1995:
283; considered as provisional here.
Myrmica schencki var. burtshakabramovitshi Karawajew, 1929: 209, w, Ukraine; Weber 1948: 297.
Synonymy by Arnoldi 1970: 1842; Radchenko 1994f: 87; Bolton 1995: 278; confirmed here.

185
Myrmica lobicornis st. (subsp., nat.) burtshakabramovitshi: Santschi 1931b: 351; Karawajew 1934: 92;
Arnoldi 1934: 167.
Myrmica schencki var. brunescens Karawajcw, 1929: 208, w, Russia; Weber 1948: 297. Synonymy by
Radchenko 19941': 87; considered as provisionai here.
Myrmica lobicornis var. (nat., subsp.) brunescens: Santschi 1931b: 350; Arnoldi 1934: 167, 1970:
1842; Bolton 1995: 278.
Myrmica lobicornis st. foreli Santschi, 1931b: 348, w, q, Austria; Kutter 1970: 144. Synonymy by
Bernard 1967: 122; Seifert 1988: 38; Radchenko 1994f: 87; Bolton 1995: 278; Seifert 2005: 6; con-
firmed here.
Myrmica lobicornis var. kievensis Karawajew, 1934: 91, w, Ukraine; Weber 1948: 286. Synonymy by
Arnoldi 1970: 1842; Radchenko 1994f: 87; Bolton 1995: 280; confirmed here.
Myrmica lobicornis subsp. alpestris Arnoldi, 1934: 168 w, ni "Caucasus" (Armenia); Weber 1948: 274;
Arnoldi 1970: 1842; Kutter 1970: 144. Synonymy by Seifert 1988: 38; Radchenko 19941': 87; Bolton
1995: 277; considered as provisionai here.
Myrmica lobicornis var. lissahorensis Stitz, 1939: 100 (first available use of Myrmica lobicornis subsp.
lobicornis var. lissahorensis Stärcke, 1927, w, Czech Republic); Weber 1948: 287; Sadil 1952: 263;
Kutter 1970: 144. Synonymy by Bernard 1967: 122; Seifert 1988: 38; Radchenko 1994f: 87; Bolton
1995: 280; Seifert 2005: 6; confirmed here.
Myrmica lobulicornis: Bernard 1967: 122; Seifert 1988: 38; Bolton 1995: 280; not confirmed, sec Notes
to M. lobulicornis, below.

Type localities and type specimens. M. lobicornis: "Hab. per totam saltern Fennien sat frequens".
M. denticornis: "I secured three males and four neuters from a nest in Scotland in July 1825; but I did
not observe any females". M. lobicornis var. arduennae: "Hockay, près de Spa. Une petite colonie de
trois nids". M. lobicornis subsp. angustifrons: "Type aus Weybridge (Surrey) 13.VU.'14 leg. H. Do-
nisthorpe, Coll. Van der Weil". M. schencki var. starki: "Chibinische Berge (Halbinsel Kolskij), Strand
des Sees Imandra, 14.VII.1928 (Nr. 4078), V. Stark, ww; - Ebenda, Tundra am. See Imandra,
22.VII.1928 (Nr. 404.6), derselbe, 1 w". M. schencki var. burtshakabramovitshi: "Umgegend des
"Wilden Sees" (Dikoje Osero), Korostenj-Distr., Wolhynien, 28.VII.1927 (Nr. 3788), N. Burtshak-
Abramovitsh, ww. Nest unter Klupen des Oligozän-Sandsteins". M. schencki var. brunescens:
"Teberda, Karatshai-Kreis, Nardabhang der Kaukasus-Bergkette, 20-25.VII.1927 (Nr. 3820),
W. Karawajew, zahlreiche ww aus der Kolonie". M. lobicornis st. foreli: "Tyrol: Schluderbach (Forel).
1 q, 57 w de la collection Forel au Muséum de Genève (communiqués par M. le Dr. J. Carl), 3 w (re-
çues aytrefois de Forel) et d'autres (communiqués par M. H. Kutter)". M. lobicornis var. kievensis:
"Клавд1ево коло Киева (VIII 1932, № 5597, В. Караваев), 2 роб. Ссрсд рослинного детриту, в земл!,
в сосновому nicì" [Klavdievo near Kiev (viii. 1932, No. 5597, W. Karawajew), 2 w. In soil detritus, in
pine forest]. M. lobicornis subsp. alpestris: no precise locality given: "alpestris ist aber eine charakte-
ristische, gut zu unterscheidende Bergform, die im Kaukasus ungefähr von 1500 m. und hinüber auf-
tritt und bis in die großen Hohen von 2200-2400 m. (Armenien) hinaufsteigt". M. lobicornis var.
lissahorensis: "6 ww, Lissa Hora (Besk.) Alfr. Hetschko leg; meine Coll.; Uebergänge Gnojnik (Sil.)".
Material examined. Lectotype of M. lobicornis (designated by Radchenko 2007a; see also Seifert
2005), w (with the male on one pin), "Uleâborg", "W. Nyland.", "Mus. Zool. H: fors Spec. typ. No.
5053 Myrmica lobicornis Nyl." "Lectotype design. Radchenko et Elmes, 2002" (HELSINKI); paralec-
totypes: 1 w, (with the male on pin), "Uleâborg", "W. Nyland.", "Mus. Fenn.", "Mus. Zool. H: fors
Spec. typ. No. 5050 Myrmica lobicornis Nyl."; 1 w (with the male on pin), "Uleâborg", "W. Nyland.",
"Mus. Fenn.", "Mus. Zool. H: fors Spec. typ. No. 5051 Myrmica lobicornis Nyl."; 1 q, "Uleâborg", "W.
Nyland.", "Mus. Fenn.", "Mus. Zool. H: fors Spec. typ. No. 5052 Myrmica lobicornis Nyl."; 1 w (with
the male on pin), Uleâborg", "W. Nyland.", "Mus. Fenn." (HELSINKI); syntypes of M. denticornis: 2
w, 2 m, "denticornis Curt." (MELBOURNE); lectotype of M. lobicornis var. arduennae (designated
here; see also Seifert 2005), w, "Hockay rocaille", "type", "Lectotype design. Radchenko et Elmes,
2000" (BRUSSELS); paralectotypes: 1 w, "Hockai 18-8-10", "Myrmica lobicornis arduennae

186
Bondroit", "type", "Coll. R. I. Sc. N. В."; 1 w, "Coli. R. I. Sc. N. В.", ", "Myrmica lobicornis arduen­
nae Bondr. , type", "R. I. Sc. N. В 21.400"; 5 w (on the same pin), "Hockai", "Coli. R. I. Sc. N. B. Bel­
gique : Prov. De Liége, Hockai", "arduennae Bondr.", "type", "R. I. Sc. N. В 21.400"; 4 w (on the
same pin), "Hockai, rocaille", "Coll. R. I. Sc. N. В. Belgique : Prov. De Liege, Hockai", "Myrmica
arduennae Bondr.", "R. I. Sc. N. В 21.400" (BRUSSELS); 2 w, "Arduennes belges", "M. lobicornis Nyl.
arduennae Bondr. Type"; 2 w, "Hockai Arduennes" (GENOA); lectotype of M. lobicornis subsp. an­
gustifrons (designated here), w, "13.8.14 Weybridgè Surrey, leg. Donisthorpe", "angustifrons Stärcke
type", "Holotype w ssp. angustifrons Stärcke" "Lectotype design. Radchenko et Elmes, 2000" (LEI-
DEN); syntypes of M. schencki var. starki, 4 w, ¡"Хибины, бср. оз. Имандра, 14.VII.28, В. Старк"
[Khibiny, shore of lake Imandra, 14.vii.28, leg. V. Stark], "4078 Coll. Karawajewi"; "M. (Myrmica)
schencki v. starki Karaw. typ.", "Syntypus Myrmica schencki var. starki Kar." (KIEV); syntypes of
M. schencki var. burtshakabramovitshi, 4 w, "Коростсн. у., Бурчак-Абрамович" [Korosten' uezd, leg.
Burtschak-Abramovich], "3788 Coli. Karawajewi", ¡'Myrmica schencki Em. v. burtshak-abramovitshi
п.", "Syntypus Myrmica schencki var. burtshak-abramovitshi Karaw." (KIEV); lectotype of
M. schencki var. brunescens (designated here), w [(upper on the pin with 3 w), "Тсбсрда, Караваев"
[Teberda, leg. Karawajew], "3820 Coli. Karawajewi", "Myrmica lobicornis v. brunescens Karaw.
cotyp", "Syntypus Myrmica lobicornis brunescens Karaw."; paralectotypes: 2 w on the pin with lecto­
type; 11 w, "Тсбсрда, Карачай. ок., 20-29. VIII. 1927, В. Караваев" [Teberda, Karachaisky Okrug, 20-
29.viii.1927, W. Karawajew], "3820 Coll. Karawajewi", "Myrmica schencki Em. v. brunescens Karaw.
typus", "Syntypus Myrmica lobicornis brunescens Karaw." (KIEV, MOSCOW); lectotype of
M. lobicornis st. foreli (designated here; see also Seifert 2005), w, "type", "Myrmica foreli Sant. Type",
"Schluderbach, Forel", "5", "lobicornis var. foreli Finzi i. t.", "Sammlung Dr. F. Santschi Kairouan",
"Lectotype design. Radchenko et Elmes, 2000" (BASLE); paralectotypes: 2 w, "type", "Myrmica foreli
Sant. Type", "Suisse Schluderbach - Forel", "M. lobicornis Nyl. w, Schluderbach", "Sammlung Dr. F.
Santschi Kairouan"; 3 w, "cotype", "Myrmica foreli Sant.", "5" (BASLE); syntypes of M. lobicornis var.
kievensis, 2 v/, "Клавдисво, Караваев" [Klavdievo, leg1. Karawajew], "5597 Coli. Karawajewi", "M. [Myr­
mica) lobicornis var. kievensis Kar. typus", "Syntypus Myrmica lobicornis var. kievensis Kar." (KIEV);
lectotype of M. lobicornis subsp. alpestris (designated here), w (upper on the pin with 4 w), "Арчанск,
2300 M, Севан, Армсн., Арнольди, 25.VII.27" [Archansk, 2300 m, Sevan, Armenia, leg. Arnoldi,
25.vii.27], "2821" (MOSCOW); paralectotypes: 3 w on the pin with lectotype; 10 w, 5 m, "Armenia,
Sevan, VII.27, K. Arnoldi", No. 2820, 2821, 2804, 3061 (MOSCOW); 2 w, "Севан, Елсновка, 2300 м"
[Sevan, Elenovka, 2300 m], "5461 Coll. Karawajewi", "Myrmica lobicornis Nyl. ssp. alpestris Arnoldi
typ.", "Paratypus Myrmica lobicornis alpestris Arnoldi"; 2 w, 1 m, "5624 Coli. Karawajewi", "2820,
Армения, Севан, Арчаноц, 2400 м, VII.27, К. Арнольди" [Armenia, Sevan, Archanots, 2400 m, vii.27,
leg. К. Arnoldi], "Myrmica lobicornis Nyl. ssp. alpestris Arnoldi typ.", "Paratypus Myrmica lobicornis
alpestris Arnoldi" (MOSCOW, KIEV); lectotype of M. lobicornis subsp. lissahorensis (designated
here), w, "Lissa Hora (Besk.) Alfr. Hetschko", "type", "Holotype v. lissahorensis Stärcke" (LEIDEN);
non-type material: > 300 w, > 50 q, > 50 m from whole range.
Distribution (Map 65). Boreo-montain species. North Europe, northern part of Central
Europe, British Isles, mountains of Central and Southern Europe (in Iberian Peninsula is
absent), Forest Zone of East Europe, Caucasus, West Siberia and NE Kazakhstan (to the
south till Saur Range), East Siberia (to the east till Transbaikalia), Mongolia. Records for
Russian Far East and China (Eidmann 1941; Kupyanskaya 1985; Wei С. et al. 2001) al­
most certainly belong to other species.
Etymology. M. lobicornis: combination of Greek loßoc (lobos) = lobe [of ear] and Latin
cornis (adj) = horned or antlered, to describe the vertical lobe or projection on the bend of
the scape. M. denticornis: combination of Latin dent = tooth and comis (adj) = horned or
antlered, to describe the tooth-like projection on the bend of the scape. M. arduennae:
the name Arduenna is derived from the Gaulish for a mountain region and gave rise to the
name for the Ardennes region of Belgium (e.g. Arduenna silva = Ardennes woods) and its

187
associated goddess Arduinna. M. angustifrons: name a combination of Latin words angus-
tus = constricted or narrow and front = forehead or frons of insects, to indicate a narrower
frons than "normal" for M. lobicornis. M. starki: named for the collector, the Russian
forestry entomologist and coleopterologist, Prof. Vladimir Nikolaevich Stark (1898-
1962) of All-Union Institute of Plant Protection, Pushkin (near St. Petersburg).
M. burtshakabramovitshi : named for the collector, the Ukrainian zoologist, Prof. Nikolai
Iosifovich Burchak-Abramovich (1902-1997) of the Zoological Museum of the Ukrainian
Ac. Sci., Kiev. M. brunescens: name derived from the Germanic word brun = brown and the
Latin suffix escens = -ish, to indicate its brownish colour. M. foreli: species dedicated to Au-
guste Forel (see biography in Chapter 1.6). M. kievensis: named from Kiev with the Latin
suffix ensis = place of origin, to indicate that the type locality is the Kiev region of Ukraine.
M. alpestris: from the name Alp with the Latin suffix estris = habitat or origin, to indicate
that it is an alpine species. M. lissahorensis: from the name Lissa Hora (Bald Mountain)
with the Latin suffix ensis = place of origin, the Lissa Hora is part of Beskid mountain
range that extends into southern Poland, where the types were taken.
Notes. Material of the nomen nudum Myrmica nodicornis Chappell, 1886 was referred
to M. lobicornis by Donisthorpe (1915).
M. lobicornis belong to the lobicornis species group. In Europe the main cha-
racters used to separate its species from the schencki-group species are the shape and size
of the lobes on the bend of the antennal scape, combined with the relative frons width (FI).
However, in Europe these features can be very variable between local populations and can
overlap those of schencki-group species, as a result authors described a large number of in-
fraspecific forms. A number belong to the closely related M. lobulicornis (see notes to that
species and Seifert 2005), which might be considered as a "southern European mountain
species". Also, there are several local endemics in the southern mountains of "Greater
Europe". All the remaining forms are included under the name M. lobicornis. In the Alps
of Central Europe one can find quite different forms that one can attribute to the various
synonymised names, living in adjacent valleys and even several forms within the same

Map 65. Distribution of M. lobicornis.

188
population. Kutter (1970) could not discriminate between the males of these forms and
proposed a semi-numerical system of describing the existing (and new) forms of M. lobicor­
nis without resorting to further names. He pointed out that Nylander's types seemed well
removed from the centre of variation of M. lobicornis generally but concluded that we could
not get much further without genetical studies (this in times pre-dating modern molecular
analysis). At the time of writing such genetical studies are possible and initial results on
M. scabrinodis, a similarly morphological and ecological variable species in Europe (see
notes to that species), suggest that it might comprise several morphologically cryptic but
genetically "good" species. We suspect thatMJ lobicornis will be shown eventually to com­
prise several cryptic species. The most probable explanation is for a post-glacial invasion of
morphologically distinct forms from different glacial refugia with mixing and hybridisation
in some regions, combined with strong selection for speciation resulting from man's
post-glacial impact creating large areas of novel habitat. This is explored a little more for
M. lobicornis (below).
We can divide the other infraspecific forms of M. lobicornis according to whether they
were collected from Europe proper or from the Caucasus region. We examined and re­
examined the type specimens of M. denticornis, var. arduennae, subsp. angustifrons, var.
burtshakabramovitshi, st. foreli, var. kievensis and var. lissahorensis, all of which were de­
scribed from plains or low mountains of Central and East Europe (see above).-All these
forms have narrow frons (FI < 0.30, i.e. typical for M. lobicornis "s. str.") and by the addi­
tional important diagnostic features, such as the shape of petiole, length of propodeal spines,
etc., all are contained in the range of variability of the "typical" M. lobicornis. On the other
hand, the Caucasian var. brunescens and subsp. alpestris have relatively wide frons (mean
FI 0.304 and 0.323 respectively) and in this respect they are similar to M. lobulicornis. The
exception to this rule is var. starki, which was originally described as form of M. schencki
from Cola Peninsula near Polar Circle. It has wide frons (FI 0.333-0.338), even wider than
in the Caucasian forms.
In a preliminary study we measured the range of frons width from several hundreds of
specimens from across the European range of M. lobicornis. We found that if one draws an
imaginary line from Finland to the north coast of Spain all areas to the north-west have
only specimens that closely conform to Nylander's type with И about 0.28 and with fairly
large distinct vertical lobes or plates on the antenna. This area includes the Lucitanian re­
gion of Northern Spain, Western France, the whole of Great Britain and Ireland as well as
Holland, Belgium, north Germany, west Finland and the rest of Scandinavia. To the south
and east of this line populations are much more variable with respect to FI and size of lobe.
Furthermore, we measured several tens of M. lobicornis workers from eastern populations
(West Siberia) and in all respects they were typical for the species, except for an extremely
variable width of the frons: FI ranged from 0.27 to 0.32 and a wide variation occurred even
within the nest samples. We note also that most species of southern regions tend to have
relatively wider frons and smaller lobes compared to their more northern counterparts (e.g.
M. lobicornis and M. lobulicornis; M. schencki and M. obscura - see Elmes et al. 2009).
It appears that in the north, the wider frons form prevails in more continental climates
and the narrow frons form in more oceanic climates, so perhaps the climate of southern
mountains more resembles continental climates. Unfortunately we have no firm data to un­
derstand the functional purpose of varying frons-width and lobe sizes but as indicated else­
where in this monograph, we have anecdotal evidence that it is probably related to
individual foraging behaviour. |

189
 Provisionally we suggest the following zoogeographical explanation for the current
situation. During the previous interglacial period M. lobicornis living in the northern low-
lands developed a cline from east to west with the narrower frons forms living in the west.
Other more locally distributed wider-frons forms lived in southern mountains (e.g. M. Lo-
bulicornis and various local endemics). During the ice period these populations retreated
towards the south finding refugia in Iberia, Italy, Caucasus etc., the southern mountain
forms retreating even further south in these refugia. During this time there undoubtedly
was some mixing between related species with different "glacial lineages" being established
in the various refugia. At the end of the glaciation the "northern" adapted species spread
northwards and eastwards from the refugia while the more "southern" adapted species re-
colonised the mountains in the north of the various refugia.
In the case of M. lobicornis, we suggest that a narrow-frons form (M. lobicornis "s. str.")
with a Lucitanian distribution expanded into Scandinavia and Northern Europe where it en-
countered one or more wide-frons varieties invading from more eastern refugia. This would
lead to the current distribution of "pure" narrow-frons forms (described above) and the
persistence of a pure race of wide-frons forms that invaded from the east, in the extreme
north. Everywhere else there has been considerable mixing and probably hybridisation be-
tween the forms combined with various levels of local selection, driven by anthropogenic
factors, for either wide-frons and small lobed forms or narrow-frons and large-lobed forms,
which has resulted in the variability in Switzerland described by Kutter (1970) and in
Western Siberia observed by us.
Like Kutter (1970) we must conclude that study of morphology and morphometries can
throw little more light on this problem, which can only be resolved by genetical studies.
Fortunately, suitable techniques are now available and we can hope that somebody will feel
inspired to apply molecular methods to material collected from the whole range of
M. lobicornis "s. 1.".
Ecology. Despite being an "old" and well-known species in Europe, the ecology of M. lo-
bicornis has not been well studied. It has been used as a model species in chemical studies
of glandular secretions in a team led by Prof. David Morgan (see Chapter 1.3.2), its role
as a host for Phengaris butterflies has been investigated recently (see papers cited in Chap-
ter 1.3.6), and Prof. Vladilen Kipyatkov has made some observations on physiological
response to seasonality (see references in Chapter 1.3.5). The most probable reason for
this is that over most of its entire distribution this species is common in the sense that one
can occasionally find the odd colony living in more or less any biotope, but unlike most
other widely distributed Myrmica species it almost never dominates any particular habitat-
type anywhere in its range and in this sense can be considered rare. It is more common in
boreal habitats and we found it to be relatively abundant in some pine forests of northern
Finland. In most northern habitats, the narrow-frons form (M. lobicornis "s. str.") tends to
be a quite cryptic-foraging species; in subarctic conditions we have observed quite small
individuals foraging on the soil surface under the moss layer that might initially be
mistaken for Leptothorax species. Even in southern England it is quite hard to observe
foraging workers.
Generally colonies are small (< 200 workers) with at most 2 queens, however in Cen-
tral Lapland, Finland we found one very large colony (probably > 2,000 workers) that had
made a large earth solarium in grass on the bank of a small stream in a forest, but we could
find no queens in this nest. Its greater abundance in Finland led to the only genetical study
of M. lobicornis (Seppä 1994), who showed that on one study site in Finland most colonies

190
averaged about 300 workers but rarely exceed 500 workers and were monogynous (or
queenless), on average queens live about two years and regular replacement of queens by
queenless colonies was inferred.
In Finland M. lobicornis is not usually found in dense old forest, M. ruginodis is usually
the only Myrmica species found in such (> 100 years) forest (Punitila et al. 1991). M. lo-
bicornis and the other two common forest Myrmica species, M. scabrinodis and M. sulci-
nodis, are found in regenerating forest but none of these can be considered as primary
colonising species like M. ruginodis, nests of which persist after clear felling and rapidly
expand into the new habitat. Nests of M. lobicornis and the other two species must be
established by queens flying into the felled areas. Punttila et al. (1991) showed that this
occured sometime after two years and that by 10 years after forest clearance the population
of M. lobicornis was well established, though individual workers were much less abundant
than those of the other three species. Numbers decline as the new forest closes over but it
is not known how long M. lobicornis colonies persist in mature forest in Finland. In the
much warmer ancient mixed (mostly pine and some deciduous) woodland near Kiev,
Ukraine we have observed colonies of up to seven Myrmica species, including M. lobicornis.
However the dominant, species are M. rubra and M. ruginodis and the other Myrmica
species are relatively rare.
In the glades of the Tatry mountains of southern Poland M. lobicornis were present at
only 7 of 23 studied sites compared with 20 and 13 sites for M. ruginodis and M. scabri-
nodis respectively; when present, nests of M. lobicornis averaged 13% of the population of
all ant nests compared to 52% and 19% respectively for the other species (Woyciechowski,
1990c). On the meadows on the southern slopes of a mountain in the Beskid range of
southern Poland, M. lobicornis nests comprised only 6% of the population of ant nests de-
spite being the third most abundant species after M. scabrinodis (30%) and Lasius flavus
F. (57%) (Woyciechowski and Miszta 1976). Nuptial flights have been observed from the
end of July to mid September, and M. lobicornis males often join swarms dominated by
other Myrmica species (see Chapter 1.3.5).

Myrmica lobulicornis N y l a n d e r , 1857

(Figs 155-157)

Myrmica lobicornis var. lobulicornis Nylander, 1857: 79, w, France; Dalla Torre 1893: 112; Bondroit
1918: 105; Emery 1921: 38; Weber 1948: 287; Radchenko 2007a: 29.
Myrmica lobulicornis: Bondroit 1920a: 151.
Junior synonym of M. lobicornis: Bernard 1967: 122; Seifert 1988: 38; Bolton 1995: 280. Revived from
synonymy and raised to species: Seifert 2005: 6; 2007:210.
Myrmica arduennae var. pyrenaea Bondroit, 1918: 106, w, France. Synonymy by Seifert 1988: 38;
Bolton 1995: 282 (as synonym of M. lobicornis)) Seifert 2005: 6 (as synonym of M. lobulicornis);
the latter confirmed here.
Myrmica pyrenaea: Bondroit 1920a: 151.
Myrmica lobicornis var. (st., subsp.) pyrenaea: Finzi 1926: 108; Santschi 1931b: 350; Weber 1948:
287.
Myrmica lobicornis subsp. alpina Stärcke, 1927: 801, w, q, Italy; Weber 1948: 274; Kutter 1970: 144.
Synonymy by Bernard 1967: 122; Seifert 1988:|з8; Radchenko 1994f: 87; Bolton 1995: 277 (as
synonym of M. lobicornis); by Seifert 2005: 6; Radchenko 2007a: 39 (as synonym of M. lobuli­
cornis); the latter confirmed here.

191
Myrmica lobicornis subsp. apennina Stärcke, 1927: 82, w, Italy; Weber 1948: 274; Kutter 1970: 144.
Synonymy by Seifert 1988: 38; Radchenko 1994f: 87; Bolton 1995: 277 (as synonym of
M. lobicornis); by Seifert 2005: 6 (as synonym of M. lobulicornis); the latter confirmed here.

Type localities and type specimens. M. lobulicornis: "Mont-Dore". M. arduennae var. pyrenaea:
"Pyrénées-Orientales: Canigou, vers 1800 m". M. lobicornis subsp. alpina: "Typen aus Val Aosta, La
Thuile leg. 1915. Meine Coll." M. lobicornis subsp. apennina: "Typen vom Monte Cimone (Apenninen
zwischen Bologna und Lucca) mir von Prof. С. Menozzi geschenkt".
Material examined. Lectotype of M. lobulicornis (designated by Radchenko 2007a; see also Seifert
2005), w, "M. Dore", "F. [Siel] lobicornis var. lobulicornis Nyl." "W. Nyland.", "Mus. Zool. H: fors
Spec. typ. No. 5118 Mymíca lobulicornis Nyl." (HELSINKI); paralectotype, w, "M. Dore", "W. Ny-
land.", "Mus. Zool. H: fors Spec. typ. No. 5119 Myrmica lobulicornis Nyl." (HELSINKI); lectotype of
M. arduennae var. pyrenaea (designated on material by Radchenko and Elmes, 2000; see also Seifert
2005), w, "Canigou 10-1917", Coli. R. I. Sc. N. B. France, Pyrén. Orient. Canigou (Mont) -X-1917",
"Myrmica pyrenaea Bondr. type", "Lectotype design. Radchenko et Elmes, 2000" (BRUSSELS);
paralectotypes: 64 w with the same labels (BRUSSELS); 1 w, "Canigou", "ex. Coll. Donisthorpe
В.M. 1934-4", "Myrmica v. pyrenaea Bondr. type", "Syntype" (LONDON); 2 w, "Myrmica var. pyre­
naea Bondroit, type, Canigou" (GENOA); lectotype of M. lobicornis subsp. alpina (designated here),
w, "Val Aosta 1915 La Thuile", "holotype ssp. alpina Stärcke", "Lectotype design. Radchenko et
Elmes, 2002" (LEIDEN); paralectotype: 1 w, "Val Aosta 1915 La Thuile", "holotype ssp. alpina
Stärcke", "holotype subspecies alpina Stärcke", "Mus. Zool. H: fors Spec. typ. No. 5322 M. lob. alpina
Stärcke" (HELSINKI); lectotype of M. lobicornis subsp. apennina (designated here), w (upper speci-
men on the pin with 5 w), "Emilia M. Cimone 10.VIII.1921 С Menozzi", "M. lobicornis st. arduennae
Bondroit Menozzi deter.", "M. lobicornis ssp. apennina typen det. Stärcke", "Holotype ssp. apennina
Stärcke", "Lectotype M. lobicornis apennina design. Radchenko et Elmes, 2000" (LEIDEN); paralec-
totypes: 4 w on the pin with lectotype (LEIDEN); non-type material: about 70 w, 3 q and 3 m from
Iberian Peninsula, Tenerife Isl. and Italy.
Distribution (Map 66). Iberian Peninsula, Canary Islands, southern France, Italy,
Switzerland, Austria.

Map 66. Distribution of M. lobulicornis.

192
 Etymology. M. lobulicornis: combination oí Greek Xoßoc {lobos) = lobe [of ear] with
diminutive suffix ulus and Latin cornis (adj) = horned or antlered, to describe the rela-
tively small vertical lobe or projection on the bend of the scape. M. pyrenaea: name (adj)
means from the Pyrénées, the mountains between France and Spain that is the type
locality for this form; the name is derived from the Greek IJvprjvaioç = Pyranaeus, a lover
of Hercules who is buried in the mountains. M. alpina: name means of Alps or mountains;
in this case specifically the European Alps where the types were found, derived from the
Latin alpes = high mountain, often white [show] implied. M. apennina: named for the
Apennine mountain range that forms the central "backbone" of Italy, the type locality is
Monte Cimone, a mountain in the Apennines of Emilia-Romagna.
Notes. M. lobulicornis belongs to the lobicornis species group and strongly resembles
M. lobicornis, differing mainly by its wider frons (mean FI 0.34 vs 0.28), its generally smaller
scape-lobes and more southerly, montane distribution in Europe (see notes to M. lobicornis
above). In Europe, together with M. lobicornis, it comprises a complex of closely related
species and infraspecific forms; Kutter (1970) divided this complex into two groups: those
with a narrow frons and those with relatively wider frons. Examination of Nylander's types
confirms that the type of M. lobicornis is typical of the narrow-frons form of north-west Eu-
rope, and the type of M. lobulicornis is typical of the wide-frons form common
in southern Europe. Nylander's type was taken at Mont Dore, in the Monts d'Auvergne in
Le Puy-de-Dôme Dept. of Central France, which is towards the northern limit of distribu-
tion of the wide-frons forms (see notes to M. lobicornis above). Recently Seifert (2005)
raised M. lobulicornis to species level and considered all the above wide-frons forms
from the mountains of southern Europe as its junior synonyms. Based on our examination
of the type specimens and non-type material from mountains in South Europe, we confirm
this synonymy. All old records of M. lobicornis ¡from the Iberian Peninsula belong to M. lo-
bulicornis (Xavier Espadaler, pers, comm.) or related endemie species (see notes to
M. xavieri).
We found in the Humboldt Museum of Zoology, BERLIN specimens that conform to
M. lobulicornis, labelled "Saint Cruz, Tenerife] Hermon" . If these labels are correct then
almost certainly this species has been accidentally introduced to the Canary Islands some
time during the last few centuries.
It seems to us that the current situation with M. lobicornis and M. lobulicornis is now
somewhat similar to the state of affairs a few years ago in Europe concerning the black ants
of genus Lasius. Two related species were recognised, L. niger and L. alienus Foerster that
could be clearly separated by morphology, ecology and to some extent distribution in most
cases. However, both species occupied a very yide range of habitat types and often it was
difficult to assign some populations to a species. Then Seifert (1992) demonstrated that L.
niger comprised two species with distinct ecologies, and L. alienus comprised three species.
We suspect that when similar detailed ecologically (and genetically) based studies are made
on M. lobicornis and M. lobulicornis some of the infraspecific forms, particularly those with
wide-frons, might be shown to be good species. The queens and males have not been de-
scribed previously, therfore we give formal descriptions.
First description of queens (Fig. 156).
Relatively small (HW< 1.10, AL < 1.90 mm). Generally, like the worker by all main di-
agnostic features. Typically the scape bend has a small dentiform lobe and petiolar node is
not strongly angled (seen in profile) ; the frons is somewhat relatively wider than that of the
workers.

193
 First description of males (Fig. 157).
Head slightly longer than broad, with weakly convex sides and more convex occipital
margin, and moderately rounded occipital corners; anterior clypeal margin slightly convex
and distinctly notched medially. Antennal scape quite long (SIj > 0.75, Sl 2 > 0.80), strongly
curved at the base (similar to that of M. lobicornis), antennae 13-segmented, with 5-seg-
mented club; second funicular segment approximately 1.3 times longer then third one.
Alitrunk relatively long, scutum slightly convex, scutellum does not project dorsally above
scutum when seen in profile. Propodeum with blunt, thick subtriangular teeth. Petiole with
short peduncle, although distinctly longer than height, its anterior surface straight and node
dorsum widely rounded; postpetiole higher than long, with weakly convex dorsum.
Head dorsum densely though not coarsely punctated, central part of clypeus almost
smooth, appearing shiny. Frons with longitudinal rugae, temples with fine reticulation.
Scutum mostly smooth and shiny, only its central part posteriorly to Mayrian furrows
with a few quite coarse longitudinal rugae, scutellum and propodeal dorsum coarsely lon-
gitudinally rugose. Sides of alitrunk longitudinally rugulose or rugose. Petiolar node very
finely punctated, but appears shiny, postpetiole smooth and shiny.
Head margins and mandibles with numerous, relatively long, slightly curved erect hairs.
Alitrunk with somewhat sparser hairs, waist and gaster with abundant standing hairs. Scape
with abundant erect to suberect hairs, the longest hairs sub-equal to the maximum scape
width. Tibiae and tarsi with relatively short subdecumbent hairs. Body colour dark brown;
appendages somewhat lighter.
Ecology. Like the taxonomy, the ecology of M. lobicornis and M. lobulicornis has been
confounded and little can be added to the information given for M. lobicornis (above). In
our experience, M. lobulicornis lives in more open habitats than M. lobicornis, often being
found under stones and having generally larger colonies. It is a slightly less cryptic forager
than M. lobicornis (see notes on ecology of that species, above) and forages among the
stems of the vegetation more similar to the mode of foraging of sympatric M. scabrinodis.

Myrmica lonae Finzi, 1926

(Figs 158-160)

Myrmica scabrinodis subsp. lonae Finzi, 1926: 103, w, q, m, Finland, Austria and Italy; Karawajew
1929: 207.
Myrmica sabuleti st. (var., subsp.) lonae: Santschi 1931b: 346; Stitz 1939: 96; Weber 1948: 294; Sadil
1952: 249.
Junior synonym of M. sabuleti: Bernard 1967: 117; Arnoldi 1970: 1841; Seifert 1988: 31; Atanassov,
Dlussky 1992: 97; Radchenko 1994d: 80; Bolton 1995: 281; not confirmed, see Notes below.
Myrmica lonae: Seifert 1994: 12; 1996: 226; Czechowski et al. 1997: 490; Radchenko et al. 1997: 490;
Seifert 2000: 195; Czechowski et al. 2002: 30; Radchenko and Elmes 2004: 232; Radchenko,
Czechowska and Czechowski 2004: 50; Seifert 2007: 199, q.
Myrmica rubra var. scabrinodolobicornis: Sadil 1952: 249; not confirmed, see Notes to M. sabuleti,
below.
Myrmica sabuleti var. salina: Sadil 1952: 249; not confirmed, see Notes to M. salina, below.

Type localities and type specimens. "Ho esaminato: w da Karislojo (Finlandia - leg. dott. Forsius) ;
w, q, m da Ratzes (Tirólo - dot. Wolf - coli. Müller); w, q, M. Baldo (Trentino - leg. Lona); w -
M. Nanos (Ven. Giulia); w, M. Disgrazia (Frasca - coli. Museo civ. di Genova)".

194
 Material examined. Syntypes, 1 w, "M. Nanos, V.23 Finzi", "4061 Coll. Karawajewi", "M. (Myr-
mica) scabrinodis Nyl. v. lonae Finzi det.", "Syntypus Myrmica scabrinodis v. lonae Finzi" (KIEV);
1 w, "M. Disgrazia, 2500 m, Frasca 85", "M. scabrinodis sub. Lonae Finzi i. L. 1924" (GENOA);
non-type material: > 150 w, > 20 q, > 30 m from Europe, West Siberia and Aitai Mts.
Distribution (Map 67). Central and North Europe, Balkans (from NW Italy to Bulgaria),
East Europe, Asia Minor, Caucasus, West Siberia, northern Kazakhstan, Altai Mts.; all pre-
vious records of M. sabuleti east of Ural Mts. belong to M. lonae (see Radchenko 1994d).
Etymology. M. lonae: named for Carlo Lonâ (1885-1971) of Trieste, a coleopterologist
and schoolteacher who participated in the 1923 Ravasini-Lona expedition to Albania; they
made their ant collection available to Finzi.
Notes. M. lonae belongs to the sabuleti-complex of the scabrinodis species group (see
also Radchenko and Elmes 2004). It is very similar to M. sabuleti and differs from it mainly
by the extremely well-developed, large lobe at the base of scape, which is raised above the
level of the dorsal plane of scape (see Key, Chapter 5.1, 5.2 and Figs 158 and 230), and by
a somewhat narrower frons. However, the males of these two species are practically indis-
tinguishable.
Since its description, M. lonae was considered by most authors to be at best
a sub-species, and later a junior synonym of MJ sabuleti (see above). If one compares just
the shape and size of the lobe of M. sabuleti iti is found to be quite variable; we have ob-
served a sort of cline in Europe from the northeast (large lobes very similar to M. lonae) to
the southwest (very small lobes that can be confused with M. scabrinodis). However, Seifert
(1994) showed that if one measures the extent to which the lobe is raised over the dorsal
plane of the scape there is a clear difference from M. sabuleti, which he considered sufficient
to make M. lonae a good species; consequently he revived the name M. lonae from
synonymy (see also Seifert 2000).
While we agree that this feature clearly separates the two forms there is no disjunction
in their distribution (as, for example, is the case between M. schencki and M. obscura).

Map 67. Distribution of M. lonae.

195
We have frequently observed M. lonae and M. sabuleti living sympatrically and on several
occasions have found totally mixed mating swarms of the two species, as males can not be
distinguished it was impossible to say whether there is like-with-like matings. We think it
is quite possible that M. lonae is an "ecological race" adapted to cooler habitats, with mor-
phological differences only expressed in the female castes (see ecological notes below). This
question can only be resolved by detailed ecological study supported by molecular geneti-
ca! analyses of sympatric populations. Until such studies are made it is best to accept
M. lonae as a good species.
Sadil (1952) synonymised M. rubra var. scabrinodolobicornis and M. salina with
M. lonae but gave no comments as to why. We do not accept this treatment; we consider
that M. salina is a good species but var. scabrinodolobicornis is a junior synonym of M. sa-
buleti (see notes for the corresponding species, and also Radchenko and Elmes 2009b).
Ecology Generally, M. lonae is a much less thermophilous species than M. sabuleti and
inhabits more shady, relatively cooler and humid microhabitats. For example, we found
a population living on "humid heath" in the east of The Netherlands (Elmes et al. 1994)
where M. lonae comprised about 15% of the population of Myrmica nests (M. ruginodis
comprised about 50% and M. scabrinodis 35%). The vegetation was typical for humid heath
and very little difference could be detected in the microhabitats used by these three species.
M. sabuleti very rarely co-exists with M. ruginodis in west Europe so the population of
M. lonae was recognised as being unusual even before noting their huge scape lobes; M. sa-
buleti were not present on the site but nests could be found in warmer, drier areas of grass-
land in the same locality. On the other hand, M. sabuleti quite often coexists with
M. scabrinodis on grassland sites (see notes to ecology of M. sabuleti). Four nests were
excavated and ranged in population from 1 queen with 19 workers to 2 queens with almost
1600 workers.
In Finland, M. lonae is common in open pine and mixed forests where it builds nests in
the moss and underlying soil. A common feature of these sites is that they are rather cold
and wet in early summer but become quite hot and dry in mid and late summer. In more
southerly mountains (e.g. Transcarpathia, Ukraine) we have observed M. lonae living in
meadows at the forest edge and in higher mountain meadows up to 1200 m a.s.l. In these
habitats nests are built in soil, often under stones, but also in decayed wood. M. sabuleti live
sympatrically at these sites and appear to prefer the more open better grazed vegetation, but
we have not made a detailed study of this. Nuptial flight occurs in August-September and
swarms often contain both M. lonae and M. sabuleti gynes.
We indicated that M. lonae might be a morph produced in response to ecological con-
ditions. Our evidence for this comes from unpublished observations on a site managed to
produce a large population of M. sabuleti as a management strategy for the conservation of
a Large Blue Butterfly in the UK (see Thomas et al. 1998, Fig 2). Before management of this
site in southwest England, the M. sabuleti population lived at the hotter end of a natural
temperature gradient created on a south-facing slope. A few M. schencki colonies lived in
the hottest places while M. scabrinodis dominated the central open grassy parts, and M. ru-
ginodis were found in the coolest conditions among gorse scrub (Ulex spp.) at the bottom
of the slope. Management consisted of a mixture of burning and grazing to generally in-
crease soil temperatures throughout the site and specifically to maximise the hot short-
sward conditions favoured by M. sabuleti. After some years the distribution of Myrmica
species reflected the desired outcome with M. sabuleti dominating the grassy middle part
of the site, M. scabrinodis lived in the cooler conditions lower on the slope (of course there

196
was an area of overlap between these two species) and the M ruginodis population had de­
creased and nests were more or less restricted to the edges of the remaining clumps of gorse
at the bottom of the slope. However, in the long grass in the glades between the gorse
M. sabuleti were apparently living, in considerably more shaded cooler conditions than
M. scabrinodis. These ants were generally somewhat larger, darker with a somewhat dif­
fering foraging behaviour than the M. sabuleti. By ecological differences, we would call
these ants M. lonae but we were unable to make any separation from M. sabuleti using the
character proposed by Seifert (see notes above); however, the scape lobes were much larger
than in the typical M. sabuleti found on the site.1 We suggest that M. sabuleti might produce
a female morph to exploit habitat that is quite cool and moist in spring but becomes quite
hot and dry in summer. Based on other studies one would expect such a morph to be
"physiologically more active" (see Chapter 1.3.5) and this might in some way be linked to
a more developed scape lobe. It would not be unusual if such characters were only expressed
in the female caste and males from either lineage might mate freely with any type of female.
However, this hypothesis would need testing by a series of careful breeding tests combined
with physiological and molecular genetical studies.

Myrmica luteola Kupyanskaya, 1990

(Figs 161-163)

Myrmica luteola Kupyanskaya, 1990: 103, w, q, Russia; Radchenko 1994b: 44,1994c: 136, 1994e: 75,
m; Bolton 1995: 281; Masuko and Terayama 2002: 224; Imai et al. 2003: 182; Radchenko and
Elmes 2003a: 239; Radchenko 2005: 143.

Type localities and type specimens. "Материал. Голотип - рабочий, Курильские острова, о-в Кунашир,
оз. Горячее, 24.VIII 1975 (Купянская); паратипы: 21 рабочий из той же серии; Приморский край, Кедровая
Падь, 18.IX 1973 (Купянская), 75 рабочих, 1 Ç" [Material. Holotype - worker, Kurily Islands, Isl.
Kunashir, Lake Goriachee, 24.viii.1975, leg. Kupyanskaya; paratypes: 21 workers from the same nest
series; Primorsky Krai, Kedrovaya Pad', 18.ix.1973, leg. Kupyanskaya, 75 workers, 1 queen].
Material examined. Holotype, w, "о-в Кунашир, оз. Горячее, 24.08.1975, Купянская" [Isl. Kunashir,
Lake Goriachee, 24.viii.1975, leg. Kupyanskaya] (MOSCOW); paratypes: 4w"o. Кунашир, оз. Горячее,
из почвы, 25.VIII.75, Купянская" " [Isl. Kunashir, Lake Goriachee, 24.viii.1975, from soil, leg. Kupyan­
skaya]; 11 w, 1 q, "Приморск. край, Кедровая Падь, 18.IX 1973, Купянская" [Primorsky Krai, Kedrovaya
Pad', 18.ix.1973, leg. Kupyanskaya] (MOSCOW, VLADIVOSTOK); non-type material: about 50 w,
9 q, 6 m from Russian Far East, Korean Peninsula, China and Japan.
Distribution (Map 68). Southern part of the Russian Far East, NE China, Korean Penin­
sula and Japan.
Etymology. M. luteola: named from the Latin diminutive adjective luteola = yellowish;
for the colour of the workers.
Notes. M. luteola is quite unusual compared to most other Myrmica species and well dif­
fers from any other known member of the genus; this led Radchenko (1994a) to define the
separate, luteola species group. This idea was recently supported by the molecular-
genetic phylogeny produced by Jansen et al. (2010) : M. luteola formed a separate clade that
probably had a common ancestor with the ritae'rgroup of species > 25 Ma, which indicates
that it is indeed a relict of an ancient Myrmica fauna.
M. luteola has many features of the so-called "inquiline syndrome" (see Wilson 1971,
1984), i.e. strongly reduced and simple non-pectinate spurs on the middle and hind tibiae,

197
somewhat developed ventral petiolar and postpetiolar processes, a hairy body and espe-
cially very small queens, which are even smaller than the workers. Moreover, uniquely for
the genus, the queens have no propodeal spines, the propodeum being only angled with at
most very short blunt denticles. Additionally, the workers and queens show another feature
that very rarely occurs in Myrmica species: the first gastral tergite is distinctly longitudinally
striated basally.
Ecology. M. luteola is widespread throughout temperate East Asia but seems to be
rare throughout its distribution. It inhabits spruce and southern-type, warm mixed forest,
where it prefers glades and the meadows bordering rivers and lakes; the highest altitude
recorded for the habitats of this species is 1820 m a.s.l. in North Korea (Radchenko 2005).
In the Russian Far East, nests were found in tree logs and stumps, but in Korea and
Japan, they also were built in soil, often under stones or around tree roots. Sexuais are
present in the nest from June to September, in Japan the nuptial flight take place from
September to mid-October (see also Kupyanskaya 1990; Imai et al. 2003; Radchenko
2005).
Based entirely on morphological characteristics, we speculated that M. luteola might be
at least a temporary social parasitic microgyne form, derived from a species such as
M. mirabilis (Elmes and Radchenko 1998, 2003a). Recently Masuko and Terayama (2002)
showed that M. luteola might be temporary social parasite on Manica yessensis Azuma;
if they are correct then M. luteola would be the first example,of intergeneric parasitism by
a Myrmica species. This is not implausible as there seems to be a tendency for social para-
sites to use less related hosts the older the species is in geological time (see Jansen et al.
2010). On the other hand, it seems unlikely because the Japanese evidence is open to
several other interpretations and there are no Manica species in the Russian Far East, Korea
and China. Thus over most of its range M. luteola must either parasitize a different host
or it establishes its nests independently. This interesting "old" species deserves more
study.

Map 68. Distribution of M. luteola.

198
 Myrmica margaritae Emery, 1889
(Fig. 164)

Myrmica margaritae Emery, 1889: 502, w, Burma; Dalla Torre 1893: 112; Forel 1903: 696; Bingham
1903: 267; Emery 1921: 38; Weber 1950: 219; Cllapman and Capco 1951: 127; Radchenko 1994b:
44; Bolton 1995: 281; Radchenko and Elmes 1998: 5; Radchenko et al. 2001: 217, nec Eidmann
1941: 15; Wu and Wang 1995: 92; Wei С et al. 2001: 561, misidentifications.

Type localities and type specimens. "Tenasserim: Mooleyit, 1000-1900 m."

Material examined. Lectotype (designated by Radchenko et Elmes 1998), w, "Tenasserim, M. Moo­
leyit, 1800-1900 m, Fea - Marzo 1887" "TYPUS", "Myrmica Margaritae Em., n. sp.", "Museo Civico
di Genova", "Lectotypus design. Radchenko et Elmes[ 1998" (GENOA); paralectotype, w "Tenasserim,
M. Mooleyit, 1000-1300 m, Fea Apr. 1887" "TYPUS", "Myrmica Margaritae Em.", "Margaritae Em."
(GENOA).
Distribution (Map 69). It is known only from the type locality in northern Burma.
Etymology. M. margaritae: С. Emery probably named this species for his wife
who was known as Rita (see etymology of M. ritae) being a diminutive of Margarita.
Emery used the name for new species in two other genera and Forel also used it in two
genera.
Notes. M. margaritae belongs to the ritee-complex of the ritae species group. It is known
until now only from the two type specimens, collected in 1887 in northern Burma. The most
distinctive feature of this species is the extremely coarse sculpture on its head dorsum,
where there are only four rugae between frontal carinae level with the eyes (only M. pul­
chella, M. sinensis and M. emeryi are known to have similar coarse rugosity). A second un­
usual feature appears to be the very few hairs on the occipital margin and alitrunk dorsum.
When we revised the ritae species group (Radchenko and Elmes 1998) we believed there to
be only one specimen that we designated as the' lectotype. We thought that the lack of hairs
was probably an artefact because the hairs of old specimens get brittle and quite often are

Map 69. Distribution of M. margaritae.

I
199
broken off. However, our recent (2007) discovery of a second type specimen in GENOA
confirms the quite unusual lack of pilosity in this species.
Before our recent revisions of the ritae-group (Radchenko and Elmes 1998, 2001a,
2009c; Radchenko et al. 2001, 2008; Radchenko, Elmes and Viet 2006), the taxonomic
situation within this group was uncertain. As a result, authors somewhat arbitrarily
assigned ritae-group species to M. ritae or M. margaritae, hence the records of
M. margaritae for China (Eidmann 1941; Wu and Wang 1995; Wei С et al. 2001) really
belong to other species of the ritae-group. For example, the record by Wei et al. (loc. cit.)
is at least partly M. angulata (corresponding material examined). Queens and males are
unknown.
Ecology is unknown, except that workers were found at altitudes between 1000 and
1900 m a.s.l.

Myrmica martensi Radchenko et Elmes, 1998

(Figs 165, 166)

Myrmica martensi Radchenko et Elmes, 1998: 21, w, q, Nepal; 2001b: 243.

Type localities and type specimens. Nepal, for details see material examined.
Material examined. Holotype, w, "Nepal, Gosainkund, Sing Gyang, 3200 m, 26.04.1973, leg. J.
Martens" (BASLE); paratypes: 3 w, 1 q, with same label (probably same nest) as holotype (BASLE).
Distribution (Map 70). It is known only from the type locality in Nepal.
Etymology. M. martensi: named for the collector, the German zoologist and Himalayan
expert, Prof. Jochen Martens of the University of Mainz.
Notes. M. martensi belongs to the boltoni-complex of the ritae species group. It is closely
related to M. boltoni and differs from it by shorter, upward-pointing and distinctly divergent

Map 70. Distribution of M. martensi

200
propodeal spines, by a distinctly shorter petiole and by its more feebly developed sculpture
on alitrunk and pedicel. Males are unknown.
Ecology is unknown.

Myrmica mirabilis Elmes et Radchenko, 1998

(Fig. 167)

Myrmica mirabile Elmes et Radchenko, 1998: 218, w, Taiwan.

Myrmica mirabilis: Radchenko et al. 2007: 1499 (emendation of spelling).
I
Type localities and type specimens. Taiwan, for details see material examined.
Material examined. Holotype, w, [Taiwan] "Alishan [= Arisan], 21.10.1977, leg. K. Yamauchi
(LONDON); paratypes: 10 w from the same nest (LONDON, KIEV, ELMES).
Distribution (Map 71). It is known only from the type locality in Taiwan.
Etymology. M. mirabilis: from the Latin mirabilis = strange, wonderful or extraordinary,
to emphasise its unusual features for a Myrmica species.
Notes. M. mirabilis is a most unusual Myrmica species because superficially, by
general shape it resembles many Pogonomyrmex species. Furthermore, it is a very
large and robust Myrmica species (HW 1.62-1.83, AL 2.56-2.92 mm), second in size
only to M. titanica, which appears smaller because it is much more slender than
M. mirabilis.
The most similar species is M. luteola and even though M. mirabilis has no "socially-
parasitic" features (e.g. its tibial spurs are not reduced, its petiole and postpetiole lack ven-
tral process, etc.) we provisionally place M. mirabilis to the luteola species group. Perhaps
the taxonomic position of this species will be clarified once queens and males are found.
Queens and males are unknown.

Map 71. Distribution of M. mirabilis.

201
 Ecology. Nothing is known about this species except that the nest was found in
decaying wood on a forest margin.

Myrmica mixta Radchenko et Elmes, 2008

(Fig. 168)

Myrmica mixta Radchenko et Elmes, 2008: 770, q, China (in: Radchenko et al., 2008).

Type localities and type specimens. China, for details see material examined.
Material examined. Holotype, q, "CH[China], N Sichuan, NW Dujiang-yan, 3 km WSW Shaping-
guan, 31-10-50 N/103-28-01 E, H = 1490 m, 12.vii.05, ¡eg. I. Belousov and I. Kabak" (PETERS-
BURG).
Distribution (Map 72). It is known only from the type locality in Sichuan Province of
China.
Etymology. M. mixta: the name is derived from the Latin mixtus = mixed or mingled,
to indicate that it has features from two different species groups.
Notes. This species was described from a single queen, collected in Sichuan Province of
China. M. mixta has an unusual combination of morphological features that seems to be in-
termediate between the ritae and inezae species groups. Its fairly coarse body sculpture,
very long propodeal spines and fig-shaped postpetiole might lead one to place it in the ritae-
group. Yet it differs from that group by its relatively short scape, not-pointed-apically
propodeal lobes, not-well-marked upper lateroventral corners of head and short and
rounded petiolar node. M. mixta is most similar to the Himalayan species M. rigatoi, but dif-
fers from it mainly by its wider frons, shorter scape and differently shaped propodeal spines
and petiole. We originally placed M.rigatoiin the n'toe-group when we described it but later
transferred it to the inezae-group (Radchenko and Elmes 2001). We now also provisionally

Map 72. Distribution of M. mixta.

202
place M. mixta in the inezae-group although a final decision will depend upon the dis-
covery of its workers and males. Workers, and males are unknown.
Ecology is unknown.

Myrmica multiplex Radchenko et Elmes, 2009

(Figs 169, 170)

Myrmica multiplex Radchenko et Elmes, 2009a: 81, w, q, China.

Type localities and type specimens. China, for details see material examined.
Material examined. Holotype, w, "China, Shaanxi Prov., Qin Ling Shan, 108 km SW Sian, Au-
torote km 93 S Zhouzhi, 107°56'E, 33°45'N, H 1650 m, No. 962, l-2.ix.1995, leg. Wrase" (KIEV);
paratypes: 3 w, 3 q, with the same labels as holotype (KIEV, ELMES, SCHULZ).
Distribution (Map 73). It is known only from the type locality in Shaanxi Province of China.
Etymology. M. multiplex: name derived from the Latin word multiplex = various or di-
verse, to reflect the mixed and various kinds of sculpture on the alitrunk.
Notes. M. multiplex belongs to the pachei species group. It is characterized by the long,
subovai head (CI > 1.20) with the convex sides and occipital margin, and the very widely
rounded, barely marked occipital corners; margins of the head with not very abundant long
suberect hairs. Its pronotum has coarse, short sinuous rugae and reticulation, while the
mesonotal and propodeal dorsum has quite coarse transversal rugosity (numbering < 20),
similar rugae are also on the upper parts of mesopleura and propodeum. It differs from the
related M. weii by its distinctly higher petiole with a relatively short dorsal plate (see also
Key, Chapter 5.7). Males are unknown.
Ecology is unknown except that workers and gynes were found on a mountain at an
altitude of about 1700 m a.s.l.

Map 73. Distribution of M. multiplex.

203
 Myrmica myrmicoxena Forel, 1895
(Fig. 171)

Myrmica myrmicoxena Forel, 1895: 143, q, Switzerland; Emery 1908a: 181, q, m; Forel 1910: 29, q,
m, 1915: 28, 30, 31, q, m (in Key); Bondroit 1918: 103; Emery 1921: 38 (misspelled as myrmeco-
xena): Finzi 1926: 114 (misspelled as myrmecoxena); Arnoldi 1930: 270 (misspelled as myrmeco-
xena); Weber 1950: 218 (misspelled as myrmecoxena); Bernard 1967: 116 (misspelled as
myrmecoxena): van Boven 1970: 129; Kutter 1973: 257, 1977: 67, q, m; Seifert 1988: 41, 1994: 17;
Bolton 1995: 281; Radchenko and Elmes 2003a: 220, q.

Type localities and type specimens. "Von Professor Bugnion in den Shweizer Alpen".
Material examined. Lectotype (designated by Radchenko and Elmes 2003a), q (upper specimen
on the pin with 3 queens), "Typus", "M. myrmicoxena, q Type bei M. lobicornis Anzeindaz", "Coli.
A. Forel", "Lectotype M. myrmicoxena Forel Des. Radchenko et Elmes, 2002" (GENEVA); paralec-
totypes: 2 q on the same pin with lectotype (both one without postpetiole and gaster); 2 q (one wit-
hout head), "Typus", "Myrmica! mixtossima" [Sicì], Anzeindaz, 20 août 69", "M. myrmicoxena For.
q Type, Anzeindaz" (GENEVA); 1 q (with 1 worker of M. lobulicornis on the pin), "lobicornis q anor-
male varieté, parmi des w normales. Anzeindaz, Alp. Vaudoisses, 1900 metres, 20 août", "Myrmica
myrmicoxena For.", "Syntypus Myrmica myrmicoxena Forel, 1895", "Musco Civico di Genova"
(GENOA).
Distribution (Map 74). Switzerland, northern Italy.
Etymology. M. myrmicoxena: from the genus name Myrmica combined with xena which
is derived from the Greek word xenos = guest or stranger, the name literally means "guest
of Myrmica [ants]".
Notes. Forel (1874, pp. 78-79) described from the Swiss Alps unusual queens and males
of Myrmica that were found in a nest of M. lobicornis (note we examined a host worker sent
with the paralectotype queen, that we determined as M. lobulicornis, see notes to M. lobi-
cornis above). However, he did not name them for another 20 years; then he made an
extremely brief description of the queens, calling them M. myrmicoxena (Forel 1895), but

Map 74. Distribution of M. myrmicoxena.

204
he did not include the males to the type series. Emery (1908a) made a formal description
of the males, based on Forel's (1874) original description, and Forel (1910, 1915) made
further short additions to the description of queens and males. Kutter (1977) made some
new notes on M. myrmicoxena and published the first drawings of the queen and male, and
Seifert (1988) followed Kutter's description. Despite these more recent descriptions we
redescribed queens based on our approach to Myrmica taxonomy (Radchenko and Elmes
2003a).
The queens of M. myrmicoxena are most similar to those of M. arnoldii, differing mainly
by a somewhat narrower frons and smaller size, this led us earlier to place both species to
the myrmicoxena species group (see Radchenko and Elmes 2003a). However, a close rela-
tionship between the European M. myrmicoxena and Siberian M. arnoldii seems unlikely.
Furthermore, the antennae of M. myrmicoxena males (as drawn by Kutter 1977) have the
normal 13-segments, whereas those of M. arnoldii are 12-segmented. Hence we now return
to Radchenko's (1994b) first opinion (M. arnoldii represents a separate monotypic species
group - see Notes to that species, above) and place M. myrmicoxena in the monotypic myr-
micoxena species group.
We have examined photographs of a specimen taken by Mr. Mike Lush (UK) in June
2009 (collection details MJL390; 14-vi-2009; SWITZ: Fiescheralp, Feisch, Brig;
46°24.558'N, 8°5.907'E; 2213m) and have no doubt that it is M. myrmicoxena.
Ecology and biology. Based on a sample of one worker the original host was M. lobuli-
cornis living at 1900 m. a.s.l. (probably under a stone). It is probably workerless and
almost certainly a social parasite.
For more than 100 years the type specimens taken in 1869 were the only record for this
species until recently Mike Lush captured a specimen in a M. lobulicornis nest (see above)
and specimens have been found by different collectors in S Tyrol, Italy and Valláis, Switzer-
land (Bernhard Seifert, pers. comm.). Mike Lush (pers, comm.) wrote "When I turned over
the stone [in alpine grassland] the ants did their normal thing of running around franti-
cally, except for a small cluster of about 20 workers that I eventually realised were holding
this queen down, preventing her movement. I didn't wait around to see whether she would
escape (it had already been a minute or so whilst I was scanning for interesting things that
were actually moving), so pooted her and the workers! She seemed unharmed by her cap-
tors, so I assumed that this might be how she acquired the colonies scent and that they
were doing it before I overturned the stone".

Myrmica nitida Radchenko et Elmes, 1999

(Figs 172-174)

Myrmica nitida Radchenko et Elmes, 1999a: 35, w, q, m, India; 2001b: 261.

Type localities and type specimens. India, for details see material examined.
Material examined. Holotype, w, "Kashmir, 1 km NE Yehmer Pass, 34°13'N, 75°10'E, 3600 m,
06.08.1978, leg. P. Ward, acc. No 3044; alpine vegetation, under stone" (LONDON); paratypes: 9 w
from the same nest as holotype; 17 w, 4 q (alate), 2 m, "Kashmir, Sanang, 2600-2750 m, leg. W.
Wittmcr" (LONDON, BASLE, WARD, KIEV, ELMES).
Distribution (Map 75). It is known only from the type localities from India
(Kashmir).

205
 Etymology. M. nitida: from the Latin word nitidus = shining, to describe its generally
smooth and shiny appearance.
Notes. By the general shape of their alitrunk, workers of M. nitida are similar to species
from the cachmiriensis complex of the ragosa-group, but its males have a long antennal
scape. By this feature of males, M. nitida should be related to the smythiesii-group but the
workers do not fit with this group, because their frontal carinae do not curve outwards and
do not merge with the rugae surrounding the antennal sockets. Moreover, the workers'
alitrunk appears smooth and shiny with only fine superficial striation on the sides; similarly
reduced sculpture is rather rare and occurs only in a few Myrmica species (i.e. M. villosa,
M. inucta). Thus, we consider the taxonomic position of M. nitida as uncertain.
Ecology is poorly known. One colony was found living in the same habitat in Kashmir
as M. rhytida, at 3600 m, under a stone, on small ridge on the west side of a lake, with scat-
tered juniper and rhododendron shrubs and a ground cover of various herbs and flowers
(Philip Ward, pers. comm.).

Map 75. Distribution of M. nitida.

Myrmica obscura Finzi, 1926

(Figs 175-177)

Myrmica schencki var. obscura Finzi, 1926: 111, w, q, m, Italy; Weber 1948: 302; Baroni Urbani
1971:36.
Junior synonym of M. schencki: Bernard 1967: 118; Radchenko 1994e: 77; Bolton 1995: 281; Seifert
2003: 145; Radchenko, Elmes and Alicata 2006: 513.
Junior synonym of M. lobicornis: Seifert 1988: 38 (part.).
Myrmica obscura: Elmes et al. 2009: 99.

Type localities and type specimens. "Io l'ho raccolta sul Monte Nanos (1200 m. - Venezia Giulia
- V. 1923 e IX. 1924) e designo come tipi gli esemplari provenienti da questa località".

206
 Material examined. Syntypes, 2 w, "M. Nanos V.23 Finzi", "typo", "B. Finzi coll.", "Pur. 1950",
"MCZ type 28794"; 2 w with the same labels (only without "tipo" label); 1 q, "M. Nanos 15.9.24
Finzi", "typo", "B. Finzi coll.", "Pur. 1950", "MCZ|type 28794"; 1 m with the same labels (only wi-
thout "tipo" label) (HARVARD) (sec also Notes below); non-type material: about 400 w, 33 q, 23 m
from Italy.
Distribution (Map 76). Italy south of Alpes.
Etymology. M. obscura: named from Latin obscure = dark or shaded, originally to indi-
cate that the types were a dark form of M. schencki.
Notes. M. obscura belongs to the schencki species group; it was originally described as
a variety of Ai. schencki and is closely related to that species. It spent most of its taxonomic
history in synonymy with M. schencki until we recently revived the name from synonymy
and raised it to species rank (Elmes et al. 2009).
When we revised the schencki species group (Radchenko, Elmes and Alicata 2006) we
examined only 4 syntypes workers, queen and] male of var. obscura, collected by Finzi in
Monte Nanos (see above), and found slight differences from typical M. schencki: the male
had a relatively shorter scape, and the workers and queen had a somewhat wider frons.
Morphometrically, the workers and queen seemed somewhat intermediate to M. caucasicola
but based on a discrimination analysis the queen and males were assigned M. schencki
rather than M. caucasicola. Subsequently, we obtained a considerable amount of both
M. schencki and M. obscura collected from several Italian sites being studied by our col-
league Francesca Barbero in connection with the conservation of Phengaris butterflies (see
Elmes et al. 2009). Morphometric analysis showed that the Apennine populations that we
equated to M. obscura, had much wider frons (mean FI = 0.266) compared with M. schen-
cki populations sampled from over its geographical range (mean FI = 0.234), furthermore
the M. obscura males had shorter scapes (mean Sl2 = 0.451 vs. 0.419) that were signifi-
cantly less angled (more obtuse) at their base (mean 133° vs. 121°).

Map 76. Distribution of M. obscura.

207
 We consider M. obscura to be a separate Italian "glacial" lineage of M. schencki with
consistently different diagnostic features sufficient to consider it as a separate species (fol-
lowing the precedent of how geographical subspecies have been treated by other authors and
in this monograph). The M. schencki populations in the southern French alps have some-
what wider frons than is typical for M. schencki and we suggested that M. obscura has
invaded to the Alps from the south and M. schencki has penetrated into central Italy from
the northand that there has probably been some gene flow between the two species in
northern Italy and south-eastern France (see Elmes et al. 2009).
We also examined 1 worker specimen from LONDON, labelled as "Kelbra, Germania",
"Kiffhäuser 9.28 Krausse", "В. Finzi Coll. Purch. 1950", "M.C.Z. type 28794", "Syntype",
"Myrmica schencki obscura Finzi det. WL Brown". This can not be a type specimen of Myr­
mica obscura, because it was collected later and from another place than the type speci­
mens. Furthermore, it is a worker of M. lobicornis. Seifert (1988: 38) examined two workers
(no location of the collection given) from the same nest series, and therefore erroneously
synonymised var. obscura with M. lobicornis.
Ecology. The ecology of M. obscura is very similar to that of M schencki (see notes on
ecology of that species, below). It is quite common on hot well-grazed alpine meadows
where it builds nests in the soil and constructs the woven funnel entrance that we have ob­
served is typical for many schencki-group species in the Old World and "schencki-gvoup-
like" species in the North America. Nests average about 400 workers with usually a single
queen but two or more queens are not uncommon. Our impression is that M. obscura might
be able to tolerate slightly hotter and drier conditions than M. schencki but at this time
there is no firm data to support this hypothesis.

Myrmica onoyamai Radchenko et Elmes, 2006

(Figs 178-180)

Myrmica onoyamai Radchenko et Elmes, 2006: 507, w, q, m, Japan (in: Radchenko, Elmes and Alicata
2006).

Type localities and type specimens. Japan, for details sec material examined.
Material examined. Holotype, w, Japan, Yamamashi Pref., ca 15 km NE of Sudarmi interchange,
nr. Hokuto City, By Shiokawariv.nr. Shiokawa Dam. с. 900 m No. J-99, 24.vu.2002, leg. G. W. Elmes
(LONDON); paratypes: 64 w, 25 q, 1 m from the nest of the holotype (LONDON, KIEV, ELMES,
HELSINKI) (see also Notes below).
Distribution (Map 77). It is known only from the type locality from Japan (Hokkaido).
Etymology. M. onoyamai: we dedicated this species to the Japanese myrmecologist Prof.
Keiichi Onoyama of Obihiro University, Japan.
Notes. Unfortunately, in the original description of this species (Radchenko, Elmes and
Alicata 2006) we wrongly gave the type locality as "Japan, Isl. Honshu, Nagano Pref., about
11 km E of Matsumoto on road to Tobira pass, ca. 1400 m a.s.l.". This site where we also
collected Myrmica samples, is some tens of kilometres from the actual site on Honshu Is­
land where we collected the holotype series (see above).
M. onoyamai belongs to the schencki species group. Its workers and queens well differ
from those of all other members of this group by a distinctly wider frons and by pro­
portionally less extended frontal lobes: in workers mean FI 0.35, mean FLI 1.20 vs. mean

208
 Map 77. Distribution of M. onoyamai.

FI < 0.33, mean FLI > 1.28 in other schencki-group species. The male of M. onoyamai
most resembles males of M. koreana, but differs from them by shorter hairs on the head mar­
gins, by the much less sculptured, almost smooth scutum and scutellum, by the not angled
dorsal plane of the scape and by wider postpetiole.
Ecology. The nest of this species was found in anthropogenic conditions that were quite
exposed, hot and dry compared to general surrounding woodland, and the location may not
be typical for this species. It was under a small stone in a pile of sandy gravel dumped by
the side of the road, possibly to grit the road in icy conditions, although it had been there
some time because its surface was covered in moss and lichens. The road was in a river val­
ley in an area of mixed woodland at an altitude ca. 900 m a.s.l., about 10 m from the river.
We suggested that M. onoyamai living in more natural conditions might be associated with
warmer exposed biotopes such as sub-alpine grassland and sandy screes at the base of cliffs.

Myrmica ordinaria Radchenko et Elmes, 1999

(Figs 181, 182)

Myrmica ordinaria Radchenko et Elmes, 1999a: 41, w, q, India, Pakistan; 2001b: 252.

Type localities and type specimens. India and Pakistan, for details see material examined.
Material examined. Holotype, w, "India, Kashmir, Seven Springs. 29.07.86, leg. P. Williams"
(LONDON); paratypes: 23 w, 1 q, with the same label as holotype; б w, "Pakistan, Kalam, 2300 m,
12.07.94, leg. S. Dakatra" (LONDON, BASLE, MILAN, KIEV, ELMES).
Distribution (Map 78). It is known only from the type localities from India (Kashmir)
and Pakistan.
Etymology. M. ordinaria: name derived from Latin word Ordinarius = regular, ordinary,
to indicate that this species falls somewhere in the middle of the range of variation of mor­
phological features for genus Myrmica as a whole and in this sense is ordinary.

209
 Notes. We placed M. ordinaria to the cachmiriensis complex of the rugosa species group
based only on the features of the female castes because they most resemble M. wardì and
M. cachmiriensis. Both queens and workers differ from M. wardì by having very finely punc-
tated, more or less smooth surfaces between the rugae of the head and alitrunk; workers
differ from those of M. wardi by having a flat (not concave) mesonotal dorsum and from
M. cachmiriensis by the much darker colour of their body and relatively longer subdecum-
bent pilosity on the sides of the head. Males are unknown.
Ecology i s u n kn о wn.

Map 78. Distribution of M. ordinaria.

Myrmica orthostyla Arnoldi, 1976

(Figs 183-185)

Myrmica orthostyla Arnoldi, 1976: 552, w, q, m, Kazakhstan and Kyrgyzstan; Tarbinsky 1976: 27 (mis­
spelled as orthostila); Radchenko 1994b: 42, 1994c: 142, 1994d: 81; Bolton 1995: 281.

Type localities and type specimens. "Тянь-Шань. Голотип w, аллотипы (Ç, S) и 3 паратипа:
заповедник Аксу-Джабаглы (Длусский), Чаткальский хребет (Тарбинекий)" [Tien-Shan. Holotype w, al­
lotypes (q, m) and 3 paratypes: Natural Reserve Aksu-Dzhabagly (leg. Dlussky), Chatkalsky Range
(leg. Tarbinsky].
Material examined. Holotype, w, "Аксу-Джабаглы, Ю. Казахстан, Длусский" [Aksu-Dzhabagly,
S Kazakhstan, leg. Dlussky], "18.8.AD8a", "Holotypus, Myrmica orthostyla K. Arnoldi" (MOSCOW);
paratypes: 3 w, 3 q, 5 m (including "allotypus") with the same label as holotype; 18 w, 1 q, 1 m
(damaged), "Киргизия, Чаткальский хр., пер. Чангама, 3500 m, 5.IX.969, Тарбинекий" [Kirgizia,
Chatkalsky Range, Changama Pas, 5.ix.969, leg. Tarbinsky] "Paratypus Myrmica orthostyla
К. Arnoldi"; 3 w, 1 q, with the same labels, but date 8.vii.968 (see Notes below); non-type material:
12 w, 1 q, 5 m from Tien-Shan and Alai.
Distribution (Map 79). T i e n - S h a n , Alai.

210
 Etymology. M. orthostyla: the name is a combination of the Greek words opQvoç (orthos)
= straight and awÁoç (stylos) = pillar or pole, to describe the straight stick or pole-like
propodeal spines.
Notes. Radchenko (1994b) originally placed M. orthostyla in the scabrinodis species
group but we now consider that it is a member of the tibetana-group: It is very similar to
M. tenuispina, but differs from it mainly by the thin, straight, long, stick-like propodeal spines.
When Arnoldi (1976) made the original description of this species he did not formally
describe the queens, however, 3 queens with the same labels as the holotype are included
in the type series and should be considered as paratypes. Arnoldi also included in the type
series material collected by Tarbinsky from Chatkalsky Range, but in his paper he gave the
number of the paratype workers as only 3 (i.e. just those workers collected by Dlussky in
Aksu-Dzhabagly). Almost certainly this was a simply a drafting error and both queens and
workers from Chatkalsky Range should be considered as the paratypes.
Ecology. M. orthostyla lives at quite high altitudes (about 2500 m) in various kinds of
mountains steppes. Nests were built in soil, usually under stones. Based on collecting data,
males were found in the nests in September, which indicates that mating flights are in late
summer (August-September).

Map 79. Distribution of M. orthostyla.

Myrmica pachei Forel, 1906

(Figs 186-188)

Myrmica pachei Forel, 1906: 79, w, q, m, Nepal; Emery 1921: 38; Menozzi 1939: 293; Weber 1947: 467;
Chapman and Capco 1951: 128; Collingwood 1970: 374; Bolton 1995: 281; Radchenko and Elmes
2001b: 256, 2009a: 69.

Type localities and type specimens. "Tseram, N. E. Nepal, à 3600 métrés, récoltée par M. Pache
dans un tronc d'abre ou elle nichant".

211
 Material examined. Lectotype (designated by Radchenko and Elmes 2001b), w, "NE Nepal,
Tseram, 3600 m, Pache" (GENEVA); paralectotypes: 2 vv (GENEVA) and 1 w (HARVARD) with
same label as lectotype; lw, lm, "Myrmica Pachei n. Nepal, Himalaya 3600 m" (GENOA); non-type
material: 23 w, 2 q from Nepal and Bhutan (for details see Radchenko and Elmes 2001b).
Distribution (Map 80). India (Kashmir), Nepal and Bhutan.
Etymology. M. pachei: dedicated to the collector, the Swiss climber Alexis Pache who
was killed in an avalanche in July 1905 while attempting to climb Kangchenjunga
(the World's third highest mountain). The expedition was led by the controversial occultist
and black magician Aleister Crowley and the Swiss doctor Jules Jacot-Guillarmod. Forel
had given Pache tubes to collect high altitude ants, eventually Pache's brother-in-law re-
turned two tubes found in his belongings, one of which contained the three castes of this
species.
Notes. M. pachei belongs to the pachei species group, a key feature of which is the trans-
versal rugulosity present on the alitrunk dorsum. Until recently we considered this unusual
feature for Myrmica species to be present in only a couple of other species from the
Himalaya (M. inezae and M. villosa). M. pachei differs from M. villosa by having much
shorter, straighter and sparser hairs on the body, by darker colour and by much more dense
sculpture on the alitrunk. It clearly differs from M. inezae (see notes to species groups
diagnosis, Chapter 2.2) therefore we place M. pachei and M. villosa together in the pachei-
group.
Recently we have found a whole suite of species from Tibet and southern China that
share, to some extent, the feature of transversal rugosity. This led us to redefine the features
of the pachei species group (Radchenko and Elmes 2009a); in that paper we added a fur-
ther twelve species to the group and provided a table of diagnostic features for the group.
Ecology is poorly known. Forel (1906) noted that type specimens were collected from
a colony in a tree trunk. All we know about the other material is that it was collected in
mountains at altitudes between 2600 and 3600 m a.s.l.

Map 80. Distribution of M. pachei.

212
 Myrmica pararitae R a d c h e n k o et Elmes, 2 0 0 8
(Figs 189, 190)

Myrmica pararitae Radchenko et Elmes, 2008: 773, w, q, China (in: Radchenko et al. 2008).

Type localities and type specimens. China, for details see material examined.
Material examined. Holotype, w, "China, Sichuan Prov., Maoxian, 40 km N Winchuan, 1800 m,
23.viii.2002, leg. S. Murzin and I. Shokhin (PETERSBURG); paratypes: > 100 workers, 17 queens
with the same label as holotype (PETERSBURG, KIEV, ELMES, GUILIN).
Distribution (Map 81). The species is known from the type series from Sichuan Province
of China.
Etymology. M. pararitae: from the Greek жара (para) = near, next to, close to, and
"ritae" to indicate its similarity to M. ritae.
Notes. M. pararitae belongs to the ritae-complex of the ritae species group and is simi­
lar to M. ritae, but well differs from any other species of the ritoe-complex by the sculpture
of head, with only longitudinal rugae and without any reticulation. It also differs from
M. ritae by the broadly subovale head, shorter scape, presence of long hairs on the temples
and genae, and by its brownish-red head that does not-strongly contrast with the darker
alitrunk.
Males and ecology are unknown.

Map 81 Distribution of M. pararitae.

Myrmica pelops Seifert, 2003

(Fig. 191)

Myrmica pelops Seifert, 2003: 148, w, Greece; Radchenko, Elmes and Alicata 2006: 527.

213
 Type localities and type specimens. "7 samples from the Peloponissos peninsula in Greece: Holo-
type and 2 paratypes, labelled "Greece, Peloponessos Taigetos Mts. Laganda Pass (20 km E Kalamata)
1.6.1994 1100-1400 m Schulz Igt.", SMN Goerlitz; 9 paratypes, labelled "GREECE - Peloponnisos
1347 Taigetos Oros, Laganda Pass, 20 km E. Kalamata, 1100-1400 mH, leg. Schulz, Vock 01.6.1994",
SMN Goerlitz; 5 paratypes, labelled "GREECE - Peloponnisos 1420 Killini, Nordseite, 42 km W Ko-
rinthos, 1200-1400 mH, leg. Schulz, Vock 05.6.1994", SMN Goerlitz. Further worker paratypes with
the following labelling are in the private collection of Andreas Schulz: "GREECE - Peloponnisos 1394
Chelmos, 30 km SSE Egion, 1800-2200 mH, leg. Schulz, Vock 04.6.1994", "GREECE- Peloponnisos
1368 Taigetos Oros, unterhalb des Profitis Ilias, 1800-2000 m, leg. Schulz, Vock 02.06.1994", and
"GREECE - Peloponnisos 1404 Killini, Nordseite, 45 km W Korinthos, 1600-1800 m, leg. Schulz,
Vock 05.6.1994".
Material examined. Paratypes: 4 w (from the nest of holotype), "Greece, Peloponnesus, Taigetos
Mts., Laganda Pass (20 km E Kalamata) 1.6.1994, 1100-1400 m A. Schulz leg."; 3 w, "Greece, Pelo-
ponnisos, Taigetos Oros, unterhalb des Profitis Ilias, 1800-2000 m, No 1368, 02.vi.1994, leg. Schulz
and Vock" (KIEV, SCHULZ, ELMES, WERNER).
Distribution (Map 82). It is known only from the type localities from Greece (Pelopon-
nesus Peninsula).
Etymology. M. pelops: named for Pelops, the mythical Greek hero who gave his
name to the region of Greece where the type series was found (the Peloponnesus Penin-
sula).
Notes. M. pelops belong to the schencki species group. Seifert stated that it relates to
M. ravasinii, based on its quite large size, a somewhat similar scape structure and the shape
and sculpture of its petiole. However, M. pelops clearly differs from the latter by a much
wider frons (mean FI > 0.22 vs. < 0.20), and by a clearly much smaller lobe at the scape
bend. On the other hand, the FI of M. pelops is similar to that of M. schencki, but it well
differs from this species by a distinctly different shaped scape lobe, by coarser rugae on the
body, and by a shorter and stouter petiole with a short anterior peduncle. Queens and males
are unknown.

Map 82. Distribution of M. pelops.

214
 Ecology is poorly known. Seifert (2003) noted that all samples of M. pelops were found
in very light, devastated coniferous forests, so, this species seems to be rather xerophilous.

Myrmica petita Radchenko et Elmes, 1999

(Fig. 192)

Myrmica petita Radchenko et Elmes, 1999a: 45, q, India; 2001b: 261.

Type localities and type specimens. India, for details see material examined.
Material examined. Holotype, q, "Kashmir, Yusmar, 2300-2400 m, 6.07.1976, leg. W. Wittmer"
(BASLE).
Distribution (Map 83). It is known only from the type localities from India (Kashmir).
Etymology. M. petita: from the French petite = little, small (or dainty when applied to
females) to indicate its small size for a Myrmica queen.
Notes. M. petita is known only from a single holotype queen and has an uncertain
taxonomic position, we can not satisfactorily place it in any species group. It most resem-
bles the workers of M. wittmeri (queen unknown) differing only by the punctated surface
of its head dorsum and relatively longer propodeal spines. However, we do not think this
queen is M. wittmeri because we know of no Myrmica species where the propodeal spines
of the queens are relatively longer than those of their workers. M. petita very small, smaller
than all known free-living queens from the Himalayan Myrmica fauna and is one of the
smallest of all known free-living Myrmica species. Small size is often associated with social
parasites but we think that M. petita is unlikely to be a social parasite because it lacks any
of the typical "parasitic" characters, e. g. reduced pectinate spurs, ventral lobes on the
petiole and postpetiole, relatively wide postpetiole and general hairiness. Workers, males
and ecology are unknown.

Map 83. Distribution of M. petila.

215
 Myrmica phalacra R a d c h e n k o et Elmes, 2 0 0 9
(Figs 193, 194)

Myrmica phalacra Radchenko et Elmes, 2009a: 74, w, q, China.

Type localities and type specimens. China, for details see material examined.
Material examined. Holotype, w, "China, Shaanxi Prov., Qin Ling Shan, 118 km E Xian, 110°06'E,
34°25'N, S top 1950-2000 m, No. 969, 19.viii.1995, leg. Wrase" (KIEV); paratypes: 1 w and 2 q with
the same label as holotype (KIEV, ELMES, SCHULZ).
Distribution (Map 84). It is known only from the type locality from Shaanxi Province
of China.
Etymology. M. phalacra: from the Greek word tpaXaxpoç (phalachros) = bald, to de-
scribe the complete absence of hairs, even subdecumbent ones, on the sides on the head.
Notes. M. phalacra belongs to the pachei species group. It is characterized by the quite
coarse transversal sinuous rugosity (numbering < 20) on the mesonotal and propodeal dor-
sum and by the absence of the decumbent hairs on the lateral margins of head posterior to
the eyes. In some respects, it resembles M. hlavacì, but differs from it by partly reduced
rugosity on the head dorsum, where the frons (level with the eyes) has < 15 fine, slightly
sinuous longitudinal rugae, with only some of them extending unbroken to the occipital
margin. Males are unknown.
Ecology is unknown except that two workers and two gynes were found on mountain
meadows at the altitudes ca. 2000 m a.s.l.

Map 84. Distribution of M. phalacra.

Myrmica pisarskii R a d c h e n k o , 1994

(Figs 195-197)

Myrmica pisarskii Radchenko, 1994a: 208, w, q. ni, Russia, Mongolia; 1994b: 43, 1994c: 142, 1994f:
90, 1994g; 109; Bolton 1995: 282; Pfeiffer et al. 2007: 5.

2JÓ
Myrmica saposhnikovi: Pisarski 1969a: 228, 1969b: 296; Dlussky, Pisarski 1970: 85, misidentifica-
tìons.

Type localities and type specimens. South Siberia and Mongolia, for details see material examined.
Material examined. Holotype, w, "Transbikalien, Chitinskaja obi., Daursky Natural Reserve,
Irelgin, steppe, nest undena stone, No. 51-90, 7.VII.1990, A. Radchenko" (KIEV); paratypes: 10 w
from the same nest; 2 w, the same locality, No. 57-90, 7 VII 1990 (A. Radchenko); Chitinskaya obi.:
5 w, Borzia, No. 3-90, 28 VI 1990 (A. Radchenko); 9 w, left bank of river Onon, near village of N. Tsas-
suchci, No. 75-90, 12 VII 1990 (A. Radchenko); 5 w, Borzinsky distr., Mt. Adon-Cholon, 1000 m, No.
92-90, 17 VII 1990 (A. Radchenko); 4 w, viii. N. Tsassuchei, No. 1-90, 28 VI 1990 (A. Radchenko);
2 w, the same locality, pine forest, 12 VII 1990 (A. Radchenko); 7 w, Ononsky distr., Mt. Kuku-Chodan,
No. 113-90, 19 VII 1990 (A. Radchenko); 15 w, 6 m, SE Altai, Kuraiskaya kotlovina, steppe, Nos.
552, 580, 609, 612, 614, 638, 1 VI - 8 VIII 1964 (Z. Zhigulskaya); Mongolia: 2 w, 1 q, 4 m, Central
Aimak, Kerulen, 45 km E from Samon Bajanderler, 1400 m, exp. Kaszab, No. 304, 26 VII 1965; 3 w,
3 m, Uburchanpai Zuil, Od Bijar, 7-8 VIII 1985; 1 w, Tevtrulech, 17 VII 1979; 1 w, Kiransky Karaul,
4 VIII 1908 (Chornos); 4 w, Ara-Changajsky Aimak, lake Uchish-Noor (A. Ponomarenko); Tuva: 2 w,
Kiachta, No. 1135, 2 VIII 1962 (Z. Zhigulskaya); 2 w, the same locality, No. 1175, 6 VIII 1962 (Z.
Zhigulskaya); 9 w, near Kyzyl, No. 18, 13 VI 1961 (Z. Zhigulskaya); 3 w, Ubsu-Noorskaja kotlovina,
lake Dus-Hol, No. 297, 25 VII 1962 (Z. Zhigulskaya); 10 w, 2 q, the same locality, 10 km N lake Ters-
Hol, No. 160, 7 VI 1962 (Z. Zhigulskaya); 10 w, same locality, 3 km S village Torgalyk, No. 220, 25
VI 1962 (Z. Zhigulskaya); 2 m, the same locality, Shevelych-Hem, No. 316, 1 VII 1962 (Z. Zhigul-
skaya); 9 w, left bank of riv. Enisei, No. 389, 6 IX 1962 (Z. Zhigulskaya); 2 w, 2 m, Mt. Tannu-Ola,
village Samagaltai, dry steppe, 7 VIII 1978 (J. Reznikova); Transbaikalia: 1 w (Th. Lukyanovitsch);
1 w, Troitskosavsk, 11 VII 1928 (Th. Lukyanovitsch); 1 w, Enchor, 20 VII 1928 (Th. Lukyanovitsch);
1 w, Minusinsk, 10 VI 1936 (Th. Lukyanovitsch); 2 w, Chita, 15 VI 1912 (Gytelman) (KIEV, MOS-
COW, WARSAW, ELMES); non-type material: > 100 w, 15 q, 9 m from South Siberia and Mongolia.
Distribution (Map 85). South Siberia from SE Altai to Transbaikalia, Mongolia.
Etymology. M. pisarskii: this species was dedicated to the memory of the well known
Polish myrmecologist Prof. Bogdan Pisarski (1928-1992), one time Director of the Mu-
seum of Institute of Zoology Polish Ac. Sci., Warsaw.

Map 85. Distribution of M. pisarskii.

217
 Notes. M. pisarskii belongs to the lobicornis species group. It is very similar to
M. kasczenkoi by the body sculpture, by shape of alitrunk and petiole, but well differs from
it by the presence of a vertical lobe at the scape base.
Records of M. saposhnikovi for Mongolia (Pisarski 1969a, b; Dlussky and Pisarski 1970)
belong to M. pisarskii (corresponding material examined).
Ecology. M. pisarskii is one of the most xerophilous Myrmica species. In Russia, it in-
habits steppes and dry meadows, occasionally it can be found in dry and sparse pine forests.
It builds nests in soil, often under stones. Micha! Woyciechowski took several series from
the Altai Mountains, Russia where the nests were found in slightly more humid conditions
in moss near a river (though such habitats can become very hot and dry in summer). Woy-
ciechowski also took a number of nest samples from northern Mongolia, all were found
living in the soil in meadow dry steppe (1000-1200 m. a.s.l.). Here soils were shallow and
rocky, trees were absent and the ground flora contained a variety of herbs (see Muehlenberg
et. al. 2000). Sexuais are present in nests in July-August, suggesting that nuptial flights
occur from late July to early September.

Myrmica pleiorhytida Radchenko et Elmes, 2009

(Figs 198, 199)

Myrmica pleiorhytida Radchenko et Elmes, 2009a: 78, w, China.

Type localities and type specimens. China, for details see material examined.
Material examined. Holotype w, "China, N Yunnan, env. Xinguan, 2400 m, 29.vii.2002, leg. S.
Murzin and I. Shokhin" (PETERSBURG); paratypes: 1 w, "China, N Yunnan, Zhongdian, 3300 m,
6-8.vii.2002, leg. S. Murzin and I. Shokhin"; 8 w, China, Yunnan, Habashan Mts., 3150-3500 m,
27°19'W, 100°08'E, 11, 16-18.vi.2004, leg. R.+ H. Fouqué (WGS 84)" (PETERSBURG, KIEV,
ELMES, VERONA); non-type material: 7 w, 2 q, China, Yunnan, env. Bailakou Pass, 3400-3800 m,
28.V.2006, leg. I. Shokhin; 7 w, lq, same locality, 28.v-7.vi.2006, leg. I. Shokhin et S. Murzin; 1 w,
China, Yunnan, Zhongdian Mts., Shangri-La, 3800-4300 m, 26.V.2006, leg. I. Shokhin (PETERS-
BURG, KIEV).
Distribution (Map 86). It is known only from Yunnan Province of China.
Etymology. M. pleiorhytida: from the combination of the Greek words nXf.iov (pleion)
= more, abundant or numerous, and pviiôa (rhytida) = crease or wrinkle (ruga), to de-
scribed the sculpture of the head and alitrunk dorsums.
Notes. M. pleiorhytida belongs to the pachei species group. It is characterized by the fol-
lowing combination of the features: the pronotal dorsum with reticulation and sides with
longitudinal rugulae; the mesonotal and propodeal dorsum with fine transversal rugae num-
bering > 35; lateral and occipital margins of the head with the long numerous suberect to
erect hairs; the head only slightly longer than broad (CI < 1.15), subsquare, with the dis-
tinctly marked occipital corners; whole head dorsum with the very dense, fine, almost
straight, slightly divergent posteriorly longitudinal rugae, their number between frontal cari-
nae level with the eyes > 30.
First description of queens.
The shape and sculpture of head is similar to that of workers (see also Radchenko and
Elmes 2009a). Scutum and scutellum longitudinally rugose, propodeal dorsum with trans-
versal rugosity. Petiolar node and postpetiolar dorsums, especially in their posterior half,

218
 Map 86. Distribution of M. pleiorhytida.

transversally rugulose. Propodeal spines somewhat shorter and stouter than in workers.
Colour and character of pilosity - as in workers.
Males are unknown.
Ecology is unknown except that the workers were found on mountain meadows at alti-
tudes between 2400 and 3500 m a.s.l.

Myrmica poldii Radchenko et Rigato, 2008

(Fig. 200)

Myrmica poldii Radchenko et Rigato, 2008: 776, w, China (in: Radchenko et al. 2008).

Type localities and type specimens. China, for details see material examined.
Material examined. Holotype, w, "China, S Sichuan, 2220 m, pass SW of Mianning, Rhododendron
primary deciduous forest, 14-21.vi.2004, leg. R. Fabbri" (MILAN).
Distribution (Map 87). It is known only from the type locality from Sichuan Province
of China.
Etymology M. poldii: this species was dedicated to the memory of the Italian myrme-
cologist Dr. Bruno Poldi (1920-2002) of Mantova.
Notes. M. poldii belongs to the draco-complex of the ritae species group (for details see
Radchenko and Elmes 1998, 2001; Radchenko et al. 2001a; Radchenko, Elmes and Viet
2006). It well differs from M. draco by its much coarser sculpture of petiole and postpeti-
ole and more reticulated sculpture of the head and alitrunk dorsum; in M. draco only the
occiput and promesonotal dorsum are partly reticulate. M. poldii differs from M. schoedli
by the presence of reticulation on the head dorsum (only sinuous rugae occur in
M. schoedli), its much more coarsely sculptured petiole and postpetiole, the reticulate
sides of alitrunk, and by the relatively shorter scape. From M. y amami it differs by the less
coarsely sculptured head dorsum, the inclination of propodeal spines, which are nearly

219
horizontal, while in M. yamanei they are inclined at more than 45°. Queens, males and
ecology are unknown.

Map 87. Distribution of M. poldii.

Myrmica polyglypta Radchenko et Rigato, 2008

(Fig. 201)

Myrmica polyglypta Radchenko et Rigato, 2008: 781, w, China (in: Radchenko et al. 2008); Radchenko
and Elmes 2009a: 72.

Type localities and type specimens. China, for details see material examined.
Material examined. Holotype, w, "China, Yunnan, Habashan Mts., 4000-4700 m, 27°19'W,
100°06'E, 8-9.vi.2002, leg. S. Bccvárand R.+ H. Fouqué" (MILAN); paratypes: 13 w with the same la-
bels as holotype (MILAN, KIEV); non-type material: 3 w China, Yunnan, Habasan Mts., 13-15.6.2004,
H 4000-4300 m, 27°19'N 100°06'E, leg. Fouqué R. + H. (MGS 84) (VERONA).
Distribution (Map 88). It is known only from Yunan Province of China.
Etymology. M. polyglypta: from a combination of the Greek words nolx> (poly) = many,
and уХлштос (glyptos) = carved, to describe the variable sculpture on the alitrunk.
Notes. M. polyglypta belongs to the pachei species group (see also Radchenko and Elmes
2009a). We placed it to this group because of the transverse rugosity on the alitrunk dor­
sum (combined with a coarsely reticulate pronotum and longitudinally rugose propodeal
dorsum); other important characters are the hairless lateral margins of the head above eyes
(only genae and occipital margin with sparse, quite long suberect to subdecumbent hairs)
and reticulation on the head dorsum posterior to the eyes, combined with longitudinal ru­
gosity on the rest of the head dorsum. Queens and males unknown.
Ecology is unknown except that this species was found in subalpine meadows at alti­
tudes 4000-4700 m a.s.l. Together with M. kozlovi, M. polyglypta seem to be one of the
most high altitude tolerant of the Myrmica species.

220
 Map 88. Distribution of M. polyglypta.

Myrmica pulchella Santschi, 1937, stat. rev., stat. nov.

(Figs 202, 203)

Myrmica margaritae var. pulchella Santschi, 1937: 368, w, Taiwan (first available replacement name
for Myrmica margaritae var. formosae Wheeler W. M., 1929: 37); Weber 1950: 219; Chapman and
Capco 1951: 127; Radchenko 1994b: 44; Bolton 1995: 282.
Myrmica margaritae var. formosae Wheeler W. M., 1929: 37, w, Taiwan {Myrmica margaritae var. for-
mosae Wheeler W. M., 1928a: 9, nomen nudum) (junior primary homonym of Myrmica formosa
Smith F., 1858: 128, transferred io Podomyrma Smith F.); Chapman and Capco 1951: 127.
Myrmica ritae subsp. formosae: Weber, 1950: 220.
Myrmica formosae: Radchenko 1994b: 44; Bolton 1995: 279; Radchenko and Elmes 1998: 7; Elmes and
Radchenko 1998: 222; Radchenko, Zhou and Elmes 2001: 217; Radchenko and Elmes, 2009c: 5.
Junior synonym of M. formosae: Radchenko and Elmes 1998: 7.

Type localities and type specimens. M. margaritae var. pulchella: "Formose: Musha, (K. Sato),
4 w (n° 726)". M. margaritae var. formosae: "Numerous specimens from Funkikó, Formosa".
Material examined. Lectotype of M. pulchella (designated by Radchenko and Elmes 1998), w
(upper specimen on the pin with 3 w), "Musha, Formosa, K. Sato" (BASLE); paralectotype: w (bot-
tom specimen on the pin with lectotype; see also Notes below) (BASLE); lectotype of M. formosae
(designated by Radchenko and Elmes 1998), w, "Funkika, Formosa, Silvestri" (HARVARD); paralec-
totypes: 9 w with same label (HARVARD, MOSCOW); non-type material: 34 w, 2 q from Taiwan.
Distribution (Map 89). Taiwan.
Etymology. M. pulchella: from the Latin word pulchella = pretty, perhaps to describe
the nice appearance of this species but more probably Santschi just translated "Formosa"
to Latin. M. formosae: from the old Portuguese name Formosa = Beautiful [Island] for the
Island of Taiwan, the type locality.
Notes. M. pulchella belongs to the ritae-complex of the ritae species group. It has ex-
tremely coarse rugosity on the head dorsum (only four sinuous rugae between the frontal
carinae level with the eyes), a feature shared with M. margaritae, M. sinensis and M. emeryi.
It well differs from M. margaritae by the presence of numerous standing hairs on the head

221
margins and alitrunk dorsum and from M. emeryi - by the more abundant standing
pilosity on the head margins (> 20 long hairs vs. < 10). The absence of reticulation on the
head dorsum distinguishes M. pulchella from both M. emeryi and M. sinensis.
Based on the direct comparison of the types of both taxa, we considered (Radchenko and
Elmes 1998; Elmes and Radchenko 1998) M. margaritae var. pulchella as a junior synonym
of M. formosae. However, the latter name is a junior primary homonym of M. formosa Smith
F., 1858, and the first available replacement name for this species is M. pulchella.
We designated as the lectotype the upper specimen of 3 type workers of M. margaritae
var. pulchella, stuck on the same pin. There is no doubt that the bottom worker is the same
species, but the middle worker is distinctly smaller and has a finer sculpture on its head
(more than six rugae between frontal carinae level with the eyes). We speculated (Rad-
chenko and Elmes 1998) that either M. formosae could be a rather polymorphic species or
that the small specimen is probably M. serica, collected in error with the M. pulchella. Now,
after studying a large amount newly collected material of both M. pulchella and M. serica,
we believe that the middle specimen mentioned above is indeed M. serica. We therefore
propose to exclude it from the type series of M. pulchella.
Below we describe queens (based on 2 queens collected by G. Elmes in Taiwan, Chiayi
Hsien, Fenchihu, Sun Moon lake, 11.viii.2002).
First description of queens: they are distinctly larger than the workers and as in most
Myrmica species there is some allometry in several body proportions. They have the same
coarse sculpture of the body seen on the workers, the main difference is the shape of the
propodeal spines that distinctly curve downwards (seen in profile) rather than straight.
Males are unknown.
Ecology. We know a little of the ecology of M. pulchella from our own experience of
collecting on the Island of Taiwan. We found three nests on a bank by the side of a small
road near Sun-moon lake, Chiayi Hsien, adjacent to a bamboo forest (quite large variety
4-5 m tall) at about 1600 m. a.s.l. The ants were found living in an intermodal cavity in

Map 89. Distribution of M. pulchella.

222
semi-decayed pieces of cut bamboo (1-2 cm diameter), each contained > 100 workers and
two contained a single queen. This was the first time we have seen any Myrmica species
living in unstructured cavities in the field, although most species will happily colonise such
spaces in laboratory conditions, provided the space can maintain a high relative humidity
(at least 90%). Some North American species can live in hollow acorns, and in boreal con-
ditions some species (e.g. M. lobicornis) build small nests in rotten twigs, though in this case
they usually hollow out the nest chambers themselves. We also observed a few foraging
workers (mid morning) nearby that appeared to emerge from holes in the soil, but we could
find no nest. Based on a sample of three nests one can not say more than small colonies of
this species adopt a cavity-living life-style. We have no way of knowing whether the colony
size is typical for this species or whether they were relatively young colonies. Perhaps if
they had grown larger they might have migrated to larger cavities in large pieces of bamboo.
We did not search far into the bamboo forest because it was too dense to penetrate.

Myrmica ravasinii Finzi, 1923

(Figs 204-206)

Myrmica ravasinii Finzi, 1923: 2, w, Albania; Müller G. 1923: 39; Finzi 1926: 112, Weber 1950: 212;
Zhizhilashvili 1967: 53, 1974: 193; Arnoldi 1970: 1843; Arnoldi, Dlussky 1978: 535; Agosti and
Collingwood 1987a: 53, 1987b: 268; Seifert 1988: 37; Arakelyan 1989: 733, q, m; 1994: 25;
Radchenko 1994b: 44, 1994c: 140, 1994e: 79; Bolton 1995: 282; Seifert 2003: 150; Radchenko,
Elmes and Alicata 2006: 519.

Type localities and type specimens. "Diverse w del Tomor (a circa 1600 m.) da nido sotto un sasso".
Material examined. Syntypes: 1 w, "Tomor", "Albania 1922, Ravasini, Lona", "4011 Coll. Karawa-
jewi", "M. ravasinii F. cotypc dct. Finzi" (KIEV); 2 w, "Tomor", "Albania 1922, Ravasini, Lona",
"M. ravasinii F. cotype det. Finzi" (MOSCOW), 1 w with the same labels (WARSAW); 2 w, "Tomor,
Albania, 7.22", "M. ravasinii cotipi" (GENOA); non-type material: > lOOw, 7 q, 11 m from Georgia,
Armenia and Turkey.
Distribution (Map 90). Albania, former Yugoslavian countries, Greece, Georgia,
Armenia and Turkey.
Etymology. M. ravasinii: named for Prof. Carlo Ravasini (1874-1959) of Trieste, a doc-
tor (urologist) and amateur coleopterologist who made the 1923 Ravasini-Lona expedition
to Albania; they made their ant collection available to Finzi.
Notes. M. ravasinii belongs to the schencki species group and is one of the most pecu-
liar Myrmica species outside of the Himalaya, China and SE Asia. Its has the narrowest
frons among all known Myrmica (mean FI 0.17) and an extraordinarily enlarged lobe on the
scape bend that well distinguish it from any other known Myrmica worldwide. At the same
time, it appears to have a quite stable morphology. Its males well differ from those of the
other species of the scheitcki-group (except M. caucasicola) by the presence of long erect
hairs on the scape, tibiae and tarsi.
Ecology. M. ravasinii is a semi-xerophilous species. The regions where it lives are very
hot in summer therefore it is a mountain species usually living between 800 and 1800 m
a.s.l. Generally, it lives in fairly open forests (oak, fir, pine), where presumably it benefits
from warmth in spring but shade in summer, though occasionally it has been found
on mountains meadows. In very dry mountains it is often associated with the edges of small

223
 Map 90. Distribution of M. ravasinii.

gullies where the sub-soil may be a little moist. Nests are built in the soil and in more shaded
places it may live under stones. In southwest Turkey we found sexuais in nests in mid Au­
gust, and Arakelyan (1989) found them in Armenia in a nest on 30th of August, so nuptial
flights probably occur August-September.

Myrmica rhytida Radchenko et Elmes, 1999

(Figs 207-209)

Myrmica rhytida Radchenko et Elmes, 1999a: 41, w, q, m, India; 2001b: 261.

Type localities and type specimens. India, for details see material examined.
Material examined. Holotype, w, "Kashmir, Up. Kainthal Nar, 34°00'N, 70°45'E, 3750 m, No. 3061,
14.08.1978, leg. P. Ward" (LONDON); paratypes: 113 w, 32 m from the same nest; 3 q, 4 m, "Kash­
mir, 3 km NE Таг Sar, 34°09'N, 75°11'E, 3300 m, No. 3037, 02.08.1978, leg. P. Ward"; about 200 w,
10 q, 5 m, "Kashmir, Таг Sar, 34°09'N, 75°09'E, 3950 m, No. 3038, 3039, 3040, 3041, 3042, 04.08.1978,
leg. P. Ward"; 25 w, "Kashmir, 1 km NE Yehmer Pass, 34°13'N, 75°10'E, 3600 m, No. 3043, 3045,
06.08.1978, leg. P. Ward"; 84 w, 7 q, 4 m, "Kashmir, 4 km S Kulan, 34°14'N, 75°10'E, 3599 m,
No. 3046, 3048, 07.08.1978; 3 workers, leg. P. Ward"; "Kashmir, Sain Nar, 34°06'N, 75°34'E, 3750 m,
No. 3054, 11.08.1978, leg. P. Ward"; 4 w, 1 q, 4 m, "Kashmir, Wampet, 34°04'N, 75°37'E, 3700 m,
No. 3057, 12.08.1978, leg. P. Ward"; 1 m, "Kashmir, Up. Kaintal Nar, Oitto, 34°00'N, 75°45'E, 4200
m, No. 3068, leg. P. Ward"; 3 w, "India, Beastel, 20 km S Rhotang, 2800 m, No. J-23, 18.08.1990, leg.
J. Heinze" (LONDON, BASLE, MILAN, SCHULZ, WARD, KIEV, ELMES, MEI).
Distribution (Map 91). It is known only from the type localities from India (Kashmir).
Etymology. M. rhytida: from the Greek word parie (rhytìs) = wrinkle (ruga) to reflect
the coarse rugosity of most of the body.
Notes. The saddle-shape alitrunk of the workers of M. rhytida suggest that it might be
close to species from the cachmiriensis-comp\ex of the rugosa species group. Both workers
and queens differ from the species of the cachmiriensis-comv\ex (M. wardi, M. ordinaria

224
 Map 91. Distribution of M. rhytida.

and M. cachmiriensis) by the distinctly coarser rugosity of head dorsum, alitrunk, petiole
and postpetiole. However, rugosa-group species are typified by males having short anten-
nal scape whereas the scape of M. rhytida males is long. Probably it could be placed in "its
own" separate species group, but for now we list it under, species whose taxonomic position
is uncertain.
Ecology. A little is known about its ecology. It is fairly common at altitudes from 3130
and 4200 m in Kashmir (Philip Ward, pers, comm.), where it nested in the soil (often under
stones) and in birch logs, in alpine meadows often containing juniper species, and at the
edges of birch forests at an altitude where rhododendrons begin to appear. It was often
sympatric with M. wardi but tended to live at higher altitudes (mean 3600 m vs. 2700 m
a.s.l.) Ward found it tending root aphids in several of its nests. Its foraging behaviour
appears to be quite cryptic, it stays close to the soil surface and was never found
foraging on plants. In this respect it behaviour is reminiscent of that of the northern
European M. lobicornis.

Myrmica rigatoi R a d c h e n k o et Elmes, 1998

(Fig. 210)

Myrmica rigatoi Radchenko et Elmes, 1998: 18, w, Pakistan; 2001b: 256.

Type localities and type specimens. Pakistan, for details see material examined.
Material examined. Holotype, w, "Pakistan, Changla Gali (between Marree and Nathia Gali), 2200
m, 16.08.1994, leg. S. Dakatra" (MILAN); non-type material: 1 w, Pakistan, Hazara Durgo Gal., 2300
m, 17.iv.1984, leg. Zoja (STOCKHOLM).
Distribution (Map 92). It is known only from Pakistan.
Etymology. M. rigatoi: this species was dedicated to our friend and colleague Dr. Fa-
brizio Rigato of Museo Civico di Storia Naturale, Milan.

225
 Map 92. Distribution of M. rigatoi.

Notes. When we first started working on the oriental Myrmica we thought that M. rigatoi
most resembled species of the n'tee-group, so we placed it there (Radchenko and Elmes
1998). Later, as we refined our concept of the ritae-group we realised that M. rigatoi well
differs from the other species of n'toe-group by having an anterior clypeal margin that is not
notched and by the propodeal lobes that are more rounded rather than pointed, so we re­
moved it to the шегде-group mainly because of the similarity of the shape of petiole and
propodeal spines (Radchenko and Elmes 2001b). On the other hand, it clearly differs from
M. inezae by the absence of transversal sculpture on the alitrunk. Therefore we still consider
its placement in the inezae-group as provisional.
Queens, males and ecology are unknown.

Myrmica ritae Emery, 1889

(Figs 211, 212)

Myrmica ritae Emery, 1889: 501, w, Burma; Dalla Torre 1893: 113; Forel 1903: 696; Bingham 1903:
267; Emery 1921: 38; Weber 1950: 219; Chapman and Capco 1951: 128; Radchenko 1994b: 44;
Bolton 1995: 282; Radchenko and Elmes 1998: 5; 1999b: 75, q; Radchenko et al. 2008: 776, nec
Collingwood 1970: 394, misidentification.

Type localities and type specimens. "Tenasserim: Mooleyit, 1000-1900 m."

Material examined. Lectotype (designated by Radchenko and Elmes 1998), w, "Tenasserim,
M. Mooleyit, 1000-1300 m, Fea, Apr. 1887", "TYPUS", "Myrmica Ritae Em., n. sp.", "Museo Civico
di Genova", "Lectotypus design. Radchenko et Elmes, 1998" (GENOA); paralectotypes: 3 w with the
same labels; 5 w, "Tenasserim, M. Mooleyit, 1800-1900 m, Fea, Marzo 1887", "TYPUS", "Myrmica
Ritae Eni." (GENOA); 5 w, "Tenasserim, M. Mooleyit, 1800-1900 m, Fea, Marzo 1887", "Birmanie",
"Emery", "Museum Paris Collection Ernst André 1914" (PARIS); non-type material: 3 w, 1 q, "Nord-
tailand, Prov. Chiang Mai, Doi Inthanon, N. P. Hipfel Region, 1800-2000 mH, 1.1.1995, leg. Schulz
and Vock"; 14 w, "Doi Inthanon N. P., 2200 m, near Chiang Mai, N. Thailand, 20.viii.1998, leg.

226
S. Yamane TH-98 SKY-22"; 1 w, "N. Thailand, 16.vi.1981, W. L. Brown and 1. Burikar, Doi Indhan-
nont, 1780 m, mountain humid forest, rotten wood"; 1 w, "N. Thailand, Doi Indhannont, 1760 m,
mountain humid forest, rotten wood, 16.iv.81, W. L. Brown" (SCHULZ, MILAN, LONDON, ELMES,
KIEV).
Distribution (Map 93). Northern Burma and northern Thailand.
Etymology M. ritae: although he does not say so, Emery probably named this species for
his wife Rita; we know her name because Forel (1899: 17), when he describe Leptogenys
ritae (collected by Emery), wrote "Je dédie cette espé à Mme. Rita Emery. Se distinguée par
sa forme svelte et grêle".
Notes. This species belongs to the n'toe-complex of the ritae species group. M. ritae
clearly differs from all other related species by the almost straight (not sinuous) longitudi-
nal rugae and complete absence of reticulation on the head dorsum, and by its yellowish
head, which distinctly contrasts with the brown alitrunk.
Collingwood's (1970) record of M. ritae for Nepal almost certainly is a misidentification;
probably Collingwood was using the Emery's very old concept of M. ritae, which more or
less embraced most n toe-group ants because he gave range of M. ritae as "Burma, Borneo",
the Borneo record should be referred to M. emeryi (see notes to M. emeryi above). Males
are unknown.
Ecology. Andreas Schultz (pers, comm.) found a queen and workers of M. ritae in Thai-
land in a piece of wood (15 cm diameter) on the ground, in dense old oak forest at an alti-
tude of 1950 m a.s.l. The forest was cool (annual mean temperature < 15°C) and quite
humid, with very few epiphytes, but with abundant moss. This fits the idea that n'toe-group
are mainly forest ants which probably forage in low shrubs, perhaps even in the foliage of
trees. Other genera collected in this area included Tetramorium, Crematogaster Lund and
Pachycondyla F. Smith.
We also found two workers of M. ritae in LONDON, collected at virtually same place
as the Schulz material (northern Thailand), in rotten wood in a mountain humid forest, at
an altitude 1780 m.

Map 93. Distribution of M. ritae.

227
 Myrmica rubra (Linnaeus, 1758)
(Figs 213-215)

Formica rubra Linnaeus, 1758: 580, w, Europe; Latreille 1802: 248, q, m.

Myrmica rubra: Latreille 1804: 179; synonymy before 1893 see Dalla Torre 1893: 113; Emery 1908a:
169; Kuznetsov-ugamsky 1927: 193; Karawajew 1927b: 258; Wheeler G. С and Wheeler J. 1953:
118, larvae; Yarrow 1955: 114; Collingwood 1958b: 67; Haustecheck 1965: 325; Arnoldi 1970:
1839, 1976: 554; Dmitricnko, Petrenko 1976: 13; Tarbinsky 1976: 40; van Boven 1977: 115;
Arnoldi, Dlussky 1978: 530; Collingwood 1979: 52; Espadaler 1981b: 189; Pearson 1981: 75;
Agosti and Collingwood 1987a: 53, 1987b: 267; Seifert 1988: 5; Atanassov, Dlussky 1992: 83; Rad­
chenko 1994b: 41, 1994c: 137, 1994e: 72, 1994g: 105; Seifert 1994: 14; Bolton 1995: 282; Seifert
1996: 230; Radchenko et al. 1997: 483; Czechowski et al. 2002: 17; Radchenko, Czechowska and
Czechowski 2004: 45; Radchenko 2005: 142; Seifert 2007: 2003; Pfeiffer et al. 2007: 5; Czechowski,
Radchenko and Czechowska 2007: 326, nec Curtis 1854: 213; Santschi 1931b: 339; Karawajew
1934: 65; Weber 1947: 448; Brian M. V. and Brian A. D. 1949: 393; Sadil 1952: 240; Onoyama
1989: 131; Imai et al. 2003: 182, misinterpretations or misidentifications.
Myrmica laevinodis Nylander, 1846a: 927, w, q, m, Finland; 1846b: 1052; Mayr 1855: 402, 1861: 64,
1863: 433; Nylander 1856: 78; Smith F. 1858: 116; Meinert 1861: 51; Mayr 1877: 14; Saunders
1880: 215; André 1883: 316; Nassonov 1889: 71; Dalla Torre 1893: 110; Ruzsky 1895: 29, 1896:
73, 1905: 662, 769; Wheeler W. M. 1908a: 337; Bondroit 1912: 351; Donisthorpe 1915: 110; Bon-
droit 1918: 104; Müller G. 1923: 40; Finzi 1926: 83; Santschi 1931b: 336; Ruzsky 1936: 94;
Karawajew 1934: 68; Stitz 1939: 78; Holgersen 1940: 184; Novak, Sadil 1941: 76; Weber 1947:
452; Sadil 1952: 239; Bernard 1967: 119; Baroni Urbani 1971: 2; Pisarski 1975: 9; Kutter 1977: 43;
Radchenko 2007a: 30, nec Curtis 1854: 213. Synonymy by Yarrow 1955: 114 (see Notes below).
Myrmica rubra subsp. (г., var.) laevinodis: Forel 1874: 76; Emery and Forel 1879: 460; Emery 1908a:
170 (misspelled as levinodis); Forel 1915: 28; Emery 1916: 121; Menozzi 1918: 82; Soudek 1922:
41; Karawajew 1927b: 259, nec Wheeler W. M. 1906b: 315;Wheeler G. С and Wheeler J. 1953: 119.
Myrmica longiscapus Curtis, 1854: 213, w, q, m, England. Synonymy by Mayr 1863: 433 (as synonym
of M. laevinodis); Yarrow 1955: 114 (as synonym of M. rubra); the latter confirmed here.
Myrmica laevinodis var. europea Finzi, 1926: 84 (first available use of Myrmica rubra subsp. cham­
plaini var. europea Forel, 191 lb: 457, w, Norway); Santschi 1931b: 339, m; Stitz 1939: 83; Bolton
1995: 279. Synonymy by Radchenko et al. 1997: 483; Czechowski et al. 2002: 17; confirmed here.
Myrmica laevinodis var. bruesi Weber, 1947: 453 (first available use of Myrmica rubra subsp. laevi­
nodis var. bruesi Wheeler W. M., 1906a: 38, w, q, m, U.S.A). Synonymy by Creighton 1950: 104
(as synonym of M. laevinodis); synonym of M. rubra: syn. nov.
Myrmica microrubra Seifert, 1993: 10, q, m, Germany; 1994: 14, 1996: 230; Kvamme 1999: 19;
Czechowski et al. 1999: 123, 2002: 19; Radchenko and Elmes 2003a: 236; Radchenko, Czechowska
and Czechowski 2004: 45. Synonymy by Steiner et al. 2005: 1; confirmed here.

Type localities and type specimens. M. rubra: no locality given. M. laevinodis: Finland (no exact
locality given): "Hab. in terris borealibus sub lapidibus, muscis, caesitibus, in arena cet. vulgaris".
M. longiscapus: "1 possess four males, two females and four neuters, found in July in Perthshire, and
I have received males and females taken out of one nest in the neighbourhood of Manchester by Mr.
R. Wood". M. laevinodis var. europea: "Bredheim, Nordfjord, Norvège (Prell)". M. rubra laevinodis var.
bruesi: "A number of workers, females and males taken by Mr. С. Т. Brues and myself during 1900 and
1902 from a few large colonies nesting under stones at the edge of Fay's Woods, Woods Hole, Mass.".
M. microrubra: "The holotype (a gyne) and 5 paratypes (3 gynes and 2 males) were fixed in a nest se­
ries from a peat bog in East Germany labelled "Oberlausitz, 6 km SE Weisswasser, 13.8.1981, Grosse
Jeserit-zen". Additional 8 paratypes (4 males and 4 gynes) were fixed in a nest series from north of
Berlin, labelled "Kr. Oranienburg, Glienicke, 14.8.89". All types are deposited in the collection of the
Staatliches Museum für Naturkunde Görlitz. The material studied included a total of 23 gynes and

228
23 males. 18 of these gynes and 13 of these males originated from 12 sites in Germany. G. W. Elmes
kindly provided 5 gynes and 10 males from his study site in Dorset/England".
Material examined. Lectotype of M. laevinodis (designated by Radchenko 2007a), w (upper
specimen on the pin with 2 w), "H: fors", "W. Nyland.", "Mus. Fcnn..", "Mus. Zool. H: fors Spec. typ.
No. 5044 Myrmica laevinodis Nyl." (HELSINKI); paralectotypes: 1 w (bottom) on the pin with
lectotype; 1 q, with the same label as lectotype, but type No. 5041; 1 m, 1 w, "H: fors", "W. Nyland.",
"Coll. Nyland.", "9/viii.45", "Mus. Zool. H: fors Spec. typ. No. 5042 Myrmica laevinodis Nyl.", "Mus.
Heis. N:o 3212"; 2 m (one without waist and gaster), "H: fors", "W. Nyland.", "Mus. Zool. H: fors
Spec. typ. No. 5043 Myrmica laevinodis Nyl."; syntypes of Ai. longiscapus, 2 w, "longiscapus Curt."
(Melbourne); non-type material: about 1000 w, several hundred of q and m from the whole range.
Distribution (Map 94). Europe (in the south mostly in mountains), Siberia, to the east
until Transbaikalia, to the north until Forest-Tundra Zone; in Transcaucasus and Middle
Asian mountains is quite rare; occasionally introduced to USA.
Etymology. M. rubra: from the Latin word rubra = red, to describe its generally reddish
colour. M. laevinodis: from a combination of the Latin words lavis (or Icevis) = smooth or
polished and nodus = knot or lump, to describe the relatively smooth surfaces of the peti­
ole and postpetiole compared to M. ruginodis. M. longiscapus: from a combination of the
Latin words longus = long (like a lance) and scapus = stalk or shaft, to describe the rela­
tive long scape (especially of males) compared to some other Myrmica species. M. europea:
named to indicate that a form of a North American M. champlaini (at that time believed to
be a variety of M. rubra, see Notes below) had a European form. M. bruesi: named for the
collector Prof. Charles Thomas Brues (1879-1955), who was recruited to the entomology
department of Harvard University and succeeded W. M. Wheeler as its head. M. microrubra:
combination of the Greek word ршрос, (mikros) = small and M. rubra to describe the small
form of queens (microgynes) of this species.
Notes. Some early authors considered the names M. rubra, M. laevinodis and M. rugi­
nodis all as separate species (e. g. Emery 1908a) though later he treated the latter pair
as subspecies of M. rubra (e. g. Emery 1921). In many cases, the name M. rubra was totally

Map 94. Distribution of M. rubra.

229
ignored and the Nylander's names were used, although some considered M. ruginodis to be
a subspecies of M. laevinodis (e. g. Ruzsky 1905). Over the years the changes in synonymy
of M. rubra has caused considerable muddle in the numerous non-taxonomic publications
on this species (see Chapter 1.4 for examples). Below we outline the history of this problem.
M. rubra was designated as the type-species of the genus Myrmica by Latreille (1810).
The original description of M. rubra was very brief and incomplete (Linnaeus 1758), but De
Geer (1771) gave a much fuller account of the biology of M. rubra (insofar as it was known
at that time) and he illustrated a male with a long scape (loc. cit., Fig. 12, Pl. 43), but made
no comment on this feature; most probably this was a specimen of M. ruginodis. When Ny-
lander (1846 a, b) described the common European Myrmica species he failed to assign the
name M. rubra to any of them and this was the source of all subsequent confusion. Curtis
(1854) noticed this omission and tried to rectify it: he gave a redescription of genus Myr-
mica (all three castes) based he said on M. rubra ("from which species all these generic
characters are taken ...", loc. cit., p. 212). He then described various species of English Myr-
mica (loc cit., p. 213-215) based he said "On comparing species oìMyrmica in my cabinet
with the Collection in the British Museum, which has been arranged by Mr. Smith [Fred
Smith] having had thereby the advantage of examining the typical specimens presented
by M. Nylander to our national museum." (we have not found any such specimens). Cur-
tis considered M. scabrinodis to be conspecific with M. rubra (loc. cit., p. 213): this is also
clear from his generic description males (loc. cit., p. 212) where he wrote "basal joint [of
antenna] generally one third of the entire length" and gave the footnote "the scape is
scarcely longer than the terminal joint in the male" also illustrated by Fig. 1 (loc. cit.).
Curtis did not say whether he had seen Linnaeus' material although it is almost certain that
either he or Smith would have examined the Linnaean collection. The synonymy of
M. scabrinodis with M. rubra was quite reasonable if Curtis based his opinion on Linnaeus'
material of "F. rubra", which is mostly M. scabrinodis (Stärcke 1951; Yarrow 1955), but it
was inconsistent with the illustration of a male made by De Geer (1771). Sadly, despite
having seen some of Nylander's material, Curtis appeared to have completely muddled Ny-
lander's (1846 a, b) treatment of the other species and this is why his treatment of the type
species for genus Myrmica was more or less ignored by subsequent authors. It is clear Cur-
tis' idea of "laevinodis" was some species from the scabrinodis- or schencki species groups,
whose males have a short scape, because under the description of M. longiscapus he wrote
"This species resembles M. laevinodis, but the males are much smaller, the antennae are
much longer, and instead of the scape being only as long as the two basal joints of the fla-
gellum, as in M. laevinodis and M. rubra (fig. 1), it is equal in length to the eight following
joints (fig. 12)" (loc. cit., p. 213). We investigated the syntype specimens (2 workers) of
M. longiscapus and have no doubt that they are M. rubra (sensu Yarrow). Curtis'
M. longiscapus, M. perelegans and M. denticornis appear to be Nylander's M. laevinodis,
M. sulci-nodis and M. lobicornis respectively; he also erroneously synonymised M. ruginodis
as M. vagans (a name incertae sedis in Formica).
Although the name M. rubra could theoretically refer to many of the later-described Myr-
mica species, by the first part of the 20th century most authors considered either M. ru-
ginodis or M. laevinodis to be the most likely candidates for synonymy because Linnaeus
emphasised that M. rubra stings freely and M. laevinodis and M. ruginodis usually are more
aggressive than the other European species. However Stärcke (1951) felt that the empha-
sis placed on stinging was misguided because the words 'pessimum nostrani pungit' were
added much later, in the 12th edition of "Systema Naturae" (Linnaeus 1767).

230
 The situation seemed to be resolved when Santschi (1931b) formally synonymised
M. ruginodis under the name M. rubra because he felt that M. ruginodis best fitted the de-
scription on other criteria. Santschi's opinion was increasingly followed by subsequent au-
thors (e. g. Karawajew 1934; Weber 1947; Sadil 1952; Brian and Brian 1949; Pisarski 1975,
etc.) while others like Stitz (1939) continued to use the Nylander name. Even as late as
1951 Stärcke wrote "the name M. rubra L. remains an uncleared hononym and it were best
to drop it altogether and maintain the well known names of Nylander. If its function as
a type-species-name of the genus prevents this, there should be chosen a neotype, found in
Upsala, and readily to be identified with either ruginodis or laevinodis" (Stärcke 1951: 324).
The quest for a type for Formica rubra Linnaeus, 1758 was finally settled by Yarrow
(1955), who was the first modern author to gain free access to Linnaeus' specimens. Ac-
cording to Yarrow, these were on 6 pins: 2 had single specimens of queen M. scabrinodis,
2 had both a queen and male of M. scabrinodis and another had a single male of M. laevi-
nodis. The last specimen was a single worker of M. laevinodis pinned through the gaster and
the photographs of this specimen show clearly that it is M. laevinodis (the photographs are
available on the Linnaean Society's web site [www.linnean-online.org/]). Supported by the
European-wide reputation of M. laevinodis for freely stinging, Yarrow felt this specimen
best fitted Linnaeus' description and labelled it "Formica rubra Linnaeus, 1758, lectotype,
Yarrow 1954" and "= M. laevinodis Nyl., det. Yarrow 1954". Most authors immediately ac-
cepted Yarrow's treatment (e. g. Brian), while others ignored it for a while. Perhaps the
strangest response was that of Bernard (1967) who no doubt influenced by the out of date
opinions of Stärcke (1951) refused to accept any of the earlier synonymies because
"L'identité exacte du type Formica rubra de Linné n'est pas, à notre avis, établie défini-
tivement. Yarrow (1955) a pu examiner les 8 exeplaires, plus ou moins bien conserves, de
la collection Linné, et pense, ... qu'il s'agit de M. laevinodis, mais il n'est pas très affirma-
tive" (Bernard 1967: 119). Clearly he had not read Yarrow's paper properly. However, this
did not stop him from giving the name "Groupe rubra" to species having males with long
antennal scape, including M. laevinodis and M. ruginodis because "... c'est le Myrmica
[M. rubra] le plus anciennement décrit (1758), on donner a son nom à ce groupe". Despite
such reactions almost every author now uses Yarrow's synonymy of M. laevinodis with
M. rubra which we accept as definitive.
Frequently in strong populations of M. rubra, miniature queens or microgynes can be
found in some nests almost always with "normal" queens (see Elmes 1973b, 1974b, 1976).
Elmes followed the traditional idea that the microgyne is a specialised morph (see for
example Wheeler 1910), he considered it had been "derived" from a true "ecological morph"
such as M. ruginodis microgyna (see notes to M. ruginodis) and had a "semi-parasitic" be-
haviour and it might be a "precursor" to a social parasite. Pearson (1981) supposed it to be
a social parasite and believed his genetical analysis based on isoenzymes, showed that it dif-
fered from the M. rubra host population, but he did not go so far as to give it a name. Pear-
son's work combined with morphometric analyses led Seifert (1993) to describe it as
a distinct socially-parasitic species - M. microrubra. Savolainen and Vepsäläinen (2003)
made a genetic comparison of M. microrubra with their host populations at several locations
and found that the "parasites" were always most closely related to their host populations
and discussed how their results might lend support for some sort of sympatric speciation
within populations. Most recently the wheel has gone full circle, Steiner et al. (2005) have
•synonymised M. microrubra with M. rubra believing it reflects queen dimorphism in M. rubra,
based on a series of morphological, biological, biochemical and molecular genetical studies.

231
 Not surprisingly there has also been confusion with the naming of non Eurasian forms
of M. rubra. Forel (1901b) described M. laevinodis var. champlaini from Canada and
Wheeler (1906a) described M. rubra laevinodis var. bruesi (first available use of this name
- Weber 1947) from the USA. Based on recent knowledge of the introduction of M. rubra
into parts of Canada and northwest USA, these ants could be either North American species
or the introduced M. rubra. In the past both forms have been considered as junior synonym
of M. rubra (e.g. see Bolton et al. 2006) but recently Groden et al. (2005) synonymised var.
champlaini with M. brevispinosa Wheeler; we have studied a syntype worker of var. cham-
plaini (GENOA) and confirm the proposed synonymy. Although we have not checked the
types of var. bruesi we are sure that it is a synonym of M. rubra. Firstly, Wheeler (1906a:
38) said that his specimens "agreed very closely with European laevinodis from Russia,
Austria, Germany, England and Scotland in my collection" being only slightly less sculp-
tured and therefore he "establish this variety with some hesitation". Secondly based on sur-
veys of populations of introduced M. rubra in north-eastern USA the species is known to
be present in parts of Massachusetts (Groden et. al. 2005).
This leaves Myrmica rubra subsp. champlaini var. europea described from Norway by
Forel (1911b) (first available use of this name-Finzi, 1926). We have not seen types of this
form, but based on Forel's description ("Epines dentiformes, comme chez le type de la
subsp. Pétiole antérieur du premier noeud plus distinct, mais plus court que chez la laevi-
nodis. Couleur d'une laevinodis pâle. Sculpture de la laevinodis") it is quite clear, that
Forel had trivial variants of M. rubra with somewhat shortened propodeal spines. There is
absolutely no evidence for the possible introduction of M. brevispinosa (M. champlaini)
into Norway and furthermore, from our understanding of Myrmica taxonomy and
faunistics the presence of an infraspecific form of M. rubra in Norway highly improbable.
Moreover, M. brevispinosa (and "champlaini") clearly differs from any of the Palaearctic
species of the rubra-group, therefore we confirm its synonymy with M. rubra.
M. rubra is superficially quite similar to M. ruginodis, and oriental populations of M. ru-
ginodis vary somewhat from western ones on which most keys were based (see Notes to
M. ruginodis). We are sure that M. rubra is not present in Japan and Far East of Asia, but
M. kotokui has behavioural and ecological similarities to M. rubra, which has led some
Japanese authors to erroneously determined some populations M. kotokui as M. rubra (see
Notes to M. kotokui above).
Wheeler G. С and Wheeler J. (1953) described larvae of M. rubra (under the name
M. rubra laevinodis) based on "numerous larvae from Massachusetts" (loc. cit., p. 119). It
is quite probable that these should be referred to Wheeler's var. bruesi but equally they
could have been M. brevispinosa or related species. In their later additional notes on larvae
of genus Myrmica (Wheeler G. and Wheeler J. 1972: 233) gave no further information on
M. rubra other than the surprising statement that they had not personally bothered to read
M. V. Brian's meticulous work on ontogeny and caste determination in M. rubra larva, but
instead recommended second-hand (and partly inaccurate) reviews.
Ecology. M. rubra is a eurytopic species distributed widely throughout Europe and West
Siberia where it can dominate some habitats. It thrives in damp habitats, especially soils
with high water tables or habitats in areas of high rainfall. However, it is seldom found
living in tussocks on true bogs, in the manner of some populations of M. scabrinodis and
M. ruginodis. In western Europe it is considered to be a species of damp meadows and is
rarely found in woods and forests, the largest populations usually occur on west-facing
slopes with heavy clayey (often limestone) soils, where it builds nests in the soil and under

232
flat stones. These sites often have high rainfall and the moisture is held in the heavy soils.
In eastern Europe (Russia, Ukraine etc.) it is considered to be more of a forest species in-
habiting many different kinds of forests (except those with light, dry soils), where it builds
nests in the soil under moss and in or under rotten wood. In central Europe M. rubra is
often very abundant in grass on forest, woodland and hedgerow edges, and in Germany,
Poland and France etc. it is particularly abundant in the longer vegetation at the edges of
water meadows used for haymaking and grazing. Throughout its entire range it is associated
with meadows bordering rivers and lakes. In recent years it has become important in nature
conservation as the primary ant host of the endangered butterfly species Phengaris nau-
sithous (Bergsträsser) (see papers in Settele et al. 2005).
M. rubra colonies do not need a long season of high temperatures to complete their life
cycles, in most habitats the ants do not become active until the end of April and are
entering a pre-hibernation state by late September (Elmes 1982). They have an active basal
physiology (compared to many other Myrmica species) that has adapted to local environ-
ments in different parts of its range (see Chapter 1.3.5 for some references). Habitat selec-
tion seems to be determined by a trade-off between sufficient insolation to complete their
life cycle and maintaining a high humidity within the soil nest (assuming other factors such
as food availability and nest site suitability being equal). Thus at sea level in the more
oceanic climates of western Europe, woodlands are too cold in summer while east-facing
meadows get too hot and dry whereas in the much more continental climates of eastern
Europe, the hot summers enable them to live in woodlands which dessicate less rapidly
than open meadows. Mountains of course, make their own local climates so that for
example, in the Carpathians populations favour more open meadow habitats at higher alti-
tudes that are ecologically very similar to the prime habitat in western Europe.
Generally, the microhabitat favoured by M. rubra colonies living by rivers and wet
meadows ranges from open grass (about 10-20 cm tall) in the north and west of Europe to
much longer grass and reeds (1-2 m tall) in southern and eastern habitats. Much less is
known about its distribution in West Siberia: M. rubra is one of the commonest ants in
various habitats of West and East Siberia (Reznikova 1983; Dmitrienko, Petrenko 1976)
and is particularly common in rivers meadows in north-eastern Kazakhstan (M. Woycie-
chowski, pers. comm.). The principal competitors of M. rubra are other Myrmica species but
in meadows it faces strong competition from Lasius niger (Czechowski 1985).
Morphologically M. rubra is comparatively stable over its very wide range (unlike, for
example, M. scabrinodis, which is probably undergoing current speciation in Europe - see
notes to that species), most of its local adaptation appears to have been physiological
and perhaps behavioural (see Chapter 1.3.5). Consequently, given its abundance, it would
be an ideal candidate to attempt phylogeographical history of its invasion of Europe using
modern molecular analytical techniques. Until this occurs we can only hypothesise
from whence M. rubra spread. It seems probable to us, that it survived the last ice age in
Middle Asia or maybe the Balkans or southeast Europe, on so-called "tundra-steppes", and
spread rapidly into Europe along the great river margins as the ice melted. Coinciding with
man's deforestation of Europe many new habitats were created in the oceanic part of
Europe, the forests of which would have been generally too cold for colonisation. Thus
in a sense M. rubra is pre-adapted to invade anthropogenic habitats (gardens, agro-
coenoses) especially in areas of high rainfall. This might help explain why it has been a very
successful invader of the eastern seaboard of USA and Canada (see Groden et al. 2005 and
Chapter 1.3).

233
 It is most useful to think of M. rubra in terms of "nests", colonies can consist of single
nests, or polycalic aggregations of several nests and even huge aggregations that can be con-
sidered as super-colonies (see Garnas et al. 2007). At a typical meadow site in Southern
England nests were on average 1.8 m apart (Elmes 1974a). Most studies to date show that
individual nests contain from a few tens of workers to up to eight thousand with a mean of
about 1,000 (Wardlaw and Elmes 1996); nests are usually polygynous with a mean of about
15 queens and nests hardly ever exceed a maximum density of 1 queen per 10 workers
(Elmes 1973a). Remarkably average nest size does not differ notably from these estimates
even in the super colonies of the USA (Garnas et al. 2007). In wild populations workers
"turn-over" quite rapidly and live on average less than 2 years (Brian 1972) and most queens
are equally short-lived (Elmes 1980; Seppä and Walin, 1996) .The density of queens in nests
varies between populations in space and time (Elmes and Pçtal 1990), a statistical analysis
of weather variables indicated that the nest population of both workers and queens in-
creased as the mean late summer temperatures deviated from about 16°C, which was more
or less the long-term average for the sites studied. It was suggested that the most probable
explanation was that the local population structure was adapted to the long term condi-
tions at a particular site and in years when conditions deviated from this, nests were less
likely to fragment (see below) and so on average, were somewhat larger. This fits with the
larger picture for local physiological adaptation by populations of Myrmica in different parts
of their range (Elmes et. al. 1999 and Chapter 1.3.5) and the observations in particular for
M. rubra (e.g. Brian 1973a; Kipyatkov 1979; Elmes and Wardlaw 1983a; Raybould and
Pearson 1992). Nests are usually built into the soil and the design of the excavated nest is
tailored to the conditions of the nest site (e.g. Evesham 1992). In coastal meadows M. rubra
are less tolerant of salted soils compared to M. scabrinodis but nests can survive periodic
inundation (Boomsma and De Vries 1980; Boomsma and Isaaks 1982). Young mated
queens are available for much of the active year and are often recruited into nests (Elmes
1982) so that local populations can vary considerable for individual relatedness and level
of effective polygyny (Seppä and Walin 1996). Consequently nest fission is a common way
of reproduction in M. rubra. Nests frequently fragment in spring and in some circumstances
become aggressive towards each other (Czechowski 1984). Often fragments recombine in
autumn or they may remain associated as a polycalic colony. In some circumstances
M. rubra populations may form huge polycalic colonies with hundreds of thousands or even
millions of workers, like in some North America populations and very occasionally in some
European populations (personal observation). In such populations it appears that there is
a high level of food availability, nests grow rapidly and fragment but do not need to move
far from the parent nest. This results in a very dense population of nests though individual
nest sizes remain on average about 1000 workers. The proclivity for fragmenting into small
colonies each containing several queens seems to be a useful adaptation to rapidly invad-
ing new habitat and at the same time incipient colonies are easily spread and introduced into
new habitats by man's activities (in plant pots, roots of transplanted shrubs and trees - see
Groden et al. 2005 and Garnas et. al. 2007).
Although workers in supercolonies appear to be very tolerant towards their neighbour-
ing colony members, they can be very aggressive towards other organisms, stinging people
and other animals freely (personal observations); however, laboratory studies suggest that
M. rubra workers do not have especially different aggressive responses compared to other
members of the genus (De Vroey and Pasteels 1978). M. rubra have a well equipped sting
apparatus (Billen 1986) and although some people react allergically to the venom, the sting

234
and venom does not seem markedly different from that of other free-stinging myrmicines
(e.g. Blum and Herman 1978). Most people (including one of us - AR) think that M. rubra
stings seem particularly painful compared to other Myrmica species. However in the
opinion of the other author (GWE) when individuals of other Myrmica are provoked into
stinging (usually in hot conditions) a single sting can be equally painfully as that of M. rubra,
perhaps even more so if the specimen is large: M. rubra having acquired its reputation be-
cause colonies are corporately aggressive and individuals sting rapidly and frequently even
when fairly cool.
M. rubra is a generalist scavenger and predator hunting various small invertebrates (e.g.
Pftal 1967), but also utilize honeydew and nectar (flowers and extrafloral nectaries, e.g.
Felton 1959), aphids and scale insects. They forage on trees and shrubs more frequently
than any other Myrmica species (except perhaps M. ruginodis) ; though in Europe arboreal
foraging is quite rare while in the supercolonies of Maine, USA very many M. rubra
workers can be seen foraging high into the canopy (personal observations). M. rubra
workers often forage in groups (e.g. Dlussky et al. 1978) and they lay and follow chemical
foraging trails (e.g. Cammaerts-Tricot and Verhaeghe 1974; Cammaerts-Tricot et al. 1977).
Single foragers (weighing about 2 mg) are able to exert pulling-forces of about 100 mg.
developing a mean power of about 5.8 ergs/s (Sudd 1965), a third to a quarter of the
strength and power exerted by Formica lugubris Zett. workers.
Winged sexuais (gynes and males) are produced in June and "mature" inside the nest
until July. The ontogeny of larval development and caste determination has been exten-
sively studied by M. V. Brian using M. rubra as a model species (see Chapter 1.3.4) and has
been found to be very complex; for example, the hormonal state of the queen influences
larval hormone production and ontogeny (Brian 1959, 1974), trophic conditions are in-
volved (Brian and Abbott 1977) as are the age and numbers of workers (Brian and Jones
1980), seasonality has an effect (Pearson and Raybould 1997) and even gut parasite load
might have some impact (Pearson and Raybould 1998). A model based on these interactions
show that gyne production might be periodic (Brian et al. 1981) and this may account for
the observation that the mean size of workers and queens in nests is positively correlated
with worker number and negatively correlated with queen number (Elmes 1974b).
Nuptial flights occur from late July and have been reported as late as October. Com-
pared to many other Myrmica species, M. rubra mating swarms can be quite large aggrega-
tions and they have frequently been reported flying quite long distances to join swarms on
church towers, high trees and mountain-tops (e.g. Hubbard and Nagell 1976; Woy-
ciechowski 1990b; personal observations). We have on occasion observed nests having re-
cruited a mixture of their own daughters and other young queens (all fertilised), but we are
not sure whether their daughters mated in or near to the nest prior to joining the parent
colony or flew to a distant swarm, mated and found their way home again. While the latter
seems improbable it is what happens in the case of honey bees.

Myrmica ruginodis Nylander, 1846

(Figs 216-218)

Myrmica ruginodis Nylander, 1846a: 929, w, q, m, Finland; 1846b: 1052; Foerster 1850a: 66; Mayr
1855: 407; Nylander 1856: 89; Smith F. 1858: 115; Mayr 1861: 63; Saunders 1880: 214; André
1883: 317; Nasonov 1889: 33; Forel 1892: 315; Dalla Torre 1893: 115; Bondroit 1912: 351;

235
 Donisthorpe 1915: 115; Bondroit 1918: 103, 1920a: 150; Santschi 1919: 244; Müller G. 1923: 41;
Finzi 1926: 85; Stitz 1939: 83; Novak and Sadil 1941: 76; Holgersen 1942: 8; Yarrow 1955: 114;
Collingwood 1958b: 68; Hauschteck 1965: 325; Bernard 1967: 120; Collingwood and Yarrow 1969:
56; Arnoldi 1970: 1839; Baroni Urbani 1971: 27; Collingwood 1976: 300; Dmitricnko, Pctrenko
1976: 13; Kutter 1977: 67; van Boven 1977: 115; Arnoldi and Dlussky 1978: 530; Mizutani and Ya-
mane 1978: 38, larvae; Collingwood 1979: 53; Dmitrienko 1979: 94; Espadaler 1981b: 189; Colling-
wood 1981: 26; Kupyanskaya 1985: 76; Agosti, Collingwood 1987a: 53, 1987b: 267; Seifert 1988:
6; Atanassov, Dlussky 1992: 86; Radchenko 1994b: 41, 1994c: 137, 1994e: 73, 1994g: 105; Seifert
1994: 14; Wu and Wang 1995: 92; Bolton 1995: 282; Seifert 1996: 232; Kim B.-J. 1996: 179; Rad-
chenko et al. 1997: 484; Wci С et al. 2001: 561; Czechowski et al. 2002: 19; Kim B.-J. 2003: 2; Rad­
chenko, Czechowska and Czechowski 2004: 45; Radchenko 2005: 144; Seifert 2007: 207;
Radchenko 2007a: 30; Pfeiffer et al. 2007: 5, nec Mayr 1877: 15.
Myrmica rubra subsp. (г., var.) ruginodis: Forel 1874: 76; Emery and Forel 1879: 460; Ruzsky 1904a:
288; Emery 1908a: 171; Bondroit 1910: 498; Forel 1915: 28; Emery 1916: 119; Menozzi 1918: 82;
Soudek 1922: 41, 1923: 110; Karawajew 1926b: 94, 1927b: 258.
Myrmica laevinodis subsp. (г., var.) ruginodis: Mayr 1886b: 450; Ruzsky 1895: 29, 1896: 73, 1902d:
29, 1905: 671; Emery 1921: 39.
Myrmica diluta Nylander, 1849:41, w, Russia; Smith F. 1858: 119. Synonymy by Mayr 1861: 63; Bol­
ton 1995: 278; Radchenko 2007a: 28; confirmed here.
Myrmica rubra r. ruginodolaevinodis Forel, 1874: 78, q, Switzerland (see Notes below); Stitz 1939: 84;
Weber 1947: 452; Sadil 1952:241. Synonymy by Bernard 1967: 120; van Boven 1977: 115; Bolton
1995: 282; confirmed here.
Myrmica rubra r. ruginodis var. ruginodolaevinodis: Forel 1915: 28, w, q, m (in Key; unavailable name).
Myrmica laevinodis var. ruginodolaevinodis: Dalla Torre 1893: 111; Donisthorpe 1915a: 123.
Myrmica ruginodolaevinodis: Stitz 1917: 347.
Myrmica ruginodis var. ruginodolaevinodis: Finzi 1926: 86
Myrmica ruginodis var. silvestrii Wheeler, 1928b: 100, w, Japan, syn. nov.
Myrmica rubra var. silvestrii: Weber 1947: 451.
Myrmica silvestrii: Collingwood 1981: 26 (see also Notes below).
Myrmica kurokii var. sontica Santschi, 1937: 367, w, Japan, syn. nov.
Myrmica kurokii subsp. sontica: Weber 1947: 470; Bolton 1995: 283.
Myrmica rubra subsp. yoshiokai Weber 1947: 451, w, Japan, syn. nov.
Myrmica yoshiokai: Collingwood 1981: 26, Bolton 1995: 284.
Myrmica rubra var. mutata Sadil, 1952: 242, w, Czech Republic. Synonymy by Seifert 1988: 6; Bolton
1995: 281; confirmed here.
Myrmica rubra var. macrogyna Brian M. V. and Brian A. D., 1949: 393, q, m, Great Britain. Synonymy
by Bolton 1995: 281; confirmed here.
Myrmica rubra var. microgyna Brian M. V. and Brian A. D., 1949: 397, q, m, Great Britain. Synonymy
by Bolton 1995: 281; confirmed here.
Myrmica rubra: Santschi 1931b: 339; Karawajew 1934: 65; Weber 1947: 448; Brian M. V. and Brian
A. D. 1949: 393; Sadil, 1952: 240; Pisarski, 1975: 10, old treatment of the species name.
Myrmica orientalis: Radchenko 1994e: 73; not confirmed, see Notes to M. kotokui, above.
Senior synonym oiMyrmica dimidaita Say, 1836: 293, q, U.S A: Weber 1947: 448; Bolton 1995: 278;
not confirmed, see Notes below.

Type localities and type specimens. M. ruginodis: no precise locality given: "Habitatio et mores
praecedentis. sed est fere adhuc vulgarior". M. diluta: "E Rossia meridionali D. v. Motschulski".
M. rubra r. ruginodolaevinodis: no locality given. M. kurokii var. sontica: "Japon: Yamakita, w (No
17)... Cho Teranishi coll.". M. ruginodis var. silvestrii: "Fifteen workers from Chuzenji Lake, near
Nikko (Hondo) (type-locality) and one somewhat smaller specimen from Nikko". M. rubra subsp.
yoshiokai: "Described from three workers from Gummakcn [Gumma Ken], Japan, 14.VII.1930

236
 (H. Yoshioka coll.) in Dr. Wheeler's collection". M. rubra var. mutata: "... one w... Kotelné Jamy,
Krkonose 15-6-1926 Bohemia". M. rubra var. macrogyna and M. rubra var. microgyna: "We have
found both varieties of rubra at Coigrain, Dumbartonshire and also in Glen Falloch, Pcrthsire, 25 miles
to the north. We have found microgyna at Ings, Westmorland, and at Great barrow, Cheshire".
Material examined. Lectotype of M. ruginodis (designated by Radchenko 2005), w (upper speci-
men on the pin with 2 w), "Kuusamo", "W. Nyland.", "Mus. Zool. H: fors Spec. typ. No 5045 Myrmica
ruginodis Nyl." (HELSINKI); paralectotypes: 1 w, bottom on the pin with lectotype; 1 m, with the
same labels as lectotype, but type No. 5046; 1 q, 1 m and 1 w on one pin, with the same labels as lec-
totype, but type No. 5046 and one additional label: "Mus. Hels. N:o 3211"; 1 q, with the same labels
as lectotype, but type No. 5046 and one additional label: "Mus. Hels. N.o 3212" (HELSINKI); syntypes
of M. diluta: 1 w, "Ross, mer.", "Motchoulsky", "Coll. Nyland.", "Russ. merid. Motch.", "M. diluta
Nyl.", "Mus. Zool. H: fors Spec. typ. No. 5104 Myrmica diluta Nyl."; 1 w, "Ross, mer.", "Motchoul-
sky", "Coli. Nyland.", "Mus. Zool. H: fors Spec. typ. No. 5103 Myrmica diluta Nyl." (HELSINKI); syn-
types of M. rubra r. ruginodolaevinodis, 3 m, "M. ruginodo-laevinodis S Fisibouh 20 Août"
(LAUSANNE); 3 q, "Typus", "M. ruginodo-laevinodis, w, Fisibach 20 Août", "M. ruginodo-laevi-
nodis"; 7 m, "Typus", "M. ruginodo-laevinodis, w, Fisibach 20 Août", "Coll. A. Forel" (GENEVA);
lectotype of M. kurokii var. sontica (designated here), w, "Japon: Yamakita" (BASLE); lectotype of
M. rubra subsp. yoshiokai (designated here), w, "Gummaken, Japan, July 14.30, H. Yoshioka", "N 8",
"Myrmica rubra subsp. Yoshiokai Weber Syntype from Wheeler collection", "MCZ syntype 20552",
"Lectotypus, design. Radchenko et Elmes, 1998" (HARVARD); non-type material: about 1000 w,
several hundred of q and m from the whole range.
Distribution (Map 95). Transpalaearctic species, distributed from Atlantic to Pacific
Oceans, including Japan; in the south - only in mountains, absent from Middle Asian
mountains. Introduced to North America.
Etymology. M. ruginodis: from a combination of the Latin words ruga = wrinkle and
nodus - knot or lump, to describe the rugose surfaces of the petiole and postpetiole. M. ru-
ginodolaevinodis: from a combination of the species names M. ruginodis and M. laevi-
nodis, to indicate the intermediate characters of this species. M diluta: from late Latin word
dilutus = dissolve, dilute to describe its somewhat reduced sculpture. M. silvestrii: named

Map 95. Distribution of M. ruginodis.

237
for the collector, the Italian Prof. Filippo Silvestri (1873-1949), who was a renowned
entomologist and avid insect collector in the early 20th century. He became Director of the
Laboratorio di Entomologia di Portici (near Naples). M. sontica: name from the Latin word
sontica = hurtful or noxious, there is no record as to why Santschi chose this name but
probably the collector (Dr. Cho Teranishi) told him they stung painfully (see notes below).
M. yoshiokai: dedicated to the collector, the Japanese myrmecologist M. H. Yoshioka who
collected many other species described by Wheeler. M. mutata: name derived from Latin
word mutans = changing, to indicate it is an unusual or mutated form. M. macrogyna: com­
bination of the Greek word ршкрос {macros) = large and yuva {gyne) = woman, to de­
scribe the normal (large) form of queens of this species. M. microgyna: combination of the
Greek word /икрос {mikros) = small and yuva {gyne) = woman to describe the small form
of queens (microgynes) of this species. M. dimidaita: name derived from the Latin word
dimidius - divided [into two halves], we do not know why Say chose this name, possibly
something to do with the wing venation.
Notes. M. ruginodis belongs to the rubra species group. For quite a long period in the
middle of the 20th century the name was synonymised with M. rubra before Yarrow (1955)
decided that really M. laevinodis was the true synonym of M. rubra (this is discussed in de­
tail in the Notes to M. rubra, above). M. ruginodis can be confused with M. rubra (= laevi­
nodis) in some parts of Europe, especially when solitary specimens are examined; however
Brian and Brian (1951) demonstrated that a clear discrimination can be made on the basis
of propodeal spine length. Despite a lengthy critique by Stärcke (1951) who appeared con-
vinced that the intermediate form (race ruginodolaevinodis) really existed, Brian and
Brian's observation has stood the test of time. The simplest way of discrimination between
the two species in the field is to examine the length and shape of the spines (seen in pro-
file) with a xlO hand lens.
There appears to be a cline in the morphology of M ruginodis across its Palaearctic dis-
tribution: in Europe individuals have relatively (to head width) "stockier" bodies with rela-
tively longer legs and propodeal spines, whereas in Japan the workers are relatively longer
and more slender with somewhat shorter spines and legs (Elmes and Clarke 1981). In that
paper it was shown that if M. ruginodis from only Japan and Spain were available, one might
well consider them as separate species. However, since 1981 we have obtained much more
material and have noted that M. ruginodis populations in, for example, the mountains of
Turkey and Caucasus more resemble the oriental form. In Japan and Korea some populations
of M. ruginodis are sometimes confused with an oriental species, M. kotokui, which itself is
sometimes confused with M. rubra and /or M. ruginodis var. microgyna (see below and notes
to M. kotokui). M. ruginodis most obviously differs from M. kotokui by its longer propodeal
spines, flattened and coarsely rugulose petiolar node. Mizutani and Yamane (1978) described
larvae of M. ruginodis from Japan, but there must be some doubt as to whether these
were M. kotokui. Mayr (1877) recorded M. ruginodis for Middle Asian mountains; most
probably this was a misidentification, for there have been no subsequent records.
We investigated types of both M. ruginodis and M. diluta and can not find any essential
differences between them and therefore confirm the long established synonymy. Weber
(1947) synonymised M. dimidiata with M. ruginodis, if this synonymy is correct as has been
the case until now (see Bolton 1995; Bolton et al. 2006), then the name dimidiata has pri-
ority over ruginodis and should be considered as the senior synonym. However, we reject
this treatment and consider that M. dimidiata Say, 1836 incertae sedis in the genus Myr-
mica. Firstly, while M. dimidiata almost certainly belongs to the genus Myrmica based on

238
the wing venation of the gyne described, it is absolutely impossible to recognize the species
from the other rather vague features mentioned "Body pale yellowish: thorax somewhat
tinged with piceous" (Say 1836: 293). Secondly, although no locality was given for this
specimen based on other species described in Say's paper it was possibly in the south-
eastern states (probably Indiana) where there are no valid records of introductions oirubra-
group species. Thirdly, a size of 5 mm (l/5 th of an inch) is too small for a M. ruginodis queen,
almost certainly it is a specimen of one of the native North American species that have quite
small queens.
Forel (1874) created the name Myrmica rubra r. ruginodolaevinodis, but made notes
only for queens, writing "Epines de longeuer intermédiaire etc." (Zoc. cit., p. 78). One must
infer that the castes of this form are intermediate between laevinodis and ruginodis as de-
scribed in his keys. The syntype series from Fisibach (on the border of northwest Switzer-
land) came from a colony that Forel observed moving its nest site (Forel 1886: 137); his
description of the process and the nest sites in moss is typical for M. ruginodis, especially
var. microgyna (see ecology below). Later, Forel (1915) provided a little more morpho-
logical data on all castes in a Key, otherwise information by other authors is also scarce
(e.g. see Finzi 1926; Stitz 1939). We examined a syntype series of this form (queens and
males), these also included some workers (formally the workers can not be considered as
types). Both queens and workers have less coarsely sculptured waists, but we have in our
collection similar M. ruginodis specimens that are quite common in southern mountain
regions (Asia Minor, Caucasus). The three queens are small, headwidths ranging from 1.00
to 1.04 mm, and fall well within the size-range recorded for microgyne M. ruginodis (see
Elmes 1991, and below).
We also studied type specimens of M. rubra subsp. yoshiokai, M. ruginodis var. silvestrii
and M. kurokii var. sontica, and although they show subtle variations in morphology we be-
lieve that all are within the range of variation for M. ruginodis (discussed above) and these
names can be considered as junior synonyms of M. ruginodis. The specimens, determined
by Collingwood (1981) as M. silvestrii are in fact M. kurokii (material is in BUDAPEST,
examined).
We have not seen the type of Myrmica rubra var. mutata, but Seifert (1988:6) noted
that "it is a teratological specimen that clearly can be recognized as ruginodis". Based on
Sadil's (1952) description and comments on this unique holotype worker, we fully accept
Seifert's opinion. Probably var. mutata represents a worker, infested by Mermis parasite
(e.g. see Czechowski, Radchenko and Czechowska 2007).
Finally we come to the question of Brian and Brian's (1949) var. macrogyna and var. mi-
crogyna. There is absolutely no question that M. ruginodis has a size dimorphism of queens
(Elmes 1991, fig. 9), each size being adapted to different ecological conditions. We have no
doubt that the syntype queens of r. ruginodolaevinodis are microgyne M. ruginodis (see
above) and should var. microgyna ever be shown to be a biologically valid taxon then the for-
mer name would have priority. However, we agree with Bolton's (1995) opinion that these
are not varieties in a taxonomical sense and accept his opinion that Brian and Brian's names
are synonyms of M. ruginodis and so far, this approach has been supported by modern ge-
netical phytogenies (Jansen et al. 2010; Vepsäläinen et al. 2009). Interestingly, Nylander's
paralectotype queen has a head width = 1.06 mm which is intermediate in size between the
two forms. While the forms have no taxonomic status they have separate, important and
unique (for Myrmica) ecological roles that have been misunderstood and confused by some
authors over the years. This is investigated more fully in the ecological notes below. ,

239
 Ecology. M. ruginodis is the species of Myrmica that is most adapted to cool tempera-
tures, at least of those species studied in detail so far, though see data for M. kamtschatica
(Berman et al. 2010). Both the workers and brood have an active basal physiology (Elmes
et. al. 1999; Nielsen et al. 1999) that, for example, enables it to complete its life cycle in the
same time as a M. sabuleti nest but living at a mean nest temperature that is 4-5°C lower
(Elmes and Wardlaw 1983a). As in the case of M. rubra (above and Chapter 1.3.5) local
populations appear to have adapted physiologically to local environments (Elmes et. al.
1998). Consequently, M. ruginodis is found in cooler habitats: all kinds of forests, scrub-
lands, alpine meadows, woodland clearings, moorlands and bogs, but generally it avoids both
very wet and dry open sites. It is widespread and abundant in these habitats over most of its
range, however M. ruginodis is much less tolerant than M. rubra to anthropogenic pressure
(grazing, mowing etc.). Like all Myrmica, latitude and altitude determines the species' basic
distribution: in the far north it lives at more or less sea level, while at southern latitudes it is
mainly a montane species. At any particular altitude and latitude the degree to which
sunlight can penetrate the canopy (be it forest, scrub or long grass) in any habitat will de-
termine the range of potential nest-site temperatures available to the ants (e.g. see Brian and
Brian 1951). Its adaptation to cooler nest-sites (above) enables it to live in cool northern
forests and so it is one of the commonest ant species in the Forest Zone of the Palaearctic.
In forests and woodlands M. ruginodis prefers to build its nests in (and under) rotten
wood, branches and even trunks of fallen trees. However, in managed forest its preferred
habits is rotten tree stumps (Brian and Brian 1951; Franch and Espadaler 1988) and often
one can find a nest in more or less every stump. In grasslands it prefers to build nests in the
surface of the soil in and among the roots of grass (particularly Molinea species - see Elmes
1978b). In wetter boggy areas nests are usually constructed in the tops of moss tussocks.
In spring the workers build a small solarium above their nest where the overwinter brood
is reared; the solaria are usually a mixture of soil particles and dead vegetation, pieces of
grass, moss etc. and can be quite warm in sunshine. Such nests are often temporary and the
ants migrate to a new site every few months, often one can find a series of nests belonging
to the same colony: first the active nest, another a meter or so away that is in the process
of construction and occupied by a few workers and brood, and an old nest a meter or so in
the opposite direction occupied by just a few workers and sometimes, an even older nest
a meter or so further away from that (personal observations). It was such a colony that
Forel observed in 1866 (see notes on var. ruginodolaevinodis above). In more open grass-
land where colonies build more permanent nests in the soil and under stones, the substrate
affects nest densities (Fedoseeva and Demchenko 1997).
Early studies on caste determination, worker behaviour and larval ontogeny were made
using M. ruginodis [Sic!] as the model species (Weir 1958c, 1959a; Brian 1951b) before
M. V. Brian and his co-workers switched to M. rubra. Also, M. ruginodis was one of the first
Myrmica species studied ecologically (see Pickles 1940 and e.g. Brian 1950) and more re-
cently interesting life history traits of some populations have resulted from molecular ge-
netical studies (see Seppä 1992, 1994; Seppä and Pamilo 1995). M. ruginodis workers are
among the species that are most prolific in producing worker-laid eggs (Wardlaw and Elmes
1998). Understanding its ecology has become important to nature conservation because
M. ruginodis one of the primary hosts for the endangered butterfly Phengaris alcon (see
papers in Settele et al. 2005). In particular, understanding the species and colony recogni-
tion odours and their mimicry by the lepidopteran caterpillars is important, the cuticular
surface chemicals of M. ruginodis are quite distinct from those of M. rubra (e.g. Elmes et.

240
al. 2002). Like the other Myrmica species, workers lave long been known to stridulate
(Swinton 1878).
The workers are generalist predators, hunting small invertebrates (e.g. Brian 1955b)
and forming trophobiosis with aphids, though this is generally less developed than is the
case for M. rubra. Some studies have been made of recruitment to food sources (Cammaerts
and Cammaerts 1980). In many habitats the eliasomes of various seeds play an important
role in the diet of M. ruginodis and they have important mutualistic interaction with the
plants (e.g. Kjellsson 1985; Mark and Olsen 1996; Gammans et al. 2006).
It is in the studies of behaviour and population ecology of M. ruginodis that the concept
of var. macrogyna and var. microgyna diverges from the taxonomic notion (see taxonomic
notes above). As the two forms have no status in taxonomy we refer to them as "normal"
and "microgyne" M. ruginodis respectively. Brian and Brian (1949, 1955a) showed that in
west Scotland the queens in polygynous colonies of M. ruginodis were visibly smaller (mi-
crogynes) than those in monogynous (normal) colonies (mean head width 1.02 ± 0.06 mm
vs. 1.13 ± 0.04 mm), intermediate sized queens (head width about 1.06 mm were quite
rare). They showed that apart from being more polygynous, microgyne colonies have
workers that are generally less aggressive and more tolerant of other workers and queens
compared to workers from normal colonies. Microgyne colonies recruit new queens and re-
produce by colony fission whereas normal queens are more likely to attempt to establish
new colonies by independent (or pleometrotic) colony foundation. Thus microgyne colonies
are adapted to spread rapidly into and monopolise habitats that are stable in the long term
(such as grassy moorland) and the original colonising gene pool might remain in such
a habitat for many hundreds of years. Normal queens are better at dispersing into new habi-
tats where they form relatively short-lived monogynous colonies, though such colonies
sometimes engage in secondary recruitment (Seppä et al. 1995) and may even recruit some
microgynes. Later we confirmed the size dimorphism shown by Brian and Brian and showed
that most populations have at least a small proportion of microgyne queens (Elmes 1978a);
also we made the interesting observation that the overall proportion of morphological
var-iation of workers, expressed at the population level, was about the same for both forms
but a much higher proportion of this (ca 60%) was expressed within individual microgyne
colonies compared to only 40% within normal colonies (Elmes and Clarke 1981). This was
compatible with a higher number of unrelated queens producing workers in the microgyne
colonies. We also showed that there were differences in the pattern of brood production
and control of castes between the two forms (Elmes and Wardlaw 1983a, b).
It is clear that the size and behavioural differences represent a true polymorphism in
M. ruginodis that enables the species to exploit and monopolise a much wider range of habi-
tats than it otherwise could. The fact that M. rubra (= laevinodis) also has a semi-parasitic,
microgyne form combined with the nomenclatural problems from 1935-1955 (see notes to
M. rubra) has confused many people. Synthesisers have often confounded the results for the
two species. We recommend that anyone interested in these problems should consult the
original literature and bear in mind the name changes. To date we have no understanding
of the underlying genetical mechanism behind the polymorphism, as far as we know the
two forms "breed true" and nobody has been able to produce a microgyne queen from
a normal mother or vice-versa.
As a general rule, monogynous (or with two or three queens) colonies usually have only
normal queens, whereas highly polygynous colonies often have a mixture of queens,
a few normal queens and many more microgynes. Normal queens in microgyne colonies

241
apparently have a more "microgyne-like" tolerant behaviour, which begs the question as to
whether normal queens have an ethological dimorphism with some having an intrinsic
microgyne-like behaviour and others having a "monogynous normal" behaviour as described
by Brian and Brian (1955). This question still remains to be resolved. Microgyne colonies
average about 6 queens (with > 20 queens being common), while normal colonies average
1 queen (with 4-5 queens being very unusual). Surprisingly, the average worker populations
of the two types of colony do not vary significantly, the best overall estimate of colony size
in Britain is about 500 workers (Elmes and Keller 1993; Wardlaw and Elmes 1996). On up-
land moorland (> 200 m a.s.l.) colonies are smaller (about 400 workers) than on lowland
southern heaths and lowland Scottish moors (700 and 1200 workers respectively), the dif-
ference between southern England and Scotland being statistically significant. There has
been considerable debate as to whether Japanese populations of M. ruginodis have a mi-
crogyne form (Mizutani 1981; Kasugai et al. 1983; Ichinose 1990; Kikuchi et al. 1999 and
see Chapter 1.3.6 for further discussion). We suggest that probably the microgyne form is
not present in the Far East and that M. kotokui populations occupy the ecological niche nor-
mally used by microgyne M. ruginodis in the west; this remains to be fully tested.
Nuptial flight occurs in August-September (in mountains as late as mid-October) and
over the years have been regularly reported in the literature (e.g. Beare 1913; O'Rourke 1940)
and swarms, comprising mostly of males, often have been taken in light traps (e.g. Elmes
and Webb 1985). Brian and Brian (1949, 1955) showed that microgynes are more likely to
mate in or near to the nest in local swarms and join existing colonies whereas normal queens
are more likely to fly to larger more distant nuptial swarms and attempt independent colony
foundation. However, in regions where the microgyne form is found, mixed swarms are com-
mon but there is no evidence to support the idea of assortative mating between the two forms
(Elmes 1991), although larger males are more likely to find a mate than smaller ones. Per-
haps only the size of the mother is important in determining the nature of her female offspring
and the parentage of the males is not important (a sort of maternally mitigated polymor-
phism). Similar ideas were discussed in relation to M. lonae (see above).
Finally, the question of the stinging-abilities of M. ruginodis: like most Myrmica species
the workers readily deploy their sting when defending their nest against intruders. Their
venom is quite potent (Jentsch 1969a, b) but the general perception is that M. ruginodis
workers are much less aggressive towards human disturbance than M. rubra and that when
provoked into stinging, the sting is less painful (see ecological notes for M. rubra). However,
in warm humid conditions on Hokkaido Island Japan we found a dense population of M. rugi-
nodis nesting on the stumps of felled trees in a spruce forest. These workers were very ag-
gressive attacking as freely and stinging as painfully as the worst M. rubra colonies we have
observed. Furthermore, in these conditions M. kotokui stung freely and painfully. Thus
willingness to sting and the amount of venom injected appears to be an interaction between
the level of basic behavioural aggressive responses (probably varying between species and
populations) and temperature; it would be interesting to test this under controlled conditions.

Myrmica rugosa Mayr, 1865

(Figs 219-221)

Myrmica rugosa Mayr, 1865: 19, w, India (?); Dalla Torre 1893: 116; Forel 1903: 697; Bingham 1903:
268: Ruzsky 1905: 658; Forel 1906: 87; Emery 1908a: 167, 1921: 40; Donisthorpe 1929: 445;

242
 Chapman and Capco 1951: 129; Radchenko 1994b: 42; Bolton 1995: 282; Radchenko and Elmes
2001b: 244, 2002: 41, q, m, nec Weber 1947: 462, misidentication.

Type localities and type specimens. "Himalaya".

Material examined. Lectotype (designated by Radchenko and Elmes 2001b), w, "Himalaya",
"M. rugosa Mayr" (BERLIN); paralectotypes: 1 w, "Himalaya", "M. rugosa sp. n. Mayr" (LONDON);
1 w, "Myrmica rugosa m. Himal."; 1 w, "Himalaya Mayr е. с ej." (GENOA); 2 w, "Гималаи", "Колл.
Майра" ["Himalaya", "Coll. Mayr"], "295" (MOSCOW); non-type material: about 200 w, 7 q, 16 m
from India, Nepal and Bhutan.
Distribution (Map 96). India, Nepal, Bhutan.
Etymology. M. rugosa: name derived from the Latin word rugosus = wrinkle or full of
wrinkles, to describe its sculpture.
Notes. M. rugosa is one of the first Myrmica species to be described after Nylander's
naming of the common European species, and the first from the Himalaya. Mayr's (1865)
description was quite brief, stating only that this species has reticulated sculpture only on
the occiput and lateral parts of the head dorsum, and petiole only densely punctated. Fur­
thermore, he gave no date or collector in his original description indicating only that the
types are in his collection and that the locality was "Himalaya" (most likely India or Nepal).
We consider that together with three other Himalayan species (M. aimonissabaudiae,
M. hecate and M. ruperstris) it belongs to the rugosa species group and differs from the first
two species by the much less developed reticulation on the head dorsum, and from the last
by the absence of coarse short sinuous rugosity on the petiole and postpetiole.
We found 1 worker in BERLIN with the labels "Himalaya" and "M. rugosa Mayr", writ­
ten by Mayr's own hand. As this specimen fully agrees with Mayr's description, we have
designated it as the lectotype. Five more workers, preserved in LONDON, GENOA and
MOSCOW are designated as paralectotypes (see above).
We located in HARVARD 3 workers, labelled as "Cotypes", "Myrmica rugosa Mayr",
"Himalaya (Smythies), Forel coll.", "M.C.Z. Cotype No. 556". Actually, they are not

Map 96. Distribution of M. rugosa.

243
M. rugosa, but correspond with Ford's (1902) description and locality data of var. debilior,
and we believe these specimens belong to the type series ofthat form. When Weber (1947)
redescribed M. rugosa, almost certainly he studied the "cotypes" from HARVARD men-
tioned above. He wrote (loc. cit., p. 462): "Surface of head coarsely sculptured: ... clypeus
and median dorsal area [frons in our terminology] coarsely rugose, sides rugoso-reticulate...
Dorsal surface of petiole deeply vermiculate, sides vermiculate-punctate. Post-petiole
coarsely vermiculate-punctate"; this contradicts Mayr' description (and lectotype speci-
men), but fits well Forel's description of var. rupestris and var. debilior (see also Radchenko
and Elmes 2001b, and Notes to M. rupestris, below).
Ecology is poorly known. M. rugosa has been found at altitudes between 1200 and 3400
m a.s.l. in semi-natural habitats, but it also inhabits cultivated areas (Philip Ward, pers.
comm.). Males and alate queens were collected from the end of May to the end of June,
which is extremely early in the year compared to Euro-Siberian Myrmica species.

Myrmica rugulosa Nylander, 1849

(Figs 222-224)

Myrmica rugulosa Nylander, 1849: 32, w, q, Finland; Mayr 1855: 405, m; Nylander 1856: 80; Smith
F. 1858: 115; Mayr 1861: 63; André 1883: 317; Dalla Torre 1893: 116; Ruzsky 1902d: 30; Nasonov
1889: 71; Bondroit 1918: 100, 1920a: 146; Müller G. 1923: 38; Lomnicki 1924: 817; Finzi 1926:
78; Karawajew 1926b: 95, 1927b: 259, 1929: 203, 1934: 70; Arnoldi 1934: 161; Holgersen 1940:
185; Stärcke 1942: 25; van Boven 1947: 175; Weber 1948: 305; Sadil 1952: 245; Bernard 1967: 115;
Arnoldi 1970: 1840; Baroni Urbani 1971: 28; Kutter 1977: 68; Arnoldi, Dlussky 1978: 532; Colling-
wood 1979: 54; Seifert 1988: 11; Atanassov, Dlussky 1992: 89; Radchenko 1994b: 42, 1994c: 142,
1994d: 75; Seifert 1994: 10; Bolton 1995: 282; Seifert 1996: 222; Radchenko et al. 1997: 486;
Czechowski et al. 2002: 23; Radchenko, Czechowska and Czechowski 2004: 47; Radchenko 2007a:
28; Seifert 2007: 194; Czechowski, Czechowska and Radchenko 2007: 142, nec Mayr 1877: 15.
Myrmica rubra r. (subsp., var.) rugulosa: Forel 1874: 77; Emery and Forel 1879: 40.
Myrmica scabrinodis subsp. (st., r.) rugulosa: Forel 1892: 315; Ruzsky 1895: 31, 1896: 73, 1905: 690;
Emery 1908a: 177; Forel 1915: 29; Emery 1916: 120, 1921: 41; Soudek 1922: 43; Karawajew 1927a:
285.
Myrmica clandestina Foerster, 1850a: 63, Germany; Schenck 1852: 85, q, m. Synonymy by Mayr 1855:
405; Dalla Torre 1893: 116; Karawajew 1934: 70; Radchenko 1994d: 75; Bolton 1995: 282; con-
firmed here.
Myrmica rugulosa var. sulcinodorugulosa Nasonov, 1889: 35, w, Latvia, Estonia and Russia; Emery
1921: 42; Finzi 1926: 88; Bolton 1995: 284. Synonymy by Radchenko 1994d: 75; confirmed here.
Myrmica sulcinodis var. sulcinodorugulosa: Weber 1948: 268.

Type localities and type specimens. M. rugulosa: "Parce ad Helsingfors sub lapidibus".
M. clandestina: "Herr Mink entdeckte deise Art bei Crefeld und schickte mir 3 Arbeiter derisible our
Insight zu". M. rugulosa var. sulcinodorugulosa: "1) С штриховатостью между шипам из Виндавы; 2) с
штриховатостью на area frontalis из Моск. губ. и Ревеля" [1) With the striation between propodeal spi­
nes from Vindava; 2) with the striation on frontal area from Moscow Prov, and Revel].
Material examined. Lectotype of M. rugulosa (designated by Radchenko 2007a), w, "H: fors", "W.
Nyland.", "Coli. Nyland.", "Mus. Zool. H: fors Spec. typ. No. 5077 Myrmica rugulosa Nyl.", "Mus.
Hels. N:o 2208", "Mus. Zool. Helsinki Loan No. 1897" (HELSINKI); paralectotypes: 2 w, "H: fors",
"W. Nyland.", "Mus. Fenn.", "Mus. Zool. H: fors Spec. typ. No. 5075 Myrmica rugulosa Nyl.", "Mus.
Zool. Helsinki Loan No. 1895"; 1 q, "H: fors", "W. Nyland.", "Coli. Nyland.", "Mus. Zool. H: fors Spec.
typ. No. 5076 Myrmica rugulosa Nyl.", "Mus. Zool. Helsinki Loan No. 1896"; 1 q (alate), 1 w, "H:

244
fors", "W. Nyland.", "Coll. Nyland.", "Mus. Zool. H: fors Spec. typ. No. 5074 Myrmica rugulosa Nyl.";
1 vv, "H: fors", "W. Nyland.", "Coli. Nyland."; 2 w, "H: fors", "W. Nyland.", "Coll. Nyland." (HEL­
SINKI); syntypes of M. rugulosa var. sulcinodorugubsa: 1 w, "Ревель" [Revel], 1 w, "Московская губ."
[Moscow Prov.] (MOSCOW); non-type material: > 200 w, several tens of q and m from whole range.
Distribution (Map 97). Europe (absent from British Isles and Iberian Peninsula), Cau­
casus. All old records of this species east of Ural Mts. belong to M. gallienii.
Etymology. M. rugulosa: name derived from the diminutive of the Latin word rugosus
= wrinkle or full.of wrinkles, to describe its fine, rugulose sculpture. M. clandestina: from
the Latin word clandestina = secret or hidden, to describe the somewhat unobtrusive
foraging behaviour of the workers. M. sulcinodorugulosa: from a combination of the names
M. sulcinodis and M. rugulosa because in Nasonov's opinion it has features somewhat
intermediate between the two.
Notes. This species typifies the ragH/osa-complex of the scabrinodis species group (see
also Radchenko and Elmes 2004). It is one of the smallest European Myrmica species.
Populations do not appear to vary morphologically to any great extent. We examined syntype
workers of var. sulcinodorugulosa and concluded that they do not differ from M. rugulosa.
We have not examined type specimens of M. clandestina, but based on the original descrip­
tion and lack of variation in M. rugulosa we agree with the already established synonymy.
M. rugulosa most resembles M. constricta but differs from it by the complete absence of
a longitudinal ridge at the base of scape. On the other hand, it can be confused with
M. gallienii (especially small specimens ofthat species), but it differs from the latter by its
somewhat more angularly curved scape and much more developed reticulated sculpture on
the head dorsum. Moreover, the other castes differ: queens of M. rugulosa are much smaller
and the length of the second funicular segment of males is subequal to the third whereas in
M. gallienii males it is > 1.5 times longer than the third one.
M. rugulosa var. slobodensis (unnecessary proposed replacement name for
M. constricta) was considered to be a junior synonym of M. rugulosa (Arnoldi 1970; Rad­
chenko 1994d; Bolton 1995) until Seifert at al. (2009) placed it under the name

Map 97. Distribution of M. rugulosa.

245
M. constricta (they revived the name constricta from synonymy and raised it to species - see
Notes for that species).
Ecology. M. rugulosa is an oligotopic thermophilous species that is mainly found in dry
meadows, particularly meadows with a very sandy soil. In the more continental parts of its
range it is also found in suitable mountain meadows, open scrubby woodland, forest
clearings, glades and edges providing its nests sites are open to direct sunlight. Generally it
avoids sites with heavy, damp soils. Despite its preference for warm (often hot) micro-
habitats it is a "northern" species and does not live in steppes and dry Mediterranean
biotopes. It is quite tolerant of anthropogenic pressure and is often found inhabiting
pastures, gardens and orchards in rural areas, and lawns, parks and roadside edges in urban
areas. We have observed a very large population in Jutland, Denmark that appeared to be
one super-colony occupying the entire floor area of a large abandoned sand quarry.
Despite its relative abundance in North and Central Europe there has been
relatively little ecological study of M. rugulosa. It appears that in the habitats where it nor-
mally lives its main competitor is Lasius niger, with which it interacts by passive behaviour
and foraging at different times (e.g. Czechowski 1979; De Vroey 1980a; Gallé et al. 1998).
Individual nests may contain up to several thousands of workers and are nearly always
polygynous, we have data from only four nests excavated in Jutland (see above), that had
a mean size of 1200 workers and 19 queens. Nests are built in the soil and usually there are
no above-ground indications (spoil heaps, solaria etc.) of its presence other than small
barely visible entrance holes. Consequently it can be difficult to find the nests of
M. rugulosa, a good tip is to search after heavy rainfall when often one can see freshly ex-
cavated sand on the soil surface.
One of the most studied aspects of M. rugulosa has been the chemical structure of its
recruitment and trail pheromones (Attygalle et al. 1983; Cammaerts, 1980a; Cammaerts
and Cammaerts 1981; Cammaerts et al. 1982). Otherwise, it has been reported to be a host
to M. karavajevi in Sweden (Douwes 1977). Nuptial flights have been recorded in August
and September.

Myrmica rupestris Forel, 1902

(Figs 225-227)

Myrmica smythiesii var. rupestris Forel, 1902: 227, w, India; 1903: 697; Bingham 1903: 269; Ruzsky
1905: 661; Forel 1906: 87; Emery 1921: 41; Stärcke 1935: 260; Menozzi 1939: 296; Weber 1947:
461; Chapman and Capco 1951: 130 (misspelled as rupestria); Collingwood 1970: 374; Bolton
1995: 282.
Myrmica rupestris: Radchenko and Elmes 2001b: 245, 2002: 44, q, m.
Myrmica rugosa var. debilior Forel, 1902: 228, w, India; 1903: 697, 1904b: 22; Ruzsky 1905: 661;
Emery 1921: 40; Menozzi 1939: 295; Chapman and Capco 1951: 129; Bolton 1995: 278, nec Weber
1947: 463. Synonymy by Radchenko and Elmes 2001b: 245; confirmed here.
Myrmica rugosa var. rugososmythiesi Forel, 1902: 228. Synonymy by Forel 1902: 228; Bolton 1995: 282
(as synonym of var. M. rugosa debilior); by Radchenko and Elmes 2001b: 245 (as synonym of
M. rupestris); the latter confirmed here.
Myrmica everesti Donisthorpe, 1929: 455, w, Tibet; Menozzi 1939: 295; Weber 1950: 223; Chapman
and Capco 1951: 125; Collingwood 1970: 374; Radchenko 1994b: 41; Bolton 1995: 279. Synonymy
by Radchenko and Elmes 2001b: 245; confirmed here.
M. rugosa: Weber 1947: 462, misidentification.

246
 Type localities and type specimens. M. smythiesii var. rupestris: "Ekra Раек, 9500'.О. Himalaya
(Smythies)". M. rugosa var. debilior: "Himalaya (Smythies); Darjeeling 3000' à 8000' (Wroughton);
Inde septentrionale (Wroughton); Deota (Smythies); Mysore (Rothney); Kamáon (Schlagintweit)".
M. rugosa var. rugososmythiesi: see Notes below. M. everesti: "Described from five w, taken by Major
R. W .G. Hingston at Jelap La (Tibctian side), at 12,000 ft, 1 Aprile 1924 (Everest Expedition). Type
and paratypes in the British Museum Collection".
Material examined. Lectotype of M. rupestris (designated by Radchenko and Elmes 2001b), w
(upper specimen on the pin with 3 w), "N-W Himalaya, Ekra Peak, 4400 ft (Smythies)", "LXXXIX/12,
M. Smythiesii For. v. rupestris For., Typus" (GENEVA); paralectotypes: 2 w on the same pin as lec-
totype (GENEVA); 1 w, "w Ai. Smythiesii For. var. rupestris Forel, Ekra Peak Himalaya" (GENOA);
lectotype M. rugosa var. debilior (designated by Radchenko and Elmes 2001b), w (upper specimen on
the pin with 3 w), "Himalaya (Smythies)", "LIX/4, M. rugosa r. debilior" (GENEVA); paralectotypes:
2 w on the same pin as lectotype (GENEVA); 1 w with same labels as lectotype (OXFORD); 2 w,
"M. rugoso-Smythiesii Forel w Himalaya Smythies, LX/4", "var. debilior For." (GENOA); 3 w, "Hi-
malaya (Smythies)", "Forel coll.", "M.C.Z. Cotype No. 556" (originally labelled as M. rugosa, see Note
below) (HARVARD); holotype of M. everesti, w, "Himalaya, Jelap La, Tibetan side, 12,000 ft.,
l.iv.1924, Hingston" (LONDON); paratypes: 2 w (LONDON), 1 w (MOSCOW) with the same label
as holotype; non-type material: > 200 w, 11 q, 16 m from India, Nepal, Bhutan and NE Afghanistan.
Distribution (Map 98). India, Nepal, Bhutan, NE Afghanistan.
Etymology. M. rupestris: name from the Latin word rupes = rock with the adjectival
suffix estris = origin or habit, probably to indicate that it was found under a rock or in
a rocky place. M. debilior: name from the Latin word debilis = frail or weak, probably to de-
scribe the much shorter propodeal spines of this form. M. rugososmythiesi: combination of
the names M. rugosa and M. smythiesi, because Forel believed it to be intermediate between
these two forms. M. everesti: named for Mount Everest in the Himalaya because the types
were taken in the Everest region on an expedition to that mountain.
Notes. Originally, Radchenko (1994b) placed M. rupestris in the rubra species group
but now we consider that it is a member of the rugosa -group (see also Radchenko and Elmes

Map 98. Distribution of M. rupestris.

247
2001b). It differs from most other members of this group, M. rugosa, M. aimonissabaudiae
and M. hecate, by the much coarser sculpture on its petiole and postpetiole; M. rupestris
most resembles M. foreliana, from which it differs by the lack of punctations on the surface
of the head and alitrunk between rugae. The socially parasitic M. ereptrix can not be con-
fused with M. rupestris.
The labels on the lectotype and paralectotypes specimens of M. smythiesii var. rupestris
from GENEVA generally agree with the data in Forel's (1902) paper (see above), with only
two exceptions: on the label, altitude and site are "4400 ft" and "N-W Himalaya", but in the
paper they are given as "9500 ft" and "N-O Himalaya". We guess that correct altitude is
9500 ft. because 4400 ft. seems too low for this species. We also studied the holotype and
paratypes workers of M. everesti and concluded that they can not be distinguished from
M. rupestris and confirm the earlier synonymy.
The situation regarding the types of M. rugosa var. debilior is more complicated, par-
ticularly concerning material from HARVARD. The data labels of the lectotype and 5 para-
lectotype workers from GENEVA, OXFORD and GENOA fully correspond with that given
in original description (see above). Moreover, these workers have characteristics, which
separate var. debilior and M. rupestris from M. rugosa, e.g. the much coarser sculpture on
their petiolar and postpetiolar nodes, which have both punctures and coarse short sinuous
rugae, and their coarsely reticulated head dorsum with only the frons being longitudinally
rugulose. We examined the HARVARD "types" of M. rugosa (3 workers), which are actu-
ally not M. rugosa but are undoubtedly the same species which we recognized as var. de-
bilior. Furthermore, their label "Himalaya (Smythies), Forel Coll." agrees with those given
by Forel, and with the data of lectotype and paralectotypes labels of var. debilior from
European Museums. Therefore, we include this HARVARD type series of "M. rugosa" as
paralectotypes of M. rugosa var. debilior. As Weber (1947) almost certainly redescribed
"M. rugosa" based on the "cotypes" from HARVARD, his description should be attributed
to M. rupestris (see also notes to M. rugosa, above).
Lastly, we also studied the "type" material of M. rugosa var. debilior from HARVARD
that have labels (not written by Forel's own hand) "Sind Valley, Cachmire, 7000 ft,
Wroughton", "M. rugosa var. debilior Forel, M.C.Z. Cotype No. 568". These specimens can
not belong to the Forel's original type series of var. debilior because the locality data does
not correspond with the original paper (Forel 1902). Furthermore, these specimens have no
sinuous rugosity on the petiole and postpetiole, which are only densely punctated, and have
well developed reticulation of the head dorsum. Based on these features we identified them
as M. aimonisabaudiae and excluded this material from the type series of any species.
Because Weber (1947) most likely redescribed var. debilior based on the "cotypes" from
HARVARD, his description has to be referred to M. aimonissabaudiae.
The situation with the name "rugososmythiesi" is also somewhat confused. Forel (1902:
228) after the description of var. debilior noted: "On pourrait appeler cette variété rugoso-
smythiesi. Elle forme le passage entre les deux espèces". Most probably, the name rugoso-
smythiesi should be considered to be an unnecessary proposed replacement name for var.
debilior.
Ecology. M. rupestris has been collected at relatively higher altitudes than M. rugosa, be-
tween 2100 and 4100 m. Nests are built in soil, often under stones, in open deciduous and
mixed forests, and places with steppe-like vegetation (Philip Ward, pers. comm.). Like
M. rugosa, queens and males have been found in nests of M. rupestris from the end of May
to the end of June, much earlier than in Euro-Siberian species.

248
 Myrmica ruzskyana nom. nov.
(Figs 228, 229)

Myrmica ruzskyana (replacement name for Myrmica smythiesi var. exigua Ruzsky, 1915; 440).
Myrmica smythiesii var. exigua Ruzsky, 1915; 440, w, m, Tibet (junior primary homonym of Myrmica
exigua Buckley, 1867; 72); Eidmann 1941: 16; Weber 1947: 461; Bolton 1995: 278.

Type localities and type specimens. "Мсстонахожд.: речка Ба-чю, 12,000', Кам, басе. р. Голубой, 2/3.
VIII. 1900 (б1, ww) (Козлов)" [Locality: river Ba-chu,'12,000', Kam, bassin of riv. Yangtze, 2/3.viii.l900
(m, ww), leg. Kozlov].
Material examined. Lectotype (designated here), w, "речка Ба-чю, 12 т', Кам, б. Голубой р., Козлов,
2/3.VI1I.00" [river Ba-chju, 12,000', Kam, bassin of riv. Yangtze, 2/3.viii. 1900] (MOSCOW); para-
lectotypes: 14 w, 1 m with the same label (MOSCOW, PETERSBURG).
Distribution (Map 99). Tibet.
Etymology. M. ruzskyana: dedicated to the famous Russian myrmecologist M. D. Ruz­
sky (see biography in Chapter 1.6) who originally named the species M. exigua. M. exigua: _
from the Latin word exiguus = slight or slender, to describe this species appearance.
- Notes. Because the name M. smythiesi var. exigua Ruzsky, 1915 is preoccupied, we pro­
pose the replacement name M. ruzskyana. We place this species to the smythiesii species
group.
M. smythiesi var. exigua Ruzsky was for many years an enigmatic taxon but based on
our recent examination of Himalayan and south-west Chinese Myrmica combined with an
examination of Ruzsky's types, we consider it to be a good species.
M. ruzskyana is very similar to M. bactriana both by "subjective" and morphometrical
diagnostic features, and differs from it by its longer scape that has more abundant and
suberect hairs (see also notes to M. bactriana, above). Queens are unknown.
Ecology is unknown, except that it was found at the altitude ca. 3660 m (12,000 ft.)
a.s.l. in a subalpine meadow.

Map 99. Distribution of M. ruzskyana.

249
 Myrmica sabuleti Meinert, 1861
(Figs 230-232)

Myrmica sabuleti Meinert, 1861: 327, w, m, Denmark; Bondroit 1918: 102, 1920a: 151; Santschi
1931b: 345; Karawajew 1934: 80; Stitz 1939: 94; Holgersen 1940: 184; Novak and Sadil 1941: 79;
Stärcke 1942: 25; Weber 1948: 288; Sadil 1952: 246; Collingwood 1958b: 70; Hauschteck 1965:
325; Bernard 1967: 117; Arnoldi 1970: 1841; Baroni Urbani 1971: 31; Pisarski 1975:14; Kutter
1977: 68; Arnoldi, Dlussky 1978: 534; Collingwood 1979: 54; Espadaler 1981b: 189; Seifert 1988:
31; Casevitz-weulersse 1990: 137; Atanassov, Dlussky 1992: 97; Seifert 1994: 12; Radchenko
1994b: 42, 1994c: 144, 1994d: 80; Bolton 1995: 282; Seifert 1996: 226; Radchenko et al. 1997:
490; Czechowski et al. 2002: 29; Radchenko, Czechowska and Czechowski 2004: 50; Seifert 2005:
7; 2007: 199; Czechowski, Czechowska and Radchenko 2007: 143.
Myrmica lobicornis subsp. sabuleti: André 1883: 318.
Myrmica scabrinodis (var., subsp.) sabuleti: Emery 1895a: 314, 1908a: 176; Bondroit 1910: 495;
Donisthorpe 1915a: 143; Emery 1916: 120, 1921: 40; Menozzi 1922: 325; Soudek 1922: 44; Finzi
1924: 12, 1926: 101; Karawajew 1929: 208; Menozzi 1936: 270, nec Forel 1915: 29.
Junior synonym of M. lobicornis: Dalla Torre 1893: 111, not confirmed, see Notes below.
Myrmica rubra var. scabrinodolobicornis Forel, 1874: 77, w, q, m, Switzerland; Dalla Torre 1893:
112; Ruzsky 1896: 73. Synonymy by Sadil 1952: 249 (as synonym of M. lonae); by Radchenko
1994d: 80; Bolton 1995: 283 (as synonym of M. sabuleti); the latter confirmed here (see also
Notes below).
Myrmica scabrinodolobicornis: Ruzsky 1896: 173.
Myrmica scabrinodis var. scabrinodolobicornis: Emery 1921: 40.
Myrmica sabuleti st. lonae var. scabrinodolobicornis: Santschi 193lb: 347 (unavailable name).
Myrmica sabuleti subsp. scabrinodolobicornis: Weber 1948: 296.
Myrmica scabrinodis: Ruzsky 1905: 680 {part); Forel 1915: 29; Bondroit 1918: 102, m, nec w, q;
1920a: 147, m; Müller G. 1923: 43, misidentifications.
Myrmica granulinodis: Emery 1908a: 177; Sadil 1952: 246; not confirmed, see Notes to M. granuli-
nodis in Chapter 3.5.
Myrmica bessarabica: Weber 1948: 288; Sadil 1952: 246; not confirmed, see Notes to M. bessarabica
in Chapter 3.5.
Myrmica sabuleti subsp. lonae: Bernard 1967: 117; Arnoldi 1970: 1841; Seifert 1988: 31; Atanassov,
Dlussky 1992: 97; Radchenko 1994d: 80; Bolton 1995: 281; not confirmed, see Notes to M. lonae,
above.
Myrmica scabrinodis var. piloscapa (Sicl): Bernard 1967: 117; not confirmed, see Notes to M. scabri-
nodis, below.
Myrmica sabuleti var. spinosior: Seifert 1988: 31; Casevitz-weulersse 1990: 137; Bolton 1995: 283; not
confirmed, see Notes to M. spinosior, below.
Myrmica scabrinodis var. scabrinodosabuleti: Radchenko 1994d: 80; not confirmed, see Notes to
M. scabrinodis, below.

Type localities and type specimens. M. sabuleti: "Jeg har kun eengang fundet den under Stene
i Buurkrat ved Norre-Vosborg i Bcgyndelsen af August". Myrmica rubra var. scabrinodolobicornis: no
locality given.
Material examined. Lectotype of M. sabuleti (designated here), w (specimen damaged, its meso-
soma was broken, head with pronotum stuck to the rest of mesosoma), "9/8. 1858, Buur-Krat Vestjyll,
Meinert" (COPENHAGEN); paralectotypes, 3 m with the same labels (COPENHAGEN); lectotype of
Myrmica rubra var. scabrinodolobicornis (designated here), w (upper w on the pin with q and
2 w), "scabrinodo-lobicornis [hardly readable text], 25.VI.8" (LAUSANNE); paralectotypes: 1 q and
1 w on the same pin as lectotype; 1 m, "<S M. scabrinodo-lobicornis Savone" (LAUSANNE) (see Notes
below); non-type material: several hundred w, about 100 q and m from whole range.

250
 Distribution (Map 100). Europe (to the north until southern Sweden, Finland and Nor-
way), Caucasus; all previous records of this species east of Ural Mts. belong to M. lonae (or
any other species in older papers).
Etymology. M. sabuleti: name derived from the Latin word sabulum = sand, sandy soil,
to indicate its habitat. M. scabrinodolobicornis: named as a combination of the names
M. scabrinodis and M. lobicornis to indicate that it had features of both these species.
Notes. We examined the type specimens (worker and 3 males) of M. sabuleti, preserved
in COPENHAGEN and designated the worker as the lectotype. Despite a rather confused
taxonomic history (see below) the modern treatment of M. sabuleti accords well with these
types. The pattern of wing venation of all the paralectotype males is unusual for genus Myr-
mica because they have completely separated cell 1 +2r on the forewings; however, this fea-
ture can vary occasionally between individual males and has no taxonomic relevance in
this case.
M. sabuleti belongs to the scabrinodis species group and it typifies the sabuleti-compiex
within that group. This complex contains all.species with males having a relatively long
scape (equal to total length of 4-4.5 basal funicular segments). The workers may be con-
fused with several species, but generally they are characterized by a quite narrow frons
(mean FI 0.33) and a well-developed lobe at the scape base. Males are distinguished from
the socially parasitic members of this group, M. hirsuta, M. vandeliandM. bibikoffi, by the
absence of long erect to suberect hairs on the head margins.
Although the form was recognised quite soon after Nylander described M. scabrinodis,
early myrmecologists found M. sabuleti to be a somewhat enigmatic taxon (e.g. Ruzsky
1905; Forel 1915; Bondroit 1918, 1920a; Müller G. 1923, etc.). A few authors treated it as
a good species but others considered as being an infraspecific form of M. rubra or M. scabri-
nodis or even a synonym of M. lobicornis (see synonymy above). Some authors confused it
with forms of Myrmica that are now considered to be good species in their own right (e.g.
M. lonae), some considered it to be synonym of species now incertae sedis in the genus
Myrmica (e.g. M. granulinodis, M. bessarabica), while others considered it synonymous

Map 100. Distribution of M. sabuleti.

251
with forms that are now synonymised with other species, usually forms of M. scabrinodis
(e.g. M. pilosiscapus). It should be noted also that Forel's (1915) "var. sabuleti" is in fact
M. scabrinodis, while Bondroit (1918, 1920a) and Müller G. (1923) considered M. sabuleti
under the name "scabrinodis".
Sadil (1952) and Collingwood (1958b) were the first people to treat M. sabuleti cor-
rectly according to the modern view; even after these publications many naturalists con-
tinued to consider M. sabuleti as synonymous with M. scabrinodis, a view that had
unfortunate consequences for British nature conservation efforts aimed at preserving the
Large Blue Butterfly Phengaris avion (L.) (Elmes and Thomas 1992).
The situation with M. rubra var. scabvinodolobicovnis seems even more complicated.
Firstly, Forel (1874) did not provide a proper description of this form. On page 77 (loc.
cit.) he wrote about workers: "Le formes scabvinodo-lobicovnis sont fréquentes variation
dans la striure de l'arie frontale et dans la position du lobe du scape des antennes", but later
he only used this name for queens and males, not providing any description of the latter.
Nevertheless, we studied the type series of var. scabvinodolobicovnis from LAUSANNE,
this originally included the following material:
a) 1 q, 2 w on one pin, "scabvinodo-lobicovnis [hardly readable text], 25.VI.8";
b) 1 m, " M. scabvinodo-lobicovnis Savone"
e) 1 w, "M. scabvinodo-lobicovnis w, [locality not readable];
d) 1 q, "M. scabvinodo-lobicovnis q Savone" ;
e) 3 w on one pin (1 specimen without head and gaster) "scabvinodo-lobicovnis w Vaun.
Gerr (?)", "M. schencki Emery";
f) 1 q, "scabvinodo-lobicovnis V. Clarmont", "M. schencki Emery $".
We determined the specimens as follows: on pins (a) and (b) are M. sabuleti (note that
males of M. sabuleti and M. lonae are practically indistinguishable); the worker on pin (с)
is M. lobulicornis; the queen on the pin (d) is M. lonae, while workers and queen on pins
(e) and (f) are M. slovaca. To avoid further confusion and to maintain stability of the no­
menclature, we designated as the lectotype of var. scabvinodolobicovnis the upper worker
from pin (a), and as paralectotypes the bottom worker and queen from pin (a) and the male
on pin (b). Based on this treatment var. scabvinodolobicovnis is a clear synonym of M. sa­
buleti. All other "types" have lost their taxonomic value and should be excluded from the
type series.
Ecology. The ecology of M. sabuleti has been relatively well studied during the last 20
years mainly because of its importance as the primary ant host of Phengavis avion, which
figures in Red Data Books throughout Europe (see Chapter 1.3.6 and Settele et al. 2005).
The average size of nest is important in determining the suitability of a site to support
a population of P. avion because Thomas and Wardlaw (1992) showed that it took the
entire annual production of a nest of about 350 workers to rear one caterpillar. Also M. sa­
buleti has evolved interesing mutualisms with other grassland lycaenid butterflies (e.g.
Thomas 1983). It is also the host for several host-specific parasitic Myvmica species:
M. hivsuta, M. bibikoffi, M. lemasnei and possibly M. lauvae as well as the more generalist
M. kavavajevi. At this time it is not known why M. sabuleti should be such a good host for
various social parasites. Perhaps it is related to the species' habitat preferences or maybe to
social factors, for example compared to most other European Myvmica species, M. sabuleti
workers generally lay far fewer eggs (Wardlaw and Elmes 1998); also it is somewhat easier
to introduce foreign queens into existing cultures (personal observations) and it seems to
have symbioses with aphids (see below).

252
 For anyone who has studied their ecology it is hard to see how the female castes of
M. sabuleti were ever confused with M. scabrinodis (see notes above): apart from the ob-
vious morphological differences, they are adapted to living in warmer habitats than
M. scabrinodis (e.g. Brian 1964; Brian et al. 1976; Doncaster 1983), both adults and larvae
having a lower basal metabolism and larvae grow and develop more slowly than those of
M. scabrinodis when reared at the same temperature (Elmes and Wardlaw 1983a) and
workers have a longer foraging period (Elmes 1982). Habitats are usually grasslands, where
M. sabuleti favour short-turf in northern latitudes and live under quite long grass in
southern Europe (Thomas et al. 1998). In Northwest Europe it could be considered a relict
species of the post-glacial optimum that has persisted because of anthropogenic modifica-
tions of the natural environment (Thomas 1993). M. sabuleti is not normally considered
a woodland species though it can live in open woodland glades, especially in eastern Europe,
where summer temperatures are higher. It is often found in mature sand-dune systems, par-
ticularly those that have become grassed and are grazed by animals, the types were proba-
bly found in such a sandy place, hence its name.
Workers appear to be generalist predator scavengers, like most of the other Myrmica
species. There is circumstantial evidence that it might eat the eliasome of Viola seeds.
M. sabuleti do not often forage on obvious aphid colonies, more usually it maintains small
cultures of aphids, particularly those species that live on the base of the stems at ground
level; one can often find these colonies enclosed with soil and a few workers inside
guarding and tending the aphids (personal observations). Workers were observed tending
colonies of aphids on the underside of leaves of low-growing Clinopodium vulgare (Wood-
Baker 1977) and we suspect that M. sabuleti may be more dependent upon such less-
obvious associations than previously appreciated.
M. sabuleti mainly competes for living space with M. scabrinodis at the cooler end of its
niche, and with M. specioides and M. schencki at the hotter end. The processes involved in
these competitive interactions have been explored in numerical spatial-simulation models
(see Chapter 1.3.6). The average numbers of workers per nest varied from 12 - > 4000 in
a study of nearly 300 nests censused in southern England; average nest size on acid grass-
land was smaller (about 150 workers) than nests on chalk grassland (about 370 workers)
but the average number of queens per nest in both types of habitat was the same (just over
1 per nest) (Elmes and Wardlaw, 1982a). A more detailed study of the chalk grassland
population showed that above-nest vegetation was implicated in determining the size and
productivity of the nest (Elmes and Wardlaw 1982b, c). A population living under stones
on limestone grassland had much larger more polygynous colonies, averaging about 1200
workers with 15 queens per nest (Elmes 1974b), this was the population that contained
the type series of the social parasite M. hirsuta (see Notes for that species above). Seppä
(1996) demonstrated lowered levels of relatedness among workers commensurate with all
queens actively reproducing. Brian (1972) followed the populations of several individual
nests for some years and showed there to be quite a high turnover of queens and workers,
one nest studied supported a population of the parasite M. karavajevi. In an appendix to
a paper Brian and Elmes (1974) showed that variations in the annual productivity of this
population was directly related to changes in the annual insolation.
M. sabuleti has been used quite frequently in comparative studies of chemical ecology
(e.g. Cammaerts et al. 1981; Evershed et. al. 1982). In a more recent study of the cuticular
hydrocarbons (Elmes et al. 2002) it was shown that M. sabuleti had a rather "simpler"
chemical profile than other species, in particular workers had less Dienes and less Di-methyl

253
alcanes than M. scabrinodis, the two species can be easily discriminated by their chemical
profiles. M. sabuleti has been used as a model species in studies of food competition and re­
cruitment (De Vroey 1980a; Cammaerts and Cammaerts 1980; Debiseau and Pasteels 1994;
Debiseau et al. 1997).
Nuptial flights occur from August to the end of September. Mixed swarms with other
Myrmica species are common (Woyciechowski 1990c and personal observations) and where
it co-exists with M. lonae mixed flights are normal (see notes to M. lonae).

Myrmica salina Ruzsky, 1905

(Figs 233-235)

Myrmica scabrinodis var. salina Ruzsky, 1905: 687, w, q, m, Russia and Kazakhstan; Emery 1908a:
177, 1921:40; Finzi 1926: 102, nec Sadil 1952: 249; Arnoldi 1970: 1842, 1976: 57; Arnoldi, Dlussky
1978: 534; Seifert 1988: 25; Radchenko 1994c: 77; Seifert 1996: 228, 2002: 96; Radchenko and
Elmes 2004: 229; Radchenko, Stankiewicz and Selezncw 2004: 55; Seifert 1994: 13; Radchenko,
Elmes and Alicata 2006: 228; Seifert 2007: 201, misidentifications.
Myrmica schencki var. salina: Weber 1948: 302.
Myrmica salina: Radchenko and Elmes 2009b: 520, w, q, m (redescription).
Junior synonym of M. lonae: Sadil 1952: 249; not confirmed, see Notes to M. lonae, above.
Myrmica slovaca: Seifert 1988: 25, 1996: 228, 2002: 96; Radchenko and Elmes 2004: 229; Seifert 2007:
211; not confirmed, see Notes to M. slovaca, below.

Type localities and type specimens. "Мсстонахожд.: Оренбургская губ., Челябинск, у., окр. оз. Сары- \
Куль, самки и самцы 2-5.VII 1.94. Тобольская, оз. Горькое, на пути из ст. Беловской в Петропавловск,
солонец, 6.VII.96; окр. оз. Камсннаго, бл. с. Курсинскаго, Кург. окр., 30.VI.96" [Localities: Orenburg-
skaya Gubernia, Chelyabinsky uezd, vicinity of lake Sary-Kul', ww and mm, 2-5.vii.94. Tobolskaya
Gubernia, lake Gor'koe, on the way from st. Belovskaya to Petropavlovsk, salted marsh, 6.vii.96;
vicinity of Lake Kamennoe, near vil. Kureinskoe, Kurgansky okrug, 20.vi.96].
Material examined. Lectotype (designated by Radchenko and Elmes 2009b), w (on the pin with q),
"Myrm. scabrinodis v. salina R. w, q Gouv. Orenburg, Sary-Kul See. Salzbiden" (GENOA); paralec-
totypes: 1 q on the pin with lectotype; 1 q, "Myrm. scabrinodis v. salina R. q Gouv. Orenburg, Sary-
Kul See", 1 m, "Myrm. scabrinodis v. salina R. m Gouv. Orenburg, Sary-Kul See" (GENOA).
Distribution (Map 101). It is known only from the type localities in the south of West
Siberia and northern Kazakhstan.
Etymology. M. salina: from mediaeval Latin salina = salt pan, or salted place being de­
rived from the Latin word salis = salt, here it describes the type habitat "salted marsh".
Notes. Myrmica scabrinodis var. salina was described by Ruzsky (1905: 687) based on
workers, queens and males from different places in West Siberia and Kazakhstan (see
above). The taxonomic history of M. salina is extremely complicated and rather puzzling
because for many decades there were no types (presumed to be lost) and the original de­
scription of this species was ambiguous. Here we outline the problem. The original text is
in Russian so first we translate the description and some of Ruzsky's further comments
{loc. cit., p. 700):
Description: "(workers). Frontal carinae well developed, raised at the base, lobe-like (in
typical scabrinodis they are smaller). Lobe at the bend of scape transversally oblique (less
oblique than in scabrinodis and less transverse than in lobicornis), and looks like trans­
versal dent or thick scale. Middle part of frontal triangle smooth and shiny. Sides of head

254
dorsum with large reticulation, surface between reticulation punctated but appears shiny.
[Propodeal] spines long and straight. Petiolar dorsum angled. Outstanding hairs more
sparse [presumably compared to scabrinodis], on the gastral tergites almost absent.
Brownish-red with dark brown or blackish-brown head dorsum and first gastral segment;
antennae, mandibles, legs and apex of gaster lighter. Length 4.7-5 mm.
(queens). With same features as workers. Colour somewhat darker than in workers,
alitrunk with brownish-black patches. Basal half of wings brownish. Length 5-6 mm.
(males). Whole head very finely punctato-striated (in typical scabrinodis head, espe-
cially on sides and rear part, with quite coarse irregular rugosity). Antennal scape thickened
in the middle. Outstanding hairs on the body, legs and antennae sparser [presumably com-
pared to scabrinodis?]. Alitrunk almost without hairs. Colour of wing as in queens. Length
of scape as in typical scabrinodis."
Additional comments : "... This variety is interesting because its queens and workers by
the dark colour and by the almost transverse, scale-like lobe at the bend of antennal scape,
is similar to M. lobicornis, but its males on main features are similar to M. scabrinodis males
and differs only by sculpture of head and sparser pilosity. This species, indubitably, is most
similar to the variety schencki Emery, and both are intermediate between scabrinodis and
lobicornis. ... M. schencki together with var. salina are intermediate between M. scabri-
nodis and M. lobicornis, but salina is more close to the first and schencki - to the latter".
Unsurprisingly, many subsequent authors treated var. salina as an infraspecific form of
either M. scabrinodis or M. schencki. Thus, Emery (1908a: 177) noted: "Nach Untersu-
chung von Originalenexemplaren halte ich diese var. [i.e. salina] für eine Übergangsform von
scabrinodis zu schencki". Finzi (1926) and Weber (1948) just repeated Emery's opinion;
moreover, Weber considered it as a var. of M. schencki. Somewhat mysteriously, Sadil
(1952) without any comments regarded salina as a junior synonym of M. lonae (despite the
first name having priority).

Map 101. Distribution of M. salina.

255
 Arnoldi (1970) considered that M. salina could be discriminated from any other related
species (i.e. those of the scabrinodis-group sensu Radchenko 1994b) by its very narrow
frons (FI < 0.30) and by raised frontal lobes, even though these were important features of
M. slovaca. Therefore he revived name salina from synonymy and raised M. salina to the
species level, designating as the lectotype a worker (MOSCOW) that at most could only be
a neotype according to its data label - "Kulundinskaya step', Blagodarnoe, 19 July 1969,
N 504 (Pavlova)". Seifert (1988) followed Arnoldi's treatment, but synonymised the name
M. slovaca with M. salina and later corrected Arnoldi's error, designating the "lectotype"
as a neotype (Seifert 2002).
A fundamentally different view was taken by Radchenko (1994e), who believed that
Ruzsky's description (see above) indicated that workers of M. salina have a transversai
lobe at the bend of scape, so Radchenko placed it in the schencki-group as a junior synonym
of M lacustris (= M. deplanata). Consequently, he considered "M. salina" sensu Arnoldi
and Seifert to be a junior synonym of M. slovaca (see also Radchenko 1994d). However,
Seifert (2002) disagreed with this approach and reaffirmed his previous opinion: he revived
M. salma from synonymy, once more considering M. slovaca as its junior synonym. Sub-
sequently we accepted Seifert's treatment of M. salina as definitive (Radchenko and Elmes
2004; Radchenko, Elmes and Alicata 2006) because without the original type specimens
any further discussion would be futile.
By serendipity we found specimens (one worker, two queens and one male) in the
Emery collection (GENOA) that we believe belong to Ruzsky's original type series of
M. salina (see above). This led us to reappraise the taxonomic position of this species.
Morphologically, these specimens did not contradict the original description and their
data-labels appear to be Ruzsky's and conform to the collection data. Therefore we
designated the worker as the lectotype and the other 3 specimens as paralectotypes, and
redescribed all three castes of M. salina based on these type specimens (Radchenko and
Elmes 2009b).
Based on the rediscovered type material, M. salina obviously belongs to the scabrinodis
species group, most probably to the specioides-complex. The worker and queens have
a moderately developed horizontal lobe at the base of their scape and a relatively wide frons
(FI 0.36). By these features, M. salina could be confused with M. scabrinodis, but the male
has the specioides-complex features of a combination of a relatively short scape and short
standing hairs on the scape and tibiae. On the other hand, the worker of M. salina well dif-
fers from M. slovaca, which have much narrower frons (FI < 0.30).
Ecology is poorly known, but most probably M. salina inhabits relatively, wet, often
halophytous biotopes in steppes of West Siberia and Kazakhstan.

Myrmica saposhnikovi Ruzsky, 1904

(Figs 236-238)

Myrmica lobicornis r. saposhnikovi Ruzsky, 1904b: 3, w, Kazakhstan.

Myrmica scabrinodis subsp. saposhnikovi: Ruzsky 1905b: 701; Emery 1908a: 180; Kuznetsov-ugam-
sky 1927: 193, w, q, m; Weber 1950: 209, nec Teranishi 1940: 15, q, misidentification.
Myrmica saposhnikovi: Arnoldi 1976: 553; Tarbinsky 1976: 35; Radchenko 1994b: 43, 1994c: 139,
1994Í: 85; Bolton 1995: 283, nec Pisarski 1969a: 228, Pisarski 1969b: 296; Pisarski, Krzysztofiak
1981: 155, misidentifications.

256
Myrmica scabrinodis subsp. intermedia Kuznetsov-ugamsky, 1927: 194, w, Kazakhstan [unresolved
junior secondary homonym of Myrmica intermedia (Wheeler, 1915: 61)]. Synonymy by Radchenko
1994Í: 85; not confirmed, considered here as incertae sedis in Myrmica (see Chapter 3.5).
Myrmica saposhnikovi subsp. bergiana Tarbinsky 1976: 38, w, q, Kyrgyzstan; Bolton 1995: 277.
Synonymy by Radchenko 1994f: 85; confirmed here.

Type localities and type specimens. M. lobicornis r. saposhnikovi: "Мсстонахожд.: Копальск. у., сев
склон Джунгарского Алатау, в долине р. Баскана, 1000 м; густой еловый лес (13.VIII.02); дол. р. Или близ
Джаркснта, степь (20.viii.02)" [Localities: Kopalsky uezd, northern slope of Dzhungarian Alatau, in
valley of riv. Baskan, 1000 m, dense fire forest (13.viii.02); valley of riv. Iii near Dhzarkent, steppe
(20.vii.02)]. M. saposhnikovi subsp. bergiana: "Материал. Голотип (рабочий) - Киргизия: Чуйская
долина, окрестности с. Калинина, 17.IX 1970. Гинетип (самка) - собрана там же в тс же сроки; паратипы
- две самки и пять рабочих из того же места" [Material. Holotype (worker) - Kirghizia: Chuiskaya val­
ley, vicinity of viii. Kalinino, 17.ix.1970. Gynetype (queen) - collected in the same place and at the
same time; paratypes - two queens and five workers from the same locality].
Material examined. Lectotype of M. lobicornis r. saposhnikovi (designated by Radchenko and
Elmes 2009b), w, "Myrmica saposhnikovi R. w Fluss Baskan,. Alatoo, Ssemiretschinsk" (Ruzsky's
original label, written by pencil; see also Notes below) (GENOA); holotype of M. saposhnikovi subsp.
bergiana, w, "Киргизия: Чуйская дол., окр. с. Калинина 17.IX 1970, Тарбинский" [Kirghizia: Chuiskaia
val., vicinity of vili. Kalinino, 17.ix.1970, Tarbinsky] (MOSCOW); paratypes: 3 w, with same label as
holotype (MOSCOW); non-type material: > 50 w, 5 q, 3 m from Kyrgyzstan.
Distribution (Map 102). Dzhungarian Range, Tien-Shan, Alai.
Etymology. M. saposhnikovi: named for the well-known Russian geographer and
botanist, Professor Vasily Vasilievich Saposhnikov (1861-1924) of Tomsk University.
M. bergiana : name derived from M. bergi, to which this form is superficially similar by some
features. '
Notes. M. saposhnikovi belongs to the lobicornis species group and well differs from
the sympatric M. kirghisorum by the very small vertical dent or even ridge at the base of the

Map 102. Distribution of M. saposhnikovi.

257
scape, by the less extended frontal lobes and by its subtriangular petiole without a declined
dorsal plate (seen in profile).
For many years the types of M. saposhnikovi were considered to be lost; therefore Rad-
chenko (1994Í) designated a neotype of this species (worker, Alma-Atinsky Natural Reserve,
No. 69, 2250 M a.s.l., 23.09.1969, leg. Antsyferov). However, recently we found in Emery's
collection (GENOA) one specimen (worker) labelled as "Myrmica saposhnikovi R. [w] FIuss
Baskan, Alatoo, Ssemiretschinsk" (Ruzsky's original label, written by pencil). The data label
corresponds with the type localities, given by Ruzsky (1904b) and all main diagnostic fea-
tures of this specimen fully correspond with the original description and later treatments of
M. saposhnikovi (see Arnoldi 1976; Tarbinsky 1976; Radchenko 1994c, f). Therefore, we
designated this worker as the lectotype of M. saposhnikovi (Radchenko and Elmes 2009b)
and the neotype (designated by Radchenko 1994f) has lost its taxonomic status.
The types of M. scabrinodis subsp. intermedia are most probably lost, but Radchenko
(1994f) synonymised it with M. saposhnikovi. Now, after obtaining additional material
from the Middle Asian mountains and careful re-investigation of Kuznetsov-Ugamsky's
(1927) treatments of all Myrmica species, we believe that subsp. intermedia should be con-
sidered as an unrecognizable taxon in Myrmica (see Chapter 4.5).
We also investigated the holotype and paratype workers of M. saposhnikovi subsp.
bergiana, which slightly differed from M. saposhnikovi by having a somewhat coarser sculp-
ture of the body; nevertheless, we consider that subsp. bergiana is a synonym of
M. saposhnikovi because they have the same main diagnostic features and we can find no
other feature to make a clear separation.
However, M. saposhnikovi has a strange dichotomy in its habitat preferences. On the
one hand, it inhabits both fir forests and subalpine meadows (relatively cool habitats) at al-
titudes over 1000 m a.s.l.; Kuznetsov-ugamsky (1927: 193-194) noted that "This form to-
gether with M. rugosa dshungarica is one of the main elements of the myrmecofauna of the
belt of fir forests in the Central Tien-Shan" (our translation from Russian). On the other
hand, it lives also in much hotter and drier steppe-like habitats at lower altitudes, as was
stated by Ruzsky (1904b). We think it possible that Ruzsky (1904b) included two mor-
phologically similar species in the type series of M. saposhnikovi: one from the mountain
forests (Dzhungarian Alatau; the lectotype specimen belongs to this series) and one from
the steppes near Dhzarkent. As the Dhzarkent "type" material appears to be lost, it is open
to question whether "M. saposhnikovi" is one species with some sort of variable ecology,
or whether it contains two hardly distinguishable sibling species, each adapted to different
habitats. Unless the Dhzarkent material is found, this question can be resolved only by
a combination of ecological and perhaps molecular genetical studies of the different popu-
lations in the Middle Asian mountains.
The records of this species for Mongolia (Pisarski 1969a: 228, 1969b: 296; Pisarski,
Krzysztofiak 1981: 155) in fact belong to M. pisarskii, M. kasczenkoi and M. angulinodis
(corresponding material in WARSAW and BUDAPEST, examined). We are sure Teranishii
(1940) record of this species from Korea is erroneous and must be referred to any of the
lobicornis-group species known from Korea (Radchenko 2005).
Ecology As emphasised by Kuznetsov-ugamsky (1927), M. saposhnikovi (together with
M. dshungarica) is one of the main elements of the ant fauna in the fir forests of the Cen-
tral Tien-Shan. It was found in fir forests and subalpine meadows at altitudes between 1000
and 2900 m and also somewhat surprisingly (as noted above), in steppe-like habitats at
altitudes less than 600 m a.s.l.
 Myrmica scabrinodis Nylander, 1846: 930
(Figs 239-241)

Myrmica scabrinodis Nylander, 1846a: 930, w, q, m, Finland; 1846b: 1052; Mayr 1855: 411; Nylander
1856: 81; Smith F. 1858: 115; Meinert 1861: 53; Saunders 1880: 215; André 1883: 319; Nasonov
1889: 36; Wasmann 1891: 298; Dalla Torre 1893: 116 (see alsoadditional synonymy here); Ruzsky
1895: 30,1896: 73; Emery 1898: 126; Ruzsky 1905: 680 {part.); Emery 1908a: 174; Bondroit 1912:
351; Stitz 1914: 71; Donisthorpe 1915a: 125; Karawajew 1916: 504; Emery 1916: 120, 1920: 60,
1921: 40; Santschi 1921: 110; Menozzi 1922: 325; Soudek 1922: 43; Müller G. 1923: 43; Finzi
1926: 98; Karawajew 1926a: 65, 1926b: 95, 1927b: 259; Santschi 1931b: 341; Karawajew 1934: 77;
Weber 1950: 202; Sadil 1952: 249; Collingwood 1958b: 70, Hauschteck 1965: 325; Bernard 1967:
116; Arnoldi 1970: 1841; Baroni Urbani 1971: 34; Tarbinsky 1976: 41; Dmitrienko, Petrenko 1976:
14; Kutter 1977: 69; Arnoldi, Dlussky 1978: 534; Collingwood 1979: 55; Espadaler 1981b: 189;
Seifert 1984: 1, 1988: 27; Atanassov, Dlussky 1992: 95; Seifert 1994: 11; Radchenko 1994b: 42,
1994c: 144, 1994d: 79, 1994g: 107; Seifert 1996: 224; Radchenko et al. 1997: 489; Radchenko,
Czechowska and Czechowski 2004: 50; Seifert 2007: 197; Radchenko 2007a: 29, nec Forel 1915:
29; Bondroit 1918: 101, 1920a: 147, misidentifications.
Myrmica rubra r. scabrinodis: Forel 1874: 76; Emery and Forel 1879: 460; Emery 1895a: 313; Forel
1904a: 374; Wheeler W. M. 1908b: 406.
Myrmica sabuleti: Forel 1915: 29, misidentification.
Myrmica specioides: Bondroit 1920a: 146, m, misidentification.
Myrmica scabrinodis var. rugulosoides Forel, 1915: 29, w, Switzerland; Kutter 1924: 8, q, m; Santschi
1931b: 342; Stitz 1939: 93; Novak and Sadil 1941: 79; Sadil 1952: 251. Synonymy by Bernard
1967: 116; Banert and Pisarski 1972: 350; Collingwood 1979: 55; Seifert 1984: 1, 1988: 27; Rad-
chenko 1994 d: 79; Bolton 1995: 282; Radchenko et al. 1997: 489; Czechowski et al. 2002: 27;
confirmed here.
Myrmica scabrinodis rugulosa var. rugulosoides: Emery 1916: 120 (unavailable name).
Myrmica rugulosoides: Bondroit 1918: 102; Müller G. 1923: 42; Finzi 1926: 94, fig. 6, w, q, m; Pçtal
1963: 379; Baroni Urbani 1971: 30; Kutter 1977: 68.
Myrmica rolandi Bondroit, 1918: 101, m, nec w, q (part.), misidentification (see Notes below and to
M. aloba).
Myrmica pilosiscapus Bondroit, 1920a: 147, w, q, m, Belgium, Germany, France, Switzerland, Corsica
and Great Britain (also described as new by Bondroit, 1920b: 301); Sadil 1952: 256; Arnoldi 1970:
1842; Baroni Urbani 1971: 26. Synonymy by Bernard 1967: 117; van Boven 1977: 120 (as syno-
nym of M. sabuleti); Kutter 1977: 69; Collingwood 1979: 55; Seifert 1988: 27; Casevitz-weulersse
1990: 137; Radchenko 1994d: 79; Bolton 1995: 281 (as synonym of M. scabrinodis); the latter con-
firmed here.
Myrmica scabrinodis var. pilosiscapus: Finzi 1926: 102; Santschi 1931b: 343; Stitz 1939: 94.
Myrmica sabuleti var. pilosiscapus: Weber 1948: 296
Myrmica scabrinodis var. ahngeri Karawajew, 1926a: 66, w, Russia; 1927b: 259; Weber 1950: 203;
Arnoldi 1970: 1841, w, nec m; Arnoldi, Dlussky 1978: 534. Synonymy by Seifert 1988: 16; Bolton
1995: 277; Güsten et al. 2006: 29 (as synonym of M. specioides); Radchenko 1994d: 79 (as syno-
nym of M. scabrinodis); the latter confirmed here.
Myrmica rolandi var. reticulata Stärcke, 1942: 25 (first available use of Myrmica scabrinodis st. rolandi
'var. reticulata Santschi, 1931b: 344, w, m, Spain; Weber 150: 208). Synonymy by Seifert 1988: 27;
Radchenko 1994d: 79; confirmed here.
Myrmica (Myrmica) scabrinodis var. scabrinodosabuleti Sadil, 1952: 253, w, m, Czechoslovakia;
Bolton 1995: 283. Synonymy by Radchenko 1994d: 80 (as synonym of M. sabuleti); as synonym
of M. scabrinodis: syn. nov.
Myrmica scabrinodis var. sancta:.Karawajew 1934: 77; Sadil 1952: 249; not confirmed, see Notes to
M. specioides, below.

259
Myrmica specioides: Sadil 1952: 249; not confirmed, see Notes to M. specioides, below.
Myrmica georgica: Radchenko 1994d: 79; not confirmed, see Notes to M. georgica, above.

Type localities and type specimens. M. scabrinodis: no locality given. M. scabrinodis var. rugulo-
soides: "Torfmoore bei Roche". M. pilosiscapus: "Cette espèce semble répandue dans les régions sub-
alpines de l'Europe centrale: Belgique: Hautes-Fagnes; Allemagne; France: Jura с. с. (A. Vandel),
Pyrénées, Corse (E. Cordier); Suisse; Grande-Bretagne (Donisthorpe)... J'ai examiné plusieurs cen-
taines d'exemplaires provenant d'environ quarante nids" (Bondroit 1920a); "Pyrénées-Orientales:
Font-Romeu, vers 1.700 m.; Doubs: très commun, dans la région de Pontarlier, entre 800 et 1.000 m.
(A. Vandel) (Bondroit 1920b). M. scabrinodis var. ahngeri: "Taganrog, 8.VI. 1926, К. Ahnger, 2 w".
M. rolandi var. reticulata: "Pyrénées centrales, Cirque d'Espingo, entre 1850 et 2000 m., août 1929, w,
q, m (types). - Val Astos, Commune d'Oô, entre 1100 et 1200 m., août. -Val de la Frèche, fond de la
vallée de la Pique, entre 1550 et 1650 m, Haute-Garonne, sous des pierres (H. Gadeau de Kerville
leg.)". M. scabrinodis var. scabrinodosabuleti: "From widely different places in Czechoslovakia. As
examples I give the following finds: Nova Rise near Tele 6-7-1938 (Zálesky); Pribram 1936 (Sipek);
Brotin 28-8-39 (Novak)".
Material examined. Lectotype of M. scabrinodis (designated by Radchenko 2007a), w (bottom
specimen on the pin with 2 w), "Kuusamo", "W. Nyland.", "Mus. Fenn." (HELSINKI); paralecto-
types: 1 w (upper on the pin with the lectotype); 2 w on one pin with the same labels as lectotype
(HELSINKI) (see also Notes below); lectotype of Ai. scabrinodis var. rugulosoides (designated here),
w (bottom specimen on the pin with 2 w), "Myrmica scabrinodis Nyl. v. rugulosoides For., type",
"Tourfiéres de Roche", "Typus" (GENEVA); paralectotypes: 1 w on the pin with lectotype; 3 w, "Myr-
mica scabrinodis Nyl. v. rugulosoides For., type", "Tourfiéres de Roche", "Col. A. Forel", "Typus"
(GENEVA) (see also Notes below); lectotype of M. pilosiscapus (designated here), w (upper specimen
on the pin with 2 w, 2 q and 2 m), "Monte Jura, nid dans sphagnum, humide" (BRUSSELS); paralec-
totypes: > 100 w, > 50 q, > 50 m from Belgium, France, Corsica, Germany and England (BRUSSELS);
1 m, "Myrmica pilosiscapa Bondr., type Monte Jura francés" (GENOA); lectotype of M. scabrinodis
var. ahngeri (designated by Arnoldi 1970), "Таганрог, 31.V.1926 К. Ahnger" [Taganrog, 31.V.1926 К.
Ahnger], "Myrmica scabrinodis var. ahngeri nova typus Karawajew", "Myrmica scabrinodis ahngeri
Karaw. K. Arnoldi det. lectotyp.", "Myrmica specioides Bondr. Det. Seifert 1988" (MOSCOW); para-
lectotype: 1 w, "Таганрог, 8.VI. 1926 K. Ahnger" [Taganrog, 8.vi. 1926 K. Ahnger], "Myrmica scabrin­
odis var. ahngeri nova typus Karawajew" (KIEV); lectotype of M. rolandi var. reticulata (designated
here), m (upper specimen on the pin with 3 m and 1 q), "M. scabrinodis rolandi Bondr. v. reticulata
Sant.", "[very hardly readable text, most probably: M° Espigno, Val d'Oueu, Ha Garon, 1200 m,
Kerville, 20.VIII.29]", "Sammlung Dr. F. Santschi Kairouan", "= M. scabrinodis Nyl." (written by
С. A. Collingwood) (BASLE); paralectotypes: 2 m, 1 q on the pin with lectotype; 3 w (on the pin with
8 w; see also Notes below), "M° Espigno, Val d'Oueu, Ha Garon, 1200 m, Kerville, 20.VIII.29", "Samm­
lung Dr. F. Santschi Kairouan" (BASLE); syntypes of M. scabrinodis var. scabrinodosabuleti: 1 w,
"Myrmica scabrinodis", "24.VIII.39 Borotin, Bohemia т., leg V. Novak", "Mus. Nat. Pragae Inv.
3404", "M. se. var. scabrinodosabuleti Sadil (det. 1951)" (PRAGUE); 5 w, "Bohemia centr., Sarka ad
Praha, 2.IX.1948, leg. J. Sadil", "M. scabrinodis var. scabrinodosabuleti Sadil det. J. Sadil", "Paraty-
pus", "Inst. Zool. P.A.N. Warszawa 102/17" (WARSAW); non-type material: > 1000 w, several hun­
dred of q and m from the whole range.
Distribution (Map 103). Europe, Caucasus, Asia Minor, northern Kazakhstan, West
Siberia; the easternmost confirmed localities - vicinity of Krasnoyarsk (ca. 93°E) and 100
km N of Kansk (ca. 96°E) (material in MOSCOW, examined).
Etymology. M. scabrinodis: from a combination of the Latin words scabres = rough­
ness or "scabby" and nodus = knot or lump, to describe the rough, scabby appearance of
the surfaces of the petiole. M. rugulosoides: from a combination of the name rugulosa and
the adjectival suffix -oides from the Greek OEIÔEÇ = like or resemble, to indicate this form
somewhat resembles M. rugulosa. M. pilosiscapus: from a combination of the Latin words

2ÓO
pilosus = hairy and scapus = stalk or stem [scape], to indicate the hairy scape compared
to M. scabrinodis. M. ahngeri: named for the collector, К. Angher. M. reticulata: from the
Latin word reticulata = netted or reticulated, to describe the head of the male which was
"more reticulated than striated". M. scabrinodosabuleti: named to indicate that its features
were intermediate between M. scabrinodis and M. sabuleti.
Notes. M. scabrinodis typifies the scabrinodis species group and belongs to the scabri-
nodis-complex of that group. For many decades after its description it was one of the "basic"
Myrmica species to which many infraspecific forms were attributed, but several authors
[possibly influenced by Curtis' (1854) treatment - see notes to M. rubra] considered it to
be a variety of M. rubra (e.g. Forel 1874; Emery and Forel 1879; Emery 1895a; Forel 1904a;
Wheeler W.M. 1908b).
Many "old" authors made a variety of different sorts of mistakes, determining different
castes of M. scabrinodis as M. sabuleti or M. specioides (e.g. Forel 1915; Bondroit 1920a)
or vice versa (e.g. see Emery 1920), or including in "M. scabrinodis" several different
species (Ruzsky 1905, etc.). Such errors are not surprising especially when only the
workers are available for identification. M. scabrinodis is so widespread and common that
specimens are only given a superficial examination and even when workers are examined
more carefully it is easy to confuse M. scabrinodis with several other scabrinodis-group
species. Rarer scabrinodis-group species are often identified as M. scabrinodis (sometimes
vice-versa) but most commonly workers (and sometimes queens) are confused with those
of M. vandeli and M. specioides, and occasionally with sympatric M. sabuleti populations.
However, M. scabrinodis males have a much shorter scape than those of M. sabuleti and
M. vandeli, and much longer erect hairs on the scape and tibiae than M. specioides, thus if
males are taken with the female castes all scabrinodis-group species currently recognised,
should be identifiable correctly (see Table 2.2 in Chapter 2.2). Thus M. scabrinodis has al­
ways been a "dust-bin" species that includes all forms that can not be clearly discriminated
by a combination of features of all three castes. Even now, we suppose that "M. scabrinodis"

Map 103. Distribution of M. scabrinodis.

261
includes at least two (perhaps more) cryptic species that are morphologically indistin-
guishable but have different ecological preferences and are adapted to quite different habi-
tats (see also Ecology, below).
We studied the type series of M. scabrinodis, preserved in the Nylander's collection
(HELSINKI) and designated the worker as the lectotype (see material examined).
Originally, the lectotype worker was pinned through its gaster by a thin and blunt pin;
during our investigation the specimen fell from this pin so we glued the lectotype to
a cardboard triangle on a proper entomological pin.
Apart from the lectotype and paralectotypes we also found in Nylander's collection some
other specimens originally determined as M. scabrinodis. Firstly, a pin with only a gaster
(the main part of the specimen having fallen off and been lost at sometime), labelled
"a Dahlbom", "Mus. Zool. H: fors Spec. typ. No. 5049 Myrmica scabrinodis Nyl.", "Mus.
Heis. N:o 3217"; almost certainly this specimen was part of the type series, but there seems
no reason include this ant gaster to the paralectotypes. Secondly, 1 male (without head) and
1 queen on a pin labelled "Bogdâ", "W. Nyland.", "Coll. Nyland.", "Emsale", "20/viii.46,
f m", "Mus. Hels. N:o2212", "Mus. Hels. N:o2216"; despite having no head the other fea-
tures, such as the very long and abundant erect hairs on the legs, fit well with the charac-
ters of M. scabrinodis males. Thirdly, 1 queen labelled "H: fors", "W. Nyland.", "2.x 47",
"Coll. Nyland.". All these specimens are indubitably M. scabrinodis but formally they
can not be included to the type series because they were collected (August 1846 and Octo-
ber 1847) after Nylander's (1846a) paper was accepted (9th of February 1846). Final-
ly a worker with the labels "H: fors", "W. Nyland.", "Coll. Nyland." was a specimen of
M. sabuleti.
Additionally to the lectotype and paralectotypes workers of M. scabrinodis var. rugulo-
soides held in GENEVA (see above), we found "type" material in BASLE (2 workers, la-
belled "Suisse Roche Forel", "M. rugulosoides Forel type", "M. rugulosoides For. Sümppe
von Roche 24.VIII. 19", "2", "Sammlung Dr. F. Santschi Kairouan") that we identified as
M. scabrinodis, and in WARSAW (1 worker and 1 male, labelled "M. rugulosoides For.
Sümppe von Roche, 24.VII.19, Kutter", "Inst. Zool. P.A.S. Warszawa 90/66", "prep. 468",
"Cotypus") that we also identified as M. scabrinodis. These specimens do not belong to the
type series of M. scabrinodis var. rugulosoides because they were collected in 1919 and this
form was described only from workers in 1915. We found the lectotype and paralectotype
workers of var. rugulosoides to be indistinguishable morphologically from M. scabrinodis;
they have a somewhat smaller lobe at the scape base and somewhat less coarse sculpture
on the alitrunk and waist compared to "typical" M. scabrinodis workers, but the difference
is too minor for the separation of species and we confirm the synonymy.
During the last 50 years the separation of M. scabrinodis and M. rugulosoides has been
considered to be mainly ecological (e.g. Petal 1963; Baroni Urbani 1971; Kutter 1977); the
general feeling being that "bog scabrinodis" should be called M. rugulosoides. Seifert (1984)
compared biometrically two "ecomorphs" of M. scabrinodis: those collected in peat bogs
(the putative M. rugulosoides) with sympatric populations collected in adjacent drier habi-
tats. He found no morphological or morphometrical differences between these two ecotypes
and so confirmed synonymy of M. rugulosoides with M. scabrinodis. Of course, it can be ar-
gued that if M. rugulosoides are not "bog scabrinodis" then Seifert's results apply only to
the ecotypes of M. scabrinodis rather than M. rugulosoides.
We examined the type series of M. pilosiscapus, comprising more than 100 workers,
more than 50 queens, and more than 50 males in the Bondroit collection (BRUSSELS) that

2б2
were taken from Belgium, France, Corsica, Germany and England. We designated as the lec-
totype a worker from Jura (see above). There is no doubt that all these specimens are
M. scabrinodis and we confirm the synonymy. At first sight it is puzzling as to why Bondroit,
who described many "good" European ant species, should have described M. pilosiscapus.
The problem seems to have arisen because he included under the name "scabrinodis"
M. sabuleti (this is especially obvious from the male characteristics, given in his publica-
tions; see Bondroit 1918,1920a, b) and therefore as "real" M. scabrinodis is common every-
where and differed from his concept ofthat species he described M. pilosiscapus.
Var. reticulata was originally described by Santschi (1931b) as scabrinodis st. rolandi
var. reticulata and considered till now as unavailable name (Bolton 1995; Bolton et al.
2006). However, Stärcke (1942) used the first available name M. rolandi var. reticulata for
this taxon. The type specimens are preserved in BASLE: 3 males and 1 queen are
M. scabrinodis but among 8 workers on one pin there are 3 M. scabrinodis and 5 M. sa-
buleti. This form was already synonymised with M. scabrinodis (Seifert 1988; Radchenko
1994d) so we confirmed the synonymy and to maintain stability in the nomenclature we
designated as the lectotype a male that is indisputably M. scabrinodis (see above).
Sadil (1952) described M. scabrinodis var. scabrinodosabuleti from various localities of
Czechoslovakia based on workers and males. He stressed, that it combines characters of
M. scabrinodis and M. sabuleti, particularly, males have short scape (even shorter than in
"typical" M. scabrinodis), but some workers can be determined as M. scabrinodis, while
others - as M. sabuleti. He also suspected that it might be a hybrid form of these two species,
an idea partly supported by the observation of multi-species mating swarms (see Chapter
1.3.5). We investigated 6 workers that we believe belong to the Sadil's type series of var.
scabrinodosabuleti (see above), but unfortunately, Sadil rarely indicated type material in his
collection by special labels. Three of the specimens seem like M. scabrinodis (with FI 0.35-
0.37) but three others have features (the shape of the lobe at the scape base, the narrower
frons, etc.) typical for M. sabuleti (FI 0.32-0.34). Based on these workers var. scabrinodo-
sabuleti could be synonymised with either species and indeed, Radchenko (1994d) syno-
nymised this variety with M. sabuleti. However, the features of the males as Sadil originally
described them conform more to M. scabrinodis, therefore we now synonymise var. scabri-
nodosabuleti with M. scabrinodis. Theoretically, Sadil might have got his hybrid ideas if he
originally found a nest of M. tulinae (M. sabuleti-likе workers and M. scabrinodis-like
males) in which case his var. scabrinodosabuleti could even be a senior synonym of
M. tulinae; however his collection lends no great support to this possibility so in our
opinion it is best to consider var. scabrinodosabuleti as junior synonym of M. scabrinodis.
Myrmica scabrinodis var. ahngeri was described by Karawajew (1926a) from 2 workers
collected by K. Ahnger, taken near Taganrog (Rostov Province of Russia). Later, Arnoldi
(1970) treated it as "steppe" subspecies of M. scabrinodis, and described males (in a Key):
"Male: scape and tibiae with not long and not abundant standing hairs" (loc. cit., p. 1837).
Perhaps, based mainly on the male characters given by Arnoldi, Seifert (1988) synonymised
var. ahngeri with M. specioides. We studied the lectotype and paralectotype workers of var.
ahngeri and additional workers and queens collected by Ahnger from Taganrog (KIEV) but
unfortunately not included to the type series by Karawajew. We found them to be morpho-
metrically indistinguishable from M. scabrinodis, with the only feature reminiscent of
M. specioides being the weakly developed dorsal plate of the petiole; so we confirm the
synonymy of Radchenko (1994d). The males ascribed to the subsp. ahngeri by Arnoldi
(1970), almost certainly belong to another species (probably M. specioides).

263
 We consider that the earlier established synonymy with M. scabrinodis, of vars. sancta,
M. specioides and M. georgica was incorrect (see Karawajew 1934; Sadil 1952; Radchenko
1994d).
Ecology. Myrmica scabrinodis uses a wide variety of habitats throughout is extensive
range, it is most commonly associated with grasslands but is also present in open forests and
woodlands, especially in the southern part of its range. In forests it nests in the soil, often
under bark and pieces of wood and sometimes in and under moss. In bogs and marshes
M. scabrinodis builds nests in grass and moss tussocks and usually colonies build a quite
large solarium from soil, chewed moss and other detritus, where they rear their larvae. In
drier grassland and on high moorlands, nests are built under flat stones or directly into the
soil, in the latter case small solaria are built in spring but these are not maintained as the
season progresses. In very hot conditions nests are built in the soil with no obvious above-
ground structures. In many hotter habitats that support a population of Lasius flavus
mounds, nests of M. scabrinodis can be found built into the side of the mound and there is
some evidence that they prey upon the L. flavus larvae, a view supported by the results of
Pontin (1969) who showed that L. flavus produced more sexuais when the nests of
M. scabrinodis were removed. M. scabrinodis colonies, like M. rubra, have been shown to
amalgamate under some conditions (Morely 1938). A cast of a M. scabrinodis nest on heath-
land was made by Brian and Downing (1958) and excavation behaviour was studied by
Sudd (1971).
As has been pointed out in the notes above, it is hard to believe that bog-living
M. scabrinodis can be the same species as sympatric populations living in very hot, dry gras-
sland. For example, in one study (Elmes et al. 1994) we showed that on sites in both Spain
and the Netherlands there appeared to be two distinct "types" of M. scabrinodis that were
easily discriminated in the field. "Type 1" appeared smaller and paler, lived in wetter con-
ditions, made extensive nests in the vegetation with few if any deep chambers, were more
timid being reluctant to run up ones hand if placed in the nest and attacking baits from un-
derneath (like M. lobicornis); "type 2" appeared slightly larger and darker, lived in some-
what drier conditions, made nests with deep chambers in the soil and were more inclined
to run up ones hand when placed into the nest, and sat on top of baits (more like M. rugi-
nodis). We also noted similar ecological differences in populations from France and else-
where in Europe (unpublished).
Subsequently we identified the "type 1" specimens from Spain as M. aloba, although
these were somewhat smaller than is typical for that species especially compared to popu-
lations from Southern Spain and Portugal. Elsewhere, no consistent differences between the
two types could be found when dead samples were compared in the laboratory; all the dif-
ferences seemed to be simply behavioural. Seifert (1984) made a careful morphometric com-
parison in the context of M. rugulosoides (see above) and Sandor Csösz (pers, comm.)
made a similar study using material from a wider range, both found no discernable mor-
phological differences. However, some unpublished preliminary studies made by a Danish
Group indicate that such populations have some genetical differences that might be suffi-
cient to indicate cryptic species. In our view M. scabrinodis is undergoing a major specia-
tion event at the current time largely in response to anthropogenically altered habitats (and
probably also related species, such as M. aloba). It seems to us that over its range it might
comprise several (perhaps four or five) cryptic species and in any region perhaps two or
maybe three of these live sympatrically being separated by their behaviour and ecology.
The problem is then that if an ecologist in say Britain recognises two ecological forms and

264
another ecologist in say Hungary recognises two ecological forms there is no reason to sup-
pose that the British pair are the same genetically as the Hungarian pair. This is a fascinating
problem that can not be resolved by normal taxonomic study, only by appropriate molecu-
lar genetical studies.
On an acid grassland site M. scabrinodis colonies average about 200 workers with 1.4
queens, 30% of nests produced sexuais and these were larger colonies with fewer queens
(Elmes and Wardlaw 1982c). Like the sympatricM.sabuleti populations colony size and pro-
ductivity was related to the above ground vegetation (Elmes and Wardlaw 1982a). In Eng-
land, colonies on limestone grassland were the largest averaging > 4 queens and about 850
workers per nest, on acid grassland numbers were about 2 queens and 600 workers while
high, cold moorland had the smallest colonies, < 0.5 queens and about 300 workers per nest;
nest densities were about 1.0, 0.25 and 0.004 nests per m2 respectively (Elmes and Wardlaw
1982b). Genetical relatedness and colony structure was investigated by Seppä (1996).
The mandibular secretions of M. scabrinodis and other ants have been shown to inhibit
pollen germination (Sanderson and Wright 1989) and M. scabrinodis have been shown to
rob insect prey from Drosera species (Thum 1989), which may be important for the ants
when insect prey is in short supply. The general role of M. scabrinodis in moorland ant
communities was studied intensively in Scotland (e.g. Brian and Brian 1951; Brian 1952a,
b, 1955b, 1956b), in North Sea sand-dunes (Boomsma and De Vries 1980; Boomsma and
Van Loon 1982; Boomsma and Isaaks 1982), in a mountain meadow (Woyciechowski and
Miszta 1976), in steppe (Reznikova and Samoshilova 1981) and in relation to Formica ants
(Vepsälänen and Savolainen 1990; Dauber and Wolters 2000), but this species has never
been subjected to the same intensive study as say, M. rubra and M. ruginodis.
There have been more laboratory studies, comparable with those on the other wide-
spread Myrmica species. For example, pheromones, glandular secretions and cuticular hy-
drocarbons have been identified (Cammaerts et al. 1978; Morgan et al. 1978, 1979; Elmes
et al. 2002), foraging behaviour described (Dlussky et al. 1978; Cammaerts 1980b; De Vroey
1980a, b). Age related polyethism was demonstrated for M. scabrinodis (Weir 1958a, b)
and the solicitation behaviour for trophollaxis was described (Lenoir 1972a, b). M. scabri-
nodis workers and larvae appear physiologically adapted to moderately cool conditions, at
any temperature they grow at a rate intermediate between that of M. rubra and M. sabuleti
(Elmes and Wardlaw 1983a). This accords to species-differences in basal respiration which
vary according to the latitude at which the population lives (Nielsen et al. 1999);
respiration of M. scabrinodis larvae were compared with those of Lasius flavus with which
it coexists in many habitats (Peakin et al. 1985). While queens of M. scabrinodis suppress
the development of large gyne-potential larvae in the same way as other Myrmica species
(Elmes and Wardlaw 1983b) some gynes can develop from small over-wintered larvae and
so escape queen-control (Elmes and Wardlaw 1981), which might explain why the impact
of queen and worker numbers on individual size in natural colonies of M. scabrinodis can
not be detected like in the cases of M. rubra and M. sabuleti (Elmes 1974b). Also, different
populations of M. scabrinodis appear more variable in the ability of the workers to oviposit
(Wardlaw and Elmes 1998).
Finally, like all the other European Myrmica species nuptial flights take place from late
July until the end of September (e.g. Donisthorpe 1934; Boomsma and Leusink 1981; Woy-
ciechowski 1990a; our observations). Very often males and gynes join large mixed mating
swarms but we have also observed small very local swarms composed entirely of M. scabri-
nodis. These local swarms often use a small bush or shrub as a swarm focus.

265
 Myrmica schencki Viereck, 1903
(Figs 242-244)

Myrmica rubra var. schencki Viereck, 1903: 72 (first available use of Myrmica rubra subsp. scabri-
nodis var. schencki Emery, 1895a: 315, w, q, m, Europe) (see also Baroni Urbani 1971: 35, and
Notes below)
Myrmica scabrinodis lobicornis var. schencki: Ruzsky 1905: 699 (unavailable name).
Myrmica scabrinodis subsp. (st., var.) schencki: Emery 1908a: 178; Forel 1915: 29; Emery 1916: 120,
1921: 41; Soudek 1922: 39, nec Stitz 1934: 2.
Myrmica schencki: Bondroit 1911: 11, 1912: 351; Donisthorpe 1915b: 265; Bondroit 1918:103; Müller
G. 1923: 44; Finzi 1926: 109; Karawajew 1926b: 95; Stärcke 1927: 84; Karawajew 1927b: 259;
Santschi 1931: 351; Karawajew 1934: 93; Arnoldi 1934: 171; Stitz 1939: 102; Weber 1948: 296;
Stärcke 1942: 25; Sadil 1952: 260; Wheeler G. С and Wheeler]. 1953: 120; Collingwood 1958b:
74; Bernard 1967: 118; Arnoldi 1970: 1843; Baroni Urbani 1971: 35; Arnoldi 1976: 557; Tarbin­
sky 1976: 42; Kutter 1977: 70; Arnoldi, Dlussky 1978: 535; Collingwood 1979: 56; Espadaler
1981b: 189; Seifert 1988: 35; Atanassov, Dlussky 1992: 99; Seifert 1994: 15; Radchenko 1994b:
44, 1994c: 140, 1994e: 77, 1994g: 107; Bolton 1995: 283; Seifert 1996: 234; Radchenko et al. 1997:
492; Czechowski et al. 2002: 32; Seifert 2003: 145; Radchenko, Czechowska and Czechowski 2004:
46; Radchenko, Elmes and Alicata 2006: 513; Seifert 2007: 210; Radchenko 2007a: 29, nec Gee
1924: 102; Kupyanskaya 1986a: 87; 1990: 109, misidentifications.
Myrmica schencki var. kutteri Finzi, 1926: 111, w, Switzerland; Weber 1948: 3 0 1 . Synonymy by
Bernard 1967: 118; Radchenko 1994e: 77; Bolton 1995: 280; Seifert 2003: 145; Radchenko, Elmes
and Alicata 2006: 513; confirmed here.
Myrmica schencki nat. subopaca Arnoldi, 1934: 172, w, Ukraine (also described as new by Arnoldi, in
Karawajew 1934: 96) [unresolved junior primary homonym of Myrmica subopaca F. Smith, 1858:
127 (transferred to Monomorium)}; Weber 1948: 304. Synonymy by Arnoldi 1970: 1843; Rad­
chenko 1994e: 77; Bolton 1995: 284; Seifert 2003: 145; Radchenko, Elmes and Alicata 2006: 513;
confirmed here.
Myrmica betuliana Ruzsky, 1946: 70, w, Russia. Synonymy by Radchenko 1994e: 77; Radchenko,
Elmes and Alicata 2006: 513; confirmed here.
Myrmica schenckioides Boer et Noordijk, 2005: 120, q, Netherlands; Radchenko, Elmes and Alicata
2006: 527, syn. nov.
Myrmica schencki var. obscura: Bernard 1967: 118; Radchenko 1994e: 77; Bolton 1995: 281; Seifert
2003: 145; Radchenko, Elmes and Alicata 2006: 513; not confirmed, see Notes to M. obscura, above.

Type localities and type specimens. M. schencki: "In Europe scheint diese Ameise im Flachland ver­
breitet zu sein. Schenk beschrieb das charakteristische [m] zuerst aus Nassau. Ich erhielt [w, q, m] aus
Limburg von Herrn Wasmann; derselbe schickte mir auch [w] von Prag, sowie einen [w] und ein [q]
von Aachen aus Förster's Sammlung" (American type material, referred to this species, do not belong
toM. schencki, see Notes below). M. schencki var. kutteri: "Zermatt (Svizzera- Kutter)". M. schencki
nat. subopaca: "Besiedelt hohe Steppen des Donetzbassins, wo sie in sehr grossen Mengen von Med-
vedev gefunden ist (Provalje)" (Arnoldi 1934); "Знаходищс: Провалля (Красная Могила) в Донсцькому
басейш, С. Медведев (№ 5632), 2 w" [Locality: Provalla (Krasnaya Mogila) in Donets Basin, S. Med-
vedev (No 5632), 2 w] (in Karawajew 1934). M. betuliana: "Бараба, ст. Тайга; Богородское на Оби,
Всршинпно, Басандайка (VI—VII—VIII 1939-1941), березовые колки" [Baraba, station Taiga; Bogorods-
koe on Ob' river, Vershinino, Basandaika (VI—VII—VIII 1939-1941), birch coppices]. M. schenckioides:
"Holotype: female. Type locality: Beekhuizerland, Harderwijk, Gelderland, The Netherlands, 52°20'N,
5°40'E. Out of a pitfall, 12.V.2004-14.X.2004"
Material examined. Lectotype of M. schencki (designated by Seifert 2003), w (upper specimen on
the pin with 2 w), "M. scabrinodis var. Schencki Emery (lobicornis Schencki)", "Myrmica lobicornis",
"Ex alen", "Museo Civico di Genova", "Lectotype [top specimen] Myrmica rubra scabrinodis var.

266
schencki Emery, 1895 designated by Seifert" (GENOA); paralectotypes: 1 w (bottom specimen) on the
pin with lectotype; 1 w, 1 q (damaged, gaster and waist stuck by glue separately), "w $ i.86 Ex", "Para-
lectotypus var. schencki Emery, 1895"; 1 w, 1 m (damaged, gaster and waist stuck by glue separately),
"S w Ex", "Paralectotypus var. schencki Emery, 1895"; 2 w, "Ex alen", "Paralectotype des. Rad­
chenko"; 1 w, 1 m, "Ex 87", "Paralectotype des. Radchenko" (GENOA) (see also Notes below); syn­
types of M. schencki var. kutteri, 3 w, "Cotypus", "Zermatt с. 1750 m VII.17" "M. schencki var. kutteri
Finzi" (HARVARD); syntypes of M. schencki nat. subopaca, 19 w, "Провалъс, Донец, VI.928,
Медведев" [Provalie, Donets, vi.928, leg. Medvedev], Nos 3687, 3688, 3691, 3692, 3693, 3694, 3695;
3665 (same locality, 20.vi.928); 3670 (same locality, 18.vi.928); 3671 (same locality, 19.V.928)
(MOSCOW); 2 w, "Провальс, Донецк., VII.31, С. Медведев" [Provalie, Donets, vii.31, leg. S.
Medvedev], "5432 Coll. Karawajewi", "Myrmica schencki Em. n. subopaca Arnoldi typ.", "Syntypus
Myrmica schencki subopaca K. Arnoldi" (KIEV); non-type material: > 1000 w, > 100 q, > 100 m
from the whole range.
Distribution (Map 104). Europe (to the north until central England and Ireland, south
of Norway, Sweden and Finland; to the south until north of Spain and Italy, Balkans), Cau­
casus, north-eastern Turkey, south of West Siberia, northern Kazakhstan, Tien-Shan, Altai
Mts; the easternmost known locality - vicinity of Krasnoyarsk (East Siberia, material in
MOSCOW, examined).
Etymology. M. schencki: named for the notable 19th century German myrmecologist,
Prof. Carl Friedrich Schenck (1803-1878) of Weilberg, who first recognised this form (see
Notes below). M. kutteri: named for the outstanding Swiss myrmecologist Heinrich Kutter
(1896-1990), although a pharmacologist by profession he made a major contribution to the
study of European ants. M. subopaca: name derived from a combination of the Latin words
sub = under or in some little degree, and opaca = dark or shady, to indicate its generally
darker but not too dark colour compared with M. schencki. M. betuliana: name derived
from a combination of the Latin words betula = birch tree and the adjectival suffix
ana = connection with or close to, to indicate the species is associated with birch woods.
M. schenckioides: from a combination of the name schencki and the adjectival suffix-o¿des
from the Greek ostSsç = like or resemble, to indicate it resembles M. schencki.

Map 104. Distribution of M. schencki.

267
 Notes. M. schencki typifies the schencki species group. Its workers are characterized by
the presence of a big vertical lobe at the scape base, by the well-developed petiolar pedun­
cle and not-angled petiolar node (seen in profile) and especially, by the very narrow frons
(mean FI 0.23) ; after M. ravasinii, it has the second narrowest frons among all known Myr­
mica species in the World.
Emery described Myrmica rubra subsp. scabrinodis var. schencki in 1895: in that paper
he said that the male was first described by Schenck (1852) (under the nameM. lobicornis)
from Nassau, in the Rhineland of Germany, which is probably why he named the variety
after Schenck. Subsequently, almost all authors ascribed the authorship of M. schencki to
Emery but based on the paragraphs 10.2, 45.5.1 and 50.3.1 of the 4 th edition of the Inter­
national Code of Zoological Nomenclature (1999), the author of M. schencki was Viereck,
who used the first available name for this taxon, This rule creates a strange situation:
Viereck (1903: 72) was listing the ants of New Mexico, USA and the almost black Myrmica
specimens that he referred to M. schencki Emery (1895a) can not be the same species that
Emery described, although they are undoubtedly one of the several American schencki-Ше
species which dominate the Myrmica fauna of the southern states.
A second more significant, taxonomic problem is that Emery's (1895a) original de­
scription of var. schencki was in a paper dealing with American fauna, and as the type lo­
calities he gave first Main, New Jersey, New Hampshire and Carolina. However, he also
ascribed var. schencki to specimens in various European collections taken from Nassau,
Limburg, Prague and Aachen, and noted that "In Europe this species seems distributed on
the plains" (loc. cit., p. 315; see also Type localities, above). In hindsight, it seems that
Emery placed importance on the short male scape separating M. schencki from M. lobicor­
nis, noting that Mayr followed Schenck's view of this in some publications and not in
others. Emery (loc. cit.) discussed the morphological features of M. schencki based mostly
on the European material, and compared it with several European but not American species.
We suspect therefore, that in his mind Emery recognised the name var. schencki for the
European series (which surely were M. schencki as recognised today) and because he was
writing on North American ants he included to the types schencki-ìike specimens that had
males with short scape.
We have examined thousands of Myrmica specimens from many hundreds of nest sam-
ples from diverse localities in North America (including extremely rich material collected
recently by G. Jansen) and have never found a specimen of "European" M. schencki. After
Forel (1914) gave the name Myrmica scabrinodis r. schencki var. emeryana (first available
use of this name - Cole, 1942) to a schencki-like ant from northeast USA, American
schencki-ìike species have always been identified as M. emeryana (or as other related
American species). Emery's North American material were most probably M. emeryana,
but could have been a mixture of several American species. Therefore we absolutely agree
with Baroni Urbani's (1971: 35) opinion that the North American specimens belong to
separate species, and the name "schencki" should be referred only to the Palaearctic
populations, as has been traditionally done for about the last 100 years.
To prevent any future speculations and to maintain stability in the nomenclature, Seifert
(2003) rightly designated a lectotype (worker, GENOA) of M. schencki from the material
that he felt fully corresponded with the characteristics of West Palaearctic M. schencki as
recognised by many authors for more than 70 years. We also examined the lectotype
and paralectotype series in GENOA (totally 7 workers, 1 queen and 2 males, see above).
There is no doubt that they fully correspond to the modem treatment of M. schencki, but

268
unfortunately, all these specimens have quite illegible locality labels. Seifert (2003) has read
the label on the pin with lectotype as "Graten" (we could not find this putative locality
in Europe or anywhere else). However, after careful re-examination we read two of the
labels (including that of the lectotype) as "Ex alen" (see Fig. 245). So, these might not be
locality labels but simply indicate the source of material ("Ex" = from). Nevertheless, all the
type specimens clearly were part of Forel's collection, being stuck to cardboard triangles
with cut tips that are very characteristic of Forel, and as they are typical of European ma-
terial the designated type series of M. schencki is fully acceptable to us.

Figure 245. Original labels of lectotype worker (A) and paralectotype male (B) of M. schencki.

- M. schencki is fairly stable morphologically. It most resembles M. koreana,

M. caucasicola and M. obscura, but differs from them by its narrower frons. Also the males
of all these forms are fairly well distinguishable (see also Radchenko, Elmes and- Alicata
2006; Elmes et al. 2009, and a Key, Chapter 5.1, 5.2). In Europe, M. schencki quite often
was confused with M. lobicornis, though workers of the latter have the wider frons
(FI > 0.27), shorter propodeal spines, and the short petiole with weakly developed
peduncle and almost straight, quite steep anterior face that meets with the dorsal one at
a sharp right or even acute angle. When males are available such confusions should not
occur because M. lobicornis males have a much longer scape.

269
 Eight infraspecific forms of M. schencki have been described from the Palaearctic Re-
gion. Three of them (M. schencki var. brunescens from Caucasus, M. schencki var. burt-
shakabramovitshi from Ukraine, and M. schencki var. starki from northern Russia, Kola
Peninsula) are now considered as synonyms of M. lobicornis (see Notes to M. lobicornis,
above). M. lobicornis var. plana from Ukraine is considered in this monograph as a synonym
of M. deplanata (see above), while M. caucasicola and M. obscura are now treated as a good
species (see corresponding Notes, above).
Only two forms (M. schencki var. kutteri and M. schencki nat. subopaca) remain
synonyms of M. schencki. We investigated the syntype specimens of var. kutteri
(3 workers) and of nat. subopaca (22 workers) (see above), but did not find any essential
differences (including morphometries) between them and "typical" M. schencki. We there-
fore confirm the earlier synonymy.
Unfortunately, the types of M. betuliana are probably lost. This species was described
from workers collected in West Siberia (in modern Novosibirsk and Tomsk Provinces and
Altai Region), where it inhabited birch coppices. The main diagnostic features of this species
noted by Ruzsky (1946: 70-71) are (our translation from Russian): "Resembles lobicornis
(by the shape of scape); by the shape of alitrunk and waist it is similar to M. ruginodis.
Scape at the base curved, almost right-angled, with a wide shovel-like lobe (which is big and
well defined)... Propodeal spines long, thin, straight and acute; ... metanotal groove weak.
Petiolar and postpetiolar dorsum ovally rounded, not shiny, submat, with fine longitudinal
reticulation and rugulosity. Head and alitrunk with quite coarse longitudinal rugulosity...
Colour rather light, brownish-reddish-yellow or brownish-yellow. 5-5.5 mm" . In our opin-
ion, all the main diagnostic features of Ruzsky's description (except perhaps only weakly
developed metanotal groove) conform to those of M. schencki. The only other candidates
for its identity are M. koreana from which it differs by its "wide shovel-like lobe", and
M. lobicornis, which apart from the lobe shape has quite another shape of petiole. Based on
the description, the type locality and ecological data, as well as on other material collected
from the region of the type locality, Radchenko (1994e) synonymised M. betuliana with
M. schencki. We confirm this synonymy here.
Boer et Noordijk (2005) described from The Netherlands M. schenckioides based on
a single unusual winged gyne, found in a pitfall trap. By the size and by the shape of its
head, frons and scape it corresponds to normal gynes of M. schencki, but differs by having
very short, though sharp, propodeal spines, a "male-shaped" petiole and a short and high
postpetiole with a quite well developed ventral lobe (or rather just enlarged postpetiolar
sternum). Based on these differences, Boer et Noordijk assumed that it is a social parasite
of M. schencki which also lives in the type locality. Nevertheless, despite a very intensive
investigation of the biology of M. schencki, especially during last two decade when many
thousands of the nests have been dug-out throughout Europe (see ecology below), no one
has ever reported finding a socially-parasitic Myrmica species in a M. schencki nests. In
our opinion, this specimen M. schenckioides is an aberrant specimen of M. schencki pro-
duced either by some genetic malformation or by infestation by a Mermis parasite (e.g. see
Czechowski, Radchenko and Czechowska 2007). We therefore synonymise it with
M. schencki.
Stitz (1934) recorded M. schencki for the Sichuan Prov, of China. We have not
examined these specimens but the region is far out of the known range of this species,
almost certainly this record belongs to the recently described M. sinoschencki or possibly
some other undescribed schencki-Yuie species.

270
 Ecology. Throughout most of its northern distribution, M. schencki is associated with
hot, dry grassland habitats including calcareous grassland, glacial sand deposits and
seaboard sand-dunes. In its southern distribution it is associated with hot sub-alpine grass-
land. Much of its distribution seems to coincide with species of Tetramorium and
Tapinoma, which are dominant species in such habitats; we have often recorded and exca-
vated M. schencki nests from the middle of Tetramorium caespitum (L.) territories: T. cae-
spitum can defend discrete territories that contain several polycalic nest centres connected
by tunnels (e.g. see Brian and Elmes 1974). The common factor appears to be the need for
relatively high soil temperatures but not completely arid conditions. In biotopes where the
soil is potentially very arid, e.g. the sand-dunes of NW Europe, the sub-alpine pastures of
the Southern Alps and limestone pavement as in the Burren, Ireland, M. schencki only
occurs if there is a relatively high rainfall, which stops complete desiccation of the soils.
M. specioides is the only European Myrmica species that can tolerate similarly hot habi-
tats. In many places its main competitors are ants from other genera and we believe that this
contributes to its habit of building a small woven funnel at its nest entrance in which
a "guard-worker" sits with its head more or less blocking the entrance. Tarbinsky (1981)
showed that in the eastern part of its range M. schencki has rather limited foraging dis-
tances which generally agrees with our observations.
A population of M. schencki living on sandy soil In Jutland, Denmark has been studied
by several authors. Elmes and Abbott (1981) showed that colonies averaged about 1 queen
and 300 workers, and the overall mean head widths were 1.19 mm and 1.04 mm for queens
and workers respectively. The workers were generally bicoloured but in some colonies a pro-
portion of workers were a uniform gingery orange colour (not to be confused with the pale
yellow of callow workers) ; no queens of this colour were noted but a few queens were al-
most completely dusky black. Thus colour is quite variable in this species. On the same site
nest densities of 0.13 nests per m2 with an estimated 47 workers per m2 (Jensen 1976),
giving an estimated colony size of about 250 workers which accords well with the later es-
timate. This population was shown to have a distinct diurnal foraging activity and was most
active when soil surface temperatures were in the range 15-30°C, relative humidity did not
affect foraging activity (Nielsen 1981a), a finding confirmed at another site (Nielsen 1981b).
M. schencki has become important to nature conservation study in recent decades be-
cause it is a primary host of the larvae of the parasitic butterfly Phengaris rebeli (Hir.) (see
Chapter 1.3.6 and papers in Settele et al. 2005). Numerous nests have been excavated by
different teams from many parts of Europe, and the'general consensus is that the popula-
tion structure outlined above is typical. This work has led to the development of a spatially
explicit simulation model to describe the interaction between M. schencki and M. sabuleti,
its main Myrmica competitor on many P. rebeli sites. These models suggest that underly-
ing heterogeneity of "habitat quality" colony fission and the ability to pre-empt a nest site
is important to the spatial pattern of nest distribution (e.g. see Elmes et al. 1996; Clarke et
al. 1997 and references therein).
An important aspect of the interaction between M. schencki and the larvae of P. rebeli
is the ability of the caterpillars to mimic various ant signals. Consequently the cuticular
chemicals of the ants have been identified (e.g. Elmes et al. 2002) and how they interact with
caterpillar recognition (e.g. Elmes et al. 2004; Schrönrogge et al. 2004). One of the most in-
teresting discoveries was that P. rebeli has a species-specific parasitoid that can detect both
the M. schencki host ants and P. rebeli larvae by their odour, and itself has a suite of
chemicals that can provoke fighting in the host ants (e.g. Thomas and Elmes 1993; Thomas

271
et al. 2002). Recently the stridulations of queens and workers of M. schencki have been
recorded and were shown to be similar to the sounds made by P. rebeli caterpillars which
respond to the recorded ant sounds (Barbero et al. 2009). In the laboratory Stärcke (1949)
made quite an important contribution to the morphology of larvae. The chemistry of some
glandular secretions of M. schencki workers have been identified (e.g. Evershed et al. 1983;
Cammaerts et al. 1982; Attygalle et al. 1983). Workers have been shown to be quite prolific
egg-layers while queens regularly lay some trophic eggs (Wardlaw and Elmes 1995, 1998).
Mating swarms occur from the end of July to early September, though often slightly earlier
than in some sympatric Myrmica species, we have observed mixed swarms and a mating
swarm was described by Pontin (1986).

Myrmica schoedli Radchenko, Elmes et Viet, 2006

(Figs 246-248)

Myrmica schoedli Radchenko, Elmes et Viet, 2006: 36, w, q, Vietnam.

Type localities and type specimens. Vietnam, for details see material examined.
Material examined. Holotype, w, "Northern Cong Troi, Sa Pa district, Lao Cai province, 2000-
2200 m, 28.iv.2002, No. Eg02-VN-152 (GWE No.VN-3), leg. K. Eguchi" (LONDON); paratypes: 3
w, 1 q from the nest of the holotype; 1 w, same locality, but 2100-2200 m, 5.V.2001, GWE No. VN-4,
leg. K. Eguchi (ELMES, KIEV, VIET); non-type material: 55 w, 2 q, 1 m, Vietnam, Lao Cai, Sa Pa,
Cong Troi, 1850-2000 m К. Eguchi 11-13.Х.2006, leg. K. Eguchi.
Distribution (Map 105). Northern Vietnam.
Etymology. M. schoedli: this species was dedicated to the memory of the Austrian en­
tomologist and myrmecologist, Prof. Stefan Schöedl (1957-2005) of the natural History
Museum, Vienna.
Notes. M. schoedli belongs to the draco-complex of the ritae species group that includes
also M. draco and M. yamanei, from which M. schoedli well differs by its much longer scape
(SIj > 1.23 ws. < 1.18, SI2 > 1.45 ws. < 1.40). Additionally, the head dorsum of M. yamanei
has coarse reticulation (vs. longitudinal rugosity in M. schoedli), and the surface of the
propodeum is punctated, not smooth. M. draco has a distinctly shorter, not sub-oval head
(SIj < 1.10 vs. > 1.15), the less developed reticulation on the mesonotal dorsum, a deeper
metanotal groove and shorter petiole. As males were previously unknown, we describe them
below.
First description of male (Fig. 248). Head slightly longer than broad, with convex sides
and occipital margin, and gradually rounded occipital corners; anterior clypeal margin
very slightly convex and very shallowly notched medially. Central part of clypeus raised,
forming trapezoid lobe-like process when seen in profile. Frontal carinae very feebly curved.
Antennal scape short, somewhat shorter than 1st to 3 rd basal funicular segments together;
antennal club 5-segmented, but not well marked. Masticatory margin of mandibles distinct,
with 5 sharp teeth.
Alitrunk relatively long, low and narrow, scutum convex, and scutellum does not
project dorsally above scutum when seen in profile. Propodeum with blunt triangular
denticles. Propodeal lobes not prominent, but angled at the apex. Petiole long and low, with
distinct peduncle, with feebly convex and gradually rounded node dorsum; postpetiole also
quite long and low, seen from above fig-shaped.

272
 Map 105. Distribution of M. schoedli.

Cell 1 +2r of forewing, in contrast to that of known males of the ritae-group species, is
not wide, length / width ratio > 1.65 (see also Radchenko and Elmes 1998; Radchenko et
al. 2001).
Frons with fine striae, surface of head not coarsely but densely punctated, clypeus with
fine superficial microreticulation; whole surface appears shiny. Scutum behind Mayrian fur-
rows with a few longitudinal rugae, scutellum (except its central part) with longitudinally-
concentric rugulosity, propodeum with not coarse longitudinal rugosity, mesopleura distally
with fine rugulosity; surface of alitrunk smooth and appears shiny. Petiole and postpetiole
at most partly finely superficially punctate, mainly smooth and shiny. Gaster smooth and
shiny.
Head margins with not abundant but long standing hairs. Alitrunk, petiole, postpetiole
and gaster with similar, while somewhat shorter hairs. Legs and scape with short sub-
decumbent to decumbent hairs. Whole body reddish-brown, head somewhat darker, ap-
pendages reddish-yellow.
Ecology is poorly known, M. schoedli was found in well-developed secondary forest at
altitudes between 1850 and 2200 m and the nest was found under vegetation on a rock, by
the side of a stream.

Myrmica schulzi Radchenko et Elmes, 2009

(Fig. 249)

Myrmica schulzi Radchenko et Elmes, 2009a: 72, w, China.

Type localities and type specimens. South China, for details see material examined.
Material examined. Holotype, w, "China, Shaanxi Prov., Qin Ling Shan, Mt. W Pass, 47 km S
Xian, Autorote km 70, 108°47'E, 33°51'N, 2300-2600 m, No. 963, 26-29.viii.1995, leg. Wrase" (KIEV);
paratypes: 3 vv with the same label as the holotype (KIEV, ELMES, SCHULZ).

273
 Distribution (Map 106). It is known from the type locality in the Shaanxi province of
China.
Etymology. M. schulzi: this species was dedicated to the German myrmecologist
Andreas Schulz, who generously provided us with this and other material for our study.
Notes. M. schulzi belongs to the pachei species group which is now known to comprise
a considerable number of rather similar species in China that are mainly distinguished by
variations in body sculpture; for the separation of M. schulzi from related species see the
Key, Chapter 5.7. This species is characterized by the following combination of features:
lateral margins of head posterior to the eyes with abundant short decumbent hairs; the frontal
carinae curving outwards to merge with the rugae that surround the antennal sockets; the
frons with quite dense, fine, slightly sinuous longitudinal rugosity, numbering ca. 20 between
the frontal carinae level with the eyes, occiput and temples with fine reticulation, surface
between the rugae densely punctated, appears dull; the pronotal dorsum with irregular
sinuous rugulosity and reticulation, while the mesonotal and propodeal dorsum with not
coarse transversal sinuous rugosity, numbering > 25. Queens and males are unknown.
Ecology is unknown except that four workers were found on mountain mea-
dows at the altitudes between 2300 and 2600 m a.s.l.

Map 106. Distribution of M. schulzi.

Myrmica sculptiventris R a d c h e n k o et Elmes, 2 0 0 9

(Fig. 250)

Myrmica sculptiventris Radchenko et Elmes, 2009a: 72, w, China.

Type localities and type specimens. South China, for details sec material examined.
Material examined. Holotype, w, "China, N Sichuan Prov., W Pingwo, 15 km NW Muzuocangzu-
xian, Doubuhe riv., 32°41'32"N, 104°22'52"E, 2175 m, 30.vii.2005, leg. Belousov and Kabak" (PE-
TERSBURG); paratype: 1 w with the same label as the holotype (KIEV).

274
 Distribution (Map 107). It is known from the type locality in the Sichuan Province of
China.
Etymology. M. sculptiventris: from a combination of the Latin words sculptus = carved
or sculptured, and ventri s = belly (gaster), to describe the unusual sculpture of the gaster.
Notes. M. sculptiventris belongs to the pachei species group and by the general shape,
character of the sculpture and pilosity is quite similar to M. schulzi, but it well differs from
that species by its frontal carinae which merges with rugae that extend to the posterior third
of head dorsum, do not curve outwards and do not merge with rugae that surround anten-
nal sockets. However, the main diagnostic feature of M. sculptiventris is the rather unusual
sculpture of its gaster: the basal third of the first gastral tergite is densely punctated and lon-
gitudinally ruguloso-striated; this sculpture gradually reduces posteriorly and the rest of
the surface of the tergite has a clear superficial hexagonal microsculpture (seen under mag-
nification > 50x). It is interesting to note that there are only two other known Myrmica
species in the Palaearctic Region with a sculptured gastral tergite: M. luteola and
M. brancuccii, but both differ from M. sculptiventris by the absence of the transversal ru-
gosity on the alitrunk dorsum. Queens and males are unknown.
Ecology is unknown except that two workers were found on mountain meadows at an
altitude 2175 m a.s.l.

Map 107. Distribution of M. sculptiventris.

Myrmica serica Wheeler W. M., 1928

(Figs 251-253)

Myrmica margaritae var. serica Wheeler W. M., 1928a: 8, w, China; Chapman and Capco 1951: 127.
Myrmica ritae var. serica: Weber 1950: 223, q.
Myrmica serica: Radchenko 1994b: 44; Bolton 1995: 283; Radchenko and Elmes 1998: 7, m; Elmes and
Radchenko 1998: 222; Radchenko et al. 2001: 217; Radchenko and Elmes 2001a: 221; Radchenko,
Elmes and Viet 2006: 39.

275
 Type localities and type specimens. "A single specimen from Yunnanfu".
Material examined. Holotype, w, "Yunnanfu, Silvestri" (HARVARD); non-type material: > 150 w,
21 q, 11 m form the central and southern China, Taiwan and Vietnam.
Distribution (Map 108). Central and southern China, Taiwan, northern Vietnam; it
seems the commonest and widespread ritae-group species.
Etymology. M. serica: named for the "Land of Serica" the Greek/Roman name for
a poorly known Kingdom in East Asia, that gave its name to the Latin serica = silk, so that
the old name for China became synonymous with "The Land of Silk".
Notes. M. serica belongs to the ritae-complex of the ritae species group. The sculpture
of the head dorsum of the workers is not very coarse, the frons between frontal carinae level
with the eyes having at least six sinuous rugae, which discriminates it the sympatric
M. pulchella and M. sinensis that have only four very coarse rugae on the frons; M. pararitae
has almost straight, not sinuous rugosity on the head dorsum, and M. serica is distinctly
smaller than M. titanica (see the Key, Chapter 5.7).
Ecology. In Taiwan this species lives in forests at about 2000 m a.s.l. Here M. serica
builds nests in open clearings, under stones, in and under pieces of wood and in tree stumps
in a way reminiscent of M. ruginodis or M. rubra in European woodland. Nests were
moderately sized though one or two could have contained as many as a 1000 workers, some
nests were polygynous but at most only 2-3 queens per nest were observed. These ants were
not particularly aggressive but we were stung several times and the sting seemed no more
or less painful than that of any other similarly sized Myrmica species. A nuptial flight oc-
curred in early August, it seemed that mating took place high above the trees, rather like
a M. rubra swarm. In Vietnam Katsujuki Eguchi (pers, com.) found them at an altitude of
between 1800 and 2200 m. a.s.l. living in similar conditions to that observed in Taiwan. In
Shaanxi province of China they live at somewhat lower altitudes (c. 1200 m a.s.l.) which
probably reflected the cooler climate further north.

Map 108. Distribution of M. serica.

276
 Myrmica siciliana Radchenko, Elmes et Alicata, 2006
(Figs 254-256)

Myrmica siciliana Radchenko, Elmes et Alicata, 2006: 502, w, q, m, Italy (Sicily).

Type localities and type specimens. Sicily, for details see material examined.
Material examined. Holotype, w, "Italy, Sicily, Monti Nebrodi, Piano Menta, 1500 m a.s.L, grass-
land in a damp area, 2.x.2004, leg. A. Alicata" (MILAN); paratypes: 5 w, 6 m from the nest of the holo-
type; 1 q, "Italy, Sicily, Monti Nebrodi, Piano Menta, 1500 m a.s.L, grassland in a damp area, 2.x.2004,
leg. A. Alicata"; 17 w, 40 m, the same locality, date and collector as previous; 5 w, 1 m, "Italy, Sicily,
Monti Nebrodi, Portella Scafi, 1450 m a.s.L, grassland near pond, 2.x.2004, leg. A. Alicata"; 8 w, 6 m,
"Italy, Sicily, Monti Nebrodi, near Pizzo Scavello, 1575 m a.s.L, bare patch in a beech wood, 1.x.2004,
leg. A. Alicata"; 22 w, 2 q, 5 m, same locality and collector as previous, but date 2.x.2004; 15 w, 5 m,
"Isc24 (Elmes coll.), Sicily, reg. Monti Nébrodi, Mt. Soro, Piano Menta с. 1550 m. N facing escarp,
grazed open place among trees, nest in soil. 28.viii.2005, leg. G. W. Elmes"; 33 w, 9 m, Isc25, same col­
lection details as previous; 48 w, ISC26, same collection details as previous; 24 w, Isc27, same collec­
tion details as previous; 3 q, Iscl5, mating swarm just ending, taken from random sample of 3 Myrmica
spp. and Lasius sp. and Aphaenogaster sp. running on the ground, same site details as previous; 30 w,
4 m, "Isc28, NE Sicily, reg. Monti Nébrodi, Mt. Soro, Piano Menta, с. 1450 m, N facing escarp,
grazed damp "lawn", nest in soil at the edge of damp place, 28.viii.2005, leg. G. W. Elmes" (MILAN,
LONDON, KIEV, ELMES, VERONA).
Distribution (Map 109). Sicily.
Etymology. M. siciliana: named for the Island of Sicily, where it is believed to be
endemic.
Notes. M. siciliana belongs to the schencki species group. Although it shares many mor­
phological features with other species of the group, it well differs from all others as follows.
The female castes well differ from the other species of the schencki-group (except for the
oriental M. onoyamai) by their less-extended frontal lobes (mean FLI 1.29 vs. > 1.40); the
frontal lobes of M. onoyamai (mean FLI 1.20) are even less extended. The distinctly wide

Map 109. Distribution of M. siciliana.

277
frons of the workers and queens of M. siciliana (mean FI 0.30) well separates them from
M. ravasinii, M. schencki and M. pelops. Finally and not least, although subjective and
difficult to quantify, the shape of the lobe at the base of scape of workers and queens of
M. siciliana appears very different to that of the other species. Males of M. siciliana most
resemble those of M. schencki but differ from them by a relatively longer and lower petiole,
and by a distinctly less angled curve at the base of the scape. They are easily distinguished
from those of M. ravasinii and M. caucasicola by the short standing hairs on legs and
antennae; from M. deplanata by a distinctly longer scape, and from M. onoyamai and
M. koreana by a somewhat longer scape that is distinctly curved at its base.
Ecology. M siciliana lives in the mountains of northeast Sicily at fairly high altitudes
(> 1500 m a.s.l.). Generally they nest in the soil in areas of grazed grassland with no obvi-
ous above-ground nest structure; we did not see woven funnel entrances that are typically
associated withM. schencki, M. koreana, M. obscura and several North American schencki-
like species. M siciliana sometimes nests in open grassy areas: glades or even bare areas in
beech-woods, but they are most abundant in damp areas of open grazed grassland. They do
not require permanently wet soil, rather they live on the drier margins of ponds and areas.tem-
porarily flooded following heavy rain. M siciliana colonies face quite strong competition from
M. sabuleti and Tetramorium, Lasius, Formica and Aphaenogaster species and like many
other species of Myrmica in such circumstances, they forage mostly in the early morning and
early evening. This species was believed to be restricted to the Mt. Nebrodi region of Sicily
but we found two specimens (queen and worker foraging in mature woodland) on the neigh-
bouring Mt. Etna region. It is probably endemic to just these Sicilian mountains but we can
not rule out the possibility that it might also live in the adjacent Calabrian mountains of
southern Italy.
When collecting we observed the end of a nuptial flight in an area of damp grassland (ca.
0.5 ha) among open mixed woodland where the holotype nest had been taken on August
28th 2005. Sexuais of both M. siciliana and M. spinosior were found leaving their nests
and we caught both alate and dealate queens on the ground. The actual swarm was not lo-
cated but we guess it was centred on one of the trees in the small meadow, and was a fairly
local affair because swarming on the day did not seem to be widespread.

Myrmica sinensis Radchenko, Zhou et Elmes, 2001

(Fig. 257)

Myrmica sinensis Radchenko, Zhou et Elmes, 2001: 212, w, China.

Type localities and type specimens. South China, for details see material examined.
Material examined. Holotype, w, "S. China, NE Guangxi Province, Hua Ping, 1460 m, 17.viii.1998,
leg. J. Fellowes" (LONDON); paratypes: 3 w from the nest of holotype; 1 w, "S. China, Guangxi
Province, Daw Ling , 26.iv.1997, DWL-62, leg. J. Fellowes"; 1 w, "S. China, Guangxi Province, Da
Yao Shan, 1290 m, 16.ix.1998, DYS-39, leg. J. Fellowes" (KIEV, GUILIN, ELMES).
Distribution (Map f fO). It is known only from the type localities from the Guangxi
Province of China.
Etymology. M. sinensis: from a combination of the classical Latin name Sinœ = China
(hence sinica) and the adjectival suffix ensis = place of origin or habitat, to indicate this is
a Chinese species.

278
 Map 110. Distribution of M. sinensis.

Notes. M. sinensis belongs to the ritae-complex of the ritae species group. By its head
dorsum, that is very coarsely rugose with the frons having only four coarse longitudinal
rugae between frontal carinae level with the eyes, it resembles M. margaritae, M. pulchella
and M. emeryi. It well differs from M. margaritae by the presence of long standing hairs on
the alitrunk dorsum and occipital margin. The much more developed reticulation on its
head dorsum, where the rear half is coarsely reticulated and only the frons has longitudinal
rugae, well distinguishes it from M. emeryi and M. pulchella. Queens, males and ecology are
unknown.

Myrmica sinoschencki Radchenko et Elmes, 2008

(Fig. 258)

Myrmica sinoschencki Radchenko et Elmes, 2008: 778, w, China (in: Radchenko et al. 2008).

Type localities and type specimens. South China, for details see material examined.
Material examined. Holotype, w, "China, Sichuan Prov., Maoxian, 40 km N Winchuan, 1800 m,
23.viii.2002, leg: S. Murzin and I. Shokhin" (PETERSBURG); paratypes: 2 workers with the same
label as holotype (PETERSBURG, KIEV).
Distribution (Map 111). It is known only from the type locality from the Sichuan
Province of China.
Etymology. M. sinoschencki: from a combination of the word sino =• pertaining to China
(derived from the classical Latin name Since = China) and the name schencki, to indicate
that this species belongs to the schencki species group.
Notes. M. sinoschencki belongs to the schencki species group. It differs from all known
species of the schencki-gvoup (except forM. onoyamai andM. inucta) by its distinctly wider
frons (mean FI > 0.33 vs. < 0.32). M. sinoschencki has much more extended frontal lobes
thanM. onoyamai (FU > 1.35 vs. < 1.25), while the very reduced sculpture of M. inucta,

279
appearing dark and shiny, well distinguishes from all other species of the schencki-group
(for more details see Radchenko, Elmes and Alicata 2006; Radchenko et al. 2008). Queens
and males are unknown.
Ecology is unknown.

Map 111. Distribution of M. sinoschencki.

Myrmica slovaca Sadil, 1952

(Figs 259-261)

Myrmica slovaca Sadil, 1952: 259, w, q, m, Czechia, Slovakia, Ukraine and Bulgaria; Arnoldi 1968:
1170, 1970: 1842; Kutter 1977: 71; Arnoldi, Dlussky 1978: 534; Radchenko 1994b: 42, 1994c:
144, 1994d: 80; Bolton 1995: 283; Radchenko and Elmes 2009b: 521.
Myrmica curvithorax Bondroit, 1920a: 148, w, Spain (?); Bolton 1995: 278 (see Notes below).
Myrmica salina: Arnoldi 1970: 1842; Arnoldi, Dlussky 1978: 534, misidentifications.
Junior synonym of M. salina: Seifert 1988: 25; 1996: 228; 2002: 96; Radchenko and Elmes 2004: 229;
Seifert 2007: 201; not confirmed, see Notes to M. salina, above.

Type localities and type specimens. M. slovaca: "Finds: Chomutov, l-VII-1951 (Sadil); Somotor,
VIII-37 (Hoffer), Slov. Nové Mesto, 1.VIII.34 (Hoffer), Dolni Almas, VI1I-37 and 20.VII.38 (Babala),
1938 (Zálesky), Chorovce, 1.VI.48 (Novak) -USSR: Berehovo (former Carpathian Russia) (Zálesky).
-Bulgaria: Ba kovo, 8.VII.38 (Heberlandt)". M. curvithorax: "Espagne (Schramm)".
Material examined. Syntypes of M. slovaca: 4 vv, "Chorovce, 1.VI.48, Slovakia, Novak",
"M. slovaca Sadil (det. 1951)", "Mus. Nat. Prague Inv. 3066" (PRAGUE); 4 w, "Bohemia occ, Chomu-
tov, 1.VIII.1951, leg. J. Sadil", "M. slovaca Sadil det. J. Sadil", "Paratypus", "Inst. Zool.
P.A.N. Warszawa 102/57" (WARSAW); 5 w, "CSSR, Bohemia, Chomutov, 3.7.51, leg. Sadil"
(MOSCOW); holotype оШ. curvithorax, w, "Espagne Schramm", "Type", "R. I. Sc. N. B. I. G. 21.400"
(BRUSSELS); non-type material: > 50 w, 5 q and 5 m from the whole range of the species.
Distribution (Map 112). Germany, Austria, Switzerland, Hungary, Czech Republic,
Slovakia, Poland, Bulgaria, Romania, Slovenia, Byelorussia, Ukraine, southern part of

280
European part of Russia (to the north up to Nizhniy Novgorod; see Zrianin and Zrianina,
2007), south of West Siberia, Kazakhstan, Altai Mts.; Spain (query-see Notes below).
Etymology. M. slovaca: named to indicate that this species was found in the central
European Republic of Slovakia. M. curvithorax: from a combination of the Latin word
curvus = curved and thorax = breast or thorax, to indicated that the thorax appears curved
when seen in profile.
Notes. M. slovaca belongs to the spcioides-complex of the scabrinodis species group.
It is easily distinguished from any other species from this group by its very narrow frons
(worker FI < 0.30). However, the males have short standing hairs on the legs and scape
and are hard to distinguish from males of M. specioides and some other species.
The taxonomic history of this species during the last few decades has been somewhat
complicated. Arnoldi (1970) considered M. slovaca to be a good species that is closely re-
lated to M. salina. However, the specimens preserved in MOSCOW that Arnoldi deter-
mined as M. slovaca and M. salina, in fact belong to the same species. Seifert's (1988)
synonymy of M. slovaca with M. salina was most probably based on this material. Later
Radchenko (1994b-d) considered M. salina to be a lobicornis-group species and revived the
name slovaca from synonymy, but Seifert (2002) disagreed with this view and once more
synonymised M. slovaca with M. salina (see notes to M. salina). This question was finally
resolved after we found some of Ruzsky's original type specimens of M. salina in GENOA
that clearly differed from those of M. slovaca (see Radchenko and Elmes, 2009b and Notes
to M. salina, above).
We also studied the holotype specimen of M. curvithorax and found that is indistin-
guishable from M. slovaca. Although the name curvithorax is older we will apply to
the Commission of Zoological Nomenclature with the proposition to suppress the
name M. curvithorax and consider the name M. slovaca as senior synonym (based on para-
graphs 23.9.1.1, 23.9.1.2 and 23.9.2 of the International Code of Zoological Nomenclature
1999).

Map 112..Distribution of M. slovaca.

281
 Apart from the fact that the name N1. slovaca is well known and has been used many
times whereas M. curvithorax was almost "forgotten", that we suspect that the type locality
of M. curvithorax (Spain) is incorrect and results from a labelling error. Based on data gar-
nered from the literature and examination of numerous collections, the distribution of
M. slovaca ranges from Germany, Austria, Switzerland and Slovenia in the west to the Altai
Mts. in the east. Therefore, we are very dubious that Bondroit's specimen actually came
from Spain. First, the fauna of the Iberian Peninsula is rather isolated from that of Western
Europe generally, and the absence of M. slovaca from the quite well-studied French fauna
makes it seem unlikely that its range could extend to Spain. Secondly, there have been
several extensive investigations of Iberian Myrmica during the last couple of decades and
no further specimens of M. slovaca {"curvithorax") have ever been found. Thirdly, it is
highly unlikely that failures to find it in Spain and France are due to misidentifications, be-
cause its distinctive narrow frons means that even a single worker is easily separated from
any other species of the scabrinodis-group (remember that Bondroit, who has a fine record
for describing "good" species, felt justified in describing his species from a single worker).
Ecology. Generally, M. slovaca is a rather thermophiious species that builds its nests
directly into the soil. In central Europe it is frequently found in dry meadows but in the
eastern part of its range, M. slovaca mainly inhabits intrazonal, relatively wet and often salted
habitats in the steppe zone. Thus it could be a relict steppe species in central Europe.

Myrmica smythiesii Forel, 1902

(Figs 262, 263)

Myrmica smythiesii Forel, 1902: 226, w, India (?); Bingham 1903: 269; Forel 1904b: 22; Emery 1908a:
166 (part., nec material from Middle Asia and Siberia), 1921: 411; Menozzi 1939: 295, 1947: 456
{part., nec material from Middle Asia and Siberia); Chapman and Capco 1951: 129; Radchenko
1994b: 41; Bolton 1995: 383; Radchenko and Elmes 2001b: 253, 2002: 37, q, nec Ruzsky 1905: 659
(as Myrmica rugosa smythiesii, part., nec material from Middle Asia and Siberia), 1915: 507;
Kuznetsov-ugamsky 1927: 193, misidentifications.
Myrmica smythiesi subsp. himalayana Weber, 1947: 460, w, India; Bolton 1995: 279. Synonymy by
Radchenko and Elmes 2001b: 253; confirmed here.

Type localities and type specimens. M. smythiesii: "Diverges localités de l'Himalaya, de 7000' à
12,000' (Smythies); Deoban 8500' (Smythies); N.-O. Himalaya 8000! (Gambi)". M. smythiesi subsp.
himalayana: "Type locality: India: Simia (Wroughton). Described from three workers in Dr. W.
M. Wheeler's collection".
Material examined. Lectotype of M. smythiesii (designated by Radchenko and Elmes 2001b), w,
"Himalaya (Smythies)", "M. smythiesii Forel", "LX/1, LX/12" (GENEVA); paralectotypes: 2 w with
the same labels as lectotype; 1 w, "M. smythiesi Forel, Himalaya, 7000-12,000 ft (Smythies)", "M.C.Z
type 20533" (HARVARD); 2 w (upper with gaster and waist stuck separately; bottom - without post-
petiole and gaster), "M. Smythiesii For. Himalaya" (GENOA); lectotype of M. smythiesi subsp. hi-
malayana (designated by Radchenko and Elmes 2001b), w, "India, Simla (Wroughton)" (HARVARD);
paralectotypes: 2 w with same labels as lectotype (HARVARD); non-type material: 8 w, 2 q from India
and Nepal.
Distribution (Map 113). India, Nepal.
Etymology. M. smythiesii: this species was dedicated to "Smythies" who collected the
material; most probably this was the Englishman Arthur Smythies, who started work for the

282
Indian Forest Service in 1873 retiring from the post of Conservator in 1902, who was the
father of the more famous forester and ornithologist E. A. Smythies. M. himalayana: named
for the Himalaya region where it is found.
Notes. M. smythiesii typifies the smythiesii species group. It seems that Forel added the
extra "г" at the end of its name in error and although many later authors have used the
more logical spelling "smythiesi" the name as spelt here is correct. It most resembles
M. fortior, differing by the sculpture of the petiole and postpetiole, somewhat shorter
propodeal spines and slightly smaller body size.
The subspecies himalayana was originally distinguished fromM. smythiesii by its much
shorter propodeal spines (Weber 1947: 960). We have studied the types of both forms and
have found that while the length of the propodeal spines of subsp. himalayana are some­
what shorter than those of M smythiesii (ESLI 0.15-0.19 vs. 0.18-0.24 respectively) the
relative lengths overlap (Radchenko and Elmes 2001b). As we can find no other differences
we consider subsp. himalayana to be a junior synonym of M. smythiesii.
In earlier publications M. smythiesii was believed to be much more widespread and com­
mon than it actually is because authors, including Forel, had much too wide a definition of
this species, attributing to it species from several species groups. For example, Forel (1904a)
included as M. smythiesii material from the Russian Far East ("Middle Ussuri"); Emery
(1908a, 1921 ) indicated that its range extended from the Himalaya to the "East Turkestan"
(i.e. Tien-Shan and Pamir Mts.), Altai and East Siberia, while Ruzsky (1905) recorded it
from the Middle Asian mountains, and Ruzsky (1915) - from Tibet. Kuznetsov-ugamsky
(1927) simply repeated Ruzsky's (1905) data on the occurrence this species on the Middle
Asian mountains, but noted that this record needs confirmation. We are sure, that all these
records should be attributed species from the rubra- and/or dshungarica-groups. Males are
unknown.
Ecology is poorly known, except that this species was found at altitudes between 2100
and 3650 m a.s.l., where, based on its representation in collections, it is probably quite rare.

Map 113. Distribution of M. smythiesii.

283
 Myrmica specioides Bondroit, 1918
(Figs 264-266)

Myrmica specioides Bondroit, 1918: 100, w, q, nec m, Belgium and France; Collingwood and Yarrow
1969: 57; Pisarski 1975: 12; Kutter 1977: 70; Collingwood 1979: 56; Espadaler 1981b: 189; Seifert
1988: 16; Radchenko 1994b: 42; Seifert 1994: 11, 1996: 224; Radchenko et al. 1997: 488;
Czechowski et al. 2002: 26; Seifert 2002: 95; Radchenko and Elmes 2004: 229; Schlick-Steiner et
al. 2003: 159; Gusten et al. 2006: 29; Seifert 2007: 197, nec Bondroit, 1920a: 146, m, misidentifi-
cation.
Tetramorium silvestrianum Emery, 1924: 167, w, Spain. Synonymy by Güsten et al. 2006: 29.
Myrmica scabrinodis var. sancta Karawajew, 1926a: 49, w, Ukraine; 1927a: 285; Weber 1950: 208.
Synonymy by Stitz 1939: 90; Sadil 1952: 249 (as synonym of M. scabrinodis); Dlussky et al. 1990:
182; Atanassov, Dlussky 1992: 93; Radchenko 1994d: 77; Bolton 1995: 283 (as synonym of
M. bessarabica); Kutter 1977: 70; Radchenko and Elmes 2004: 229 (as synonym of M. specioides);
the latter confirmed here.
Myrmica sancta: Arnoldi 1970: 1840, m; Arnoldi, Dlussky 1978: 532; Seifert 1988: 21, q.
Leptothorax nevodovskii Karawajew, 1926b: 164, w, Georgia. Synonymy by Radchenko 1994d: 77;
Arakelyan 1994: 23 (as synonym of M. bessarabica); as synonym of M. specioides: syn. nov.
Myrmica rugulosoides var. striata Finzi, 1926: 96, w, q, m, Italy. Synonymy by Collingwood 1979: 56;
Bolton 1995: 283; Güsten et al. 2006: 29; consider as provisional here (see also Notes below).
Myrmica scabrinodis st. striata: Santschi 193lb: 344.
Myrmica puerilis Stärcke, 1942: 27, w, q, m, Netherlands; Weber 1948: 287. Synonymy by Colling-
wood and Yarrow 1969: 57; Collingwood 1979: 56; Seifert 1988: 16; Bolton 1995: 282; Güsten et
al. 2006: 29; confirmed here.
Myrmica puerilis ab. dolens Stärcke, 1942: 27, w, q, m, Netherlands, syn. nov.
Myrmica bessarabica subsp. dolens: Bolton 1995: 278.
Myrmica balcanica Sadil, 1952: 253, w, q, m, Czechoslovakia, Serbia and Bulgaria. Synonymy by
Arnoldi 1970: 1840 (as synonym of M. sancta); Dlussky et al. 1990:182; Atanassov, Dlussky 1992:
93; Radchenko 1994d: 77; Bolton 1995: 277 (as synonym of M. bessarabica); Pisarski 1975: 12;
Kutter 1977: 70; Collingwood 1979: 56; Seifert 1988: 16; Güsten et al. 2006: 29 (as synonym of
M. specioides); the latter confirmed here.
Myrmica balcanica var. scabrinodoides Sadil, 1952: 255, w, q, m (part.), Czechoslovakia. Synonymy
by Atanassov, Dlussky 1992: 93; Bolton 1995: 283 (as synonym of M. bessarabica); by Pisarski
1975: 12; Kutter 1977: 70; Collingwood 1979: 56; Güsten et al. 2006: 29 (as synonym of
M. specioides); the latter confirmed here.
Myrmica sancta subsp. tshuliensis Arnoldi, 1976: 553, w, m, Turkmenistan; Dlussky, Zabelin 1985:
210. Synonymy by Dlussky et al. 1990: 182; Radchenko 1994d: 77; Bolton 1995: 284 (as synonym
of M. bessarabica); as synonym of M. specioides: syn. nov.
Myrmica scabrinodis ahngeri: Seifert 1988: 16; Güsten et al. 2006: 29; not confirmed, see Notes to
M. scabrinodis, above.
Myrmica scabrinodis subsp. rugulosa var. rugulosoides: Collingwood 1979: 56; not confirmed, see
Notes to M. scabrinodis, above.
Junior synonym of M. scabrinodis: Emery 1921: 40; Finzi 1926: 99; Sadil 1952: 249; Bernard 1967: 116;
not confirmed, see Notes to M. scabrinodis, above.
Junior synonym of M. rugulosoides: Santschi 1931b: 342; Weber 1950: 208; not confirmed, see Notes
to M, scabrinodis, above.
Junior synonym of M. bessarabica: Atanassov, Dlussky 1992: 93; not confirmed (sec notes to
M. bessarabica in Chapter 3.5).

Type localities and type specimens M. specioides: " Belgique et France, commun partout". T. sil-
vestrianum: "Manresa al N. di Barcelona, 2 esemplari". M. scabrinodis var. sancta: "Umgegend von

2S4
Karadagh, nicht weit von Theodosia, Krym, lO.v.1920, Nr. 2854, leg. Karawajew, ww". L. nevodovskii:
"Kodschory bei Tiflis, 1. VII. 1912, Newodowsky, 1 w". M. rugulosoides var. striala: "La descrizione
di questa varietà è fatta su esemplari catturati da me ad Opcina (dintorni di Trieste). Posseggo nume-
rose w, q e m dell' Istria settenrionale, cacciate sulle rive di ruscelli. La varietà è sparsa pure in Italia
(Bologna, ecc)". M. puerilis: "Zandvoort, 6-8-'30, w, q, m, v. d. Wiel; Vlieland l-9-'29, w, q, m, A. Re-
claire; Vlieland 31-7-31, w, q, m, v. d. Wiel; Wollmisse (Thiir.) 12-7-47, q, 15-7-'15, m, Cohrs; Broût-
Vernet, m, H. du Buysson; Broût-Vernet (Allier) 12-6-'27 m, H. du Buysson; St. Angel, m, coll. Du
Buysson. Typen m. с. Homoiotypen coll. v. d. Wiel". M. puerilis ab. dolens: "Broût-Vernet (Allier), 8-
5-'09, q H. du Buyasson 8-8-'09, w, q, m id. St. Angel (Dép. ?) m, coll. du Buysson. Typen Spr.'s с
Paratypen coll. v. d. Wiel". M. balcanica: "Czechoslovakia: Praha-Huboëepy, 1936 (Kult), 24-6-49
(Sadil); Listice near Beroun, 14-8-41 (Frasky); Sárka near Prague, 14-8-43 (Sadil); Vinafická hora
near Kladno, 26-3-43 (Sadil); Katovka near Zákolany, 20-6-43 (Sadil); Hradist near Litoméfice,
13-6-48 (Sadil), 20-8-50 (Zednicek); Chomutov, 2-8-51 (Sadil); Brno-Hády, 5-7-36 (Hoffer); Pav-
lovské vhry, 15-7-38 (Sadil); Slov. Nov. Mesto, 27-7-36 (Hoffer); Somotor, 21-7-51 (Sadil); Yugo-
slavia: Petrovo Selo in Croatis, 26-7-38 (Novak); Vlasenica in Serbia, 30-6-38 (Novak); Konjic,
4-7-38 (Novak); Bulgaria: Ljulin, 6-35 (Tàborsky)". M. balcanica var. scabrinodoides: "Praha-Boh-
nice, 5-38 (Sadil); Oblik near Louny, 9-8-38 (Sadil); Liten near Beroun, 2-5-38 (Trakal); Praha-
Huboccpy, 24-4-49 (Sadil); Sárka near Prague, 14-8-43 (Sadil); Popice near Brno, 18-7-38 (Sadil);
Okof near Noutonìce, 13-6-43 (Sadil); Kylesovice in Silesia, 4-9-37 (Ing. Palásek)". M. sancta subsp.
tshuliensis: "Копстдаг. Голотип (w), 18 паратипов, аллотип (m) - ущелье Чули, западнее Фирюзы
(Арнольди, 1935, № 6609)" [Kopetdagh. Holotype (w), 18 paratypes, allotype (m) -gorge Chuli, west
of Firiuza (Arnoldi, 1935, No 6609)].
Material examined. Lectotype of M. specioides (designated in material by Pisarski, but published
here first), w, "St. Afrique", "Myrmica specioides Bondr. type", "R. I. Sc. N. B. 21.400", "Lectotype
det. B. Pisarski" (BRUSSELS); paralectotypes: lq, "Mt. Valérein"; 6 w, "Paris, f. de Buffon"; 15 w, "St.
Afrique"; 1 q, "St. Afrique, Août-Sept., F. Rabaud"; 1 w, "Issy-Pes-Moulineaux, 19-9-1915"; 5 w,
"Paris Museum"; 1 w, "Cahors"; 1 w, "Le Pouligum"; 7 w, "Prades"; 12 w, "Banyuls"; 4 w, "Paris, V-
6-1917"; 4 w, "Meriel S. et O."; 2 w, "Sayat, P. de Dôtne"; 3 w, "St. Aquan, M. Pic"; 1 w, "Hyëres,
col. de Gaulle"; 1 q, "La Héve, 1-10-16"; 3 w, "Fonteney aux Roses, 5-1915"; 2 q, "Haisans-Saff.";
1 w, "Calaisis"; 1 q, "Latrecy, Ht. Marne"; 2 m, "St. Afrique, 1916, 8 Aug., Rabaud" (BRUSSELS);
1 w, "M. specioides Bondr. Type Paris" (GENOA); syntypes of M. scabrinodis var. sancta: 11 w, "окр.
Карадага, Крым, Караваев" [vicinity of Karadagh, Crimea, Karawajew], "2854 Coll. Karawajewi", "Myr­
mica scabrinodis Nyl. Karawajew det.", "Syntypus Myrmica scabrinodis var. sancta Karaw." (KIEV)
(see Notes below); holotype of L. nevodovskii, w, "Kodschory, 1400 m. prope Tiflis 1.VII. 12", "Mus.
Caucas. 48-12 G. Nevodovski", "4240 Coll. Karawjewi", "Leptothorax (Leptothorax) nevodovskii
Karawajew typ.", "Holotypus Leptothorax (Leptothorax) nevodovskii", "Myrmica nevodovskii
(Karaw.) K. Arnoldi det., "Myrmica bessarabica Nason. det. Radchenko" (KIEV); lectotype of
M. rugulosoides var. striata (designated here), w, "Opcina querica 15.viii.20", "Typo", "B. Finzi coli.
purch. 1950", "B. Finzi coli, purch. 1950", "Type series M. rugulosoides striata Finzi" (HARVARD);
holotype oiM. puerilis, m, "Zandvoort, 6-8-1930, v. d. Wiel", "holotype puerilis Stärcke", "Myrmica
specioides Bondroit, 1918 det. A. J. van Loon 1992" (LEIDEN); holotype of M. puerilis ab. dolens, m,
"Everts Woeste Hoeve, 7", " " "scabrinodis subsp. ? det A. Stärcke", "ab. Dolens Stärcke Holotype",
"Myrmica specioides Bondroit, 1918 det A. J. van Loon" (LEIDEN); syntypes of M. balcanica: 1 w
"Slovenske Nove Mesto, 27-7-36, leg. Hoffer", "M. scabr. rugulosoides, det. Dr. Zálesky", "Cotype",
"M. balcanica Sad. Sadil det. 1951", "Mus. Nat. Pragae, Inv. 3409", "Myrmica puerilis Stärcke, det.
С A. Collingwood"; 2 w, "M. rugulosa", "26.VII.38. Petrovo Selo, Chorvatsko, leg V. Novak", "Co­
type", "M. balcanica Sad. Sadil det. 1951", "Mus. Nat. Pragae, Inv. 3405", "Myrmica puerilis Stärcke,
det. С A. Collingwood"; 2 w, "Ljulin Pl. Bulgaria, VI.35, leg. Dr. Tàborsky", "Cotype", "M. balcanica
Sad. Sadil det. 1951", "Mus. Nat. Pragae, Inv. 3406", "Mus. Nat. Pragae, Inv. 3058", "Myrmica puerilis
Stärcke, det. C.A. Collingwood"; 8w, "Hradistéu Litoméfice, 19.6.48, Sadil", "854", "Mus. Nat. Pra-
gae, Inv. 3043"(PRAGUE); 3 w, "Bohemia осе, Chomutov, 2.VII1.1951, leg. Sadil", "M. balcanica

285
Sadil det. Sadil", "Paratypus", "Inst. Zool. P.A.N. Warszawa, 102/17" (WARSAW); lectotype of
M..balcanica var. scabrinodoides (designated here), w (middle specimen on the pin with 3 w), "Bo­
hemia centr. Hubocepy ad Praha, 15.8.1951, leg. Sadil", "M. balcanica var. scabrinodoides Sadil det.
Sadil", "Paratypus", "Inst. Zool. P.A.N. Warszawa 102/17" (WARSAW) (see Notes below); paralec-
totypes: 2 w and 3 w heads on the pin with lectotype; 2 m, "Bohemia centr. Hubocepy ad Praha,
15.8.1951, leg. Sadil", "M. balcanica var. scabrinodoides Sadil det. Sadil", "Paratypus", "Inst. Zool.
P.A.N. Warszawa 102/17", "prep. 467" (WARSAW); holotype of M. sancta subsp. tshuliensis, w,
"6609, Копстдаг, Чули, 1935, К. Арнольдп" [6609, Kopetdagh, Chuli, 1935, К. Arnoldi]; paratypes: 27
w, 13 m with the same labels as in holotype (MOSCOW); 6 w, 3 m with the same labels as in holotype
(PETERSBURG) (see Notes below); non-type material: several hundred w, > 50 q, > 50 m from the
whole area.
Distribution (Map 114). Europe (to the north to southern England and Denmark), Cau­
casus, Asia Minor, Iran, Turkmenistan, south of West Siberia and northern Kazakhstan, to
the east until Altai Mts.; introduced to North America (see Jansen and Radchenko 2009).
Etymology. M. specioides: from a combination of the Latin words speciosus = good
looking and aides = like or resemble. M. silvestrianum: from the name Silvestri with the ad­
jectival suffix anum = from, connected with, presumably to indicate the material was from
the Italian Prof. Filippo Silvestri (1873-1949), who was a renowned entomologist and avid
insect collector in the early 20(h century. He became Director of the Laboratorio di Ento­
mologia di Portici (near Naples). M. sancta: from the Latin word sancta = sacred, proba­
bly because the types were found in or near a monastery or church. M. nevodovskii: named
for the collector Gavriil Stepanovich Nevodovski (1874-1952), a well-known Russian
botanist and mycologist. M. striata: from the Latin words stria = furrow, channel combined
with the adjectival suffix ata to give furrowed, to described the distinct striations of the
male's head. M. puerilis: name means child like, derived from Latin puer = boy, probably
to indicate both the appearance of the head of the male and that it is one of the "newest"
species named. M. dolens: the name means grievous from the Latin word dolor = grief,

Map 114. Distribution of M. specioides.

286
because the long creases behind the eye give the male that appearance compared with var.
purilis. M. balcanica: name derived from The Balcans = Balkans (Serbia was recorded as
one of the type localities). M. scabrinodoides: from a combination of the name scabrinodis
with oides = like or resemble, to indicate that it resembles M. scabrinodis. M. tshuliensis:
named from a combination of the type locality, Tshuli Valley (alternative spelling for Chuli)
in the Kopet Dag, and the Latin suffix ensis - place of origin.
Notes. M. specioides typifies the specioides-compìex. of the scabrinodis species group
(see also Radchenko and Elmes 2004). It is quite a variable species, especially in the im-
portant diagnostic characters such as frons width, the shape of petiole, colour and sculpture.
The workers and queens are most often confused with those of M. scabrinodis, differing
from the latter by the wider frons and the less extended frontal lobes (means FI 0.39 and
FLI 1.29 vs. 0.36 and 1.38), by the somewhat shorter propodeal spines and by the shape of
the petiole, with weakly developed dorsal plate. At the same time, males of M. specioides
well differ from those of M. scabrinodis by the much shorter standing hairs on the scape and
tibiae, while by these features they are very similar to several other species (e.g. M. salina,
M. hellenica, M. turcica, M. tobiasi, M. gallienii, M. rugulosa and some others; see Table
2.2 in Chapter 2.2). The taxonomy of this species has a history of confusion (see below) and
is not yet finally resolved, we would not be surprised if modern genetically based studies
eventually show that M. specioides comprises two (or perhaps even more) somewhat cryp-
tic species.(as discussed for M scabrinodis).
Bondroit's confusion of some of the species in the scabrinodis-species group at least
partly provoked the very complicated and confused taxonomic history of M. specioides.
Firstly, Bondroit recognised M. sabuleti (in its modern treatment) under the name "scabri-
nodis" (see Bondroit 1918, 1920a, and Notes to M. scabrinodis, above). As a result, when
he described males of M. specioides he was in fact describing males of M. scabrinodis; he
repeated the same mistake later (Bondroit 1920a). That is why based on the male charac-
ters given by Bondroit, all subsequent authors until the end of the 1960's treated
M. specioides as a junior synonym of either M. scabrinodis or M. rugulosoides (now a ju-
nior synonym of M. scabrinodis, see Notes to that species) (e.g. Emery 1921: 40; Finzi 1926:
99; Santschi 1931b: 342; Weber 1950: 208; Sadil 1952: 249; Bernard 1967: 116).
It was more than 50 years after its first description before Collingwood and Yarrow
(1969) gave M. specioides the correct treatment. Thereafter, M. specioides was considered
by most "Western" myrmecologists to be a West European species and in the literature of
the former Soviet Union this name was almost completely ignored. Most "East European"
myrmecologists recognised as a distinct species M. sancta, which is distributed from
Balkans and Czechoslovakia until Kopetdagh Mts. (e.g. Arnoldi 1970, 1976; Arnoldi,
Dlussky 1978), though they recognised it as a sort of "sister" species to M. specioides. More-
over, the same taxa that some "Western" specialists considered to be junior synonyms of
M. specioides were often treated as synonyms of M. sancta by "Eastern" specialists (see
synonymy, above).
Dlussky and co-workers (Dlussky et al. 1990; Atanassov, Dlussky 1992) further "mud-
died the waters" when they revived the name M. bessarabica from synonymy, raising it to
species level, designating a lectotype and considering it to be the senior synonym of M. spe-
cioides and M. sancta (including several of their infraspecific forms). This idea was accepted
by Radchenko (1994b, c, d) and the name M. bessarabica was quite widely used through-
out Europe for about 10 years. However, Seifert (2002) disagreed with Dlussky et al.'s
opinion and proposed that M. bessarabica be considered as incertae sedis in the genus

287
Myrmica; we agree with Seifert's treatment (for details see Notes to M. bessarabica in Chap-
ter 3.5; see also Radchenko and Elmes 2004). As things now stand, M. specioides is re-
garded as a good species and the senior synonym of many subsequently described taxa,
including M. sancta.
We (Radchenko and Elmes 2004) hypothesised that M. specioides might have a northern
and western range in Europe while M. sancta might have a more southern and eastern
range in Europe, Asia Minor, Iran and Turkmenistan with perhaps overlapping distribu-
tions in Ukraine and the Balkans. We studied the lectotype and paralectotypes (workers,
queens, males) of M. specioides (several tens of specimens), the syntypes (workers) of
M. sancta (and types of the earlier established synonyms of these species, see below), and
several hundred non-type specimens of these forms from the whole ranges of both taxa.
We concluded that workers and queens of M. specioides are indistinguishable from
M. sancta by all the main features, including morphometries. Males of these two species are
somewhat less similar: the European populations differ from the Caucasian and Turkish
ones by somewhat shorter standing hairs on the tibiae and tarsi, but this feature is quite
variable and is not sufficient to warrant a formal separation of these populations. Thus, we
confirm the earlier proposed synonymy of M. sancta with M. specioides (see Kutter 1977).
It should be noted that although Arnoldi (1970) designated a lectotype of M. sancta ("I pro-
pose a lectotype from the type series of Karawajew from Karadagh, Crimea)" (loc. cit.,
p. 1840) he did not select and label an actual specimen from the corresponding material;
therefore, we have to ignore Arnoldi's lectotype and consider all type specimens still to be
syntypes.
We examined the holotype (worker) of Leptothorax nevodovskii, described from Geor-
gia, and considered that it clearly belongs to M. specioides.
We examined the worker and the male from HARVARD from the type series of
M. rugulosoides var. striata, labelled as: worker- "Opcina querica 15.viii.20", "Typo", "B.
Finzi coll. purch. 1950", "B. Finzi coll. purch. 1950", "Type series M. rugulosoides striata
Finzi"; males - "Opcina querica 15.viii.20", " ", "Typo", "B. Finzi coll. purch. 1950", "B.
Finzi coll. purch. 1950", "Type series M. rugulosoides striata Finzi", "Myrmica sabuleti
Meinert det Seifert 1990". The worker has wide frons (FI 0.39), moderately extended frontal
lobes (FLI 1.31), rather long propodeal spines (ESLI 0.41), well developed dorsal plate on
the petiolar node, and quite big lobe at the scape base. More importantly, the male has
relatively long scape (like in M. sabuleti). Based on this combination of features of both
the worker and male, we identify these specimens as M. spinosior and not M. specioides.
However, Finzi (1926: 96) stressed that male of var. striata has scape equal in length to the
three basal funicular joints ("Scapo ... lungo come i primi tre articoli del funicolo") in which
case the male is neither M. sabuleti, nor M. spinosior. We have no idea why there is such
a discrepancy between Finzi's description and investigated type specimens, except that it
may be possible that the type series of Finzi's var. striata included two different species. This
question can be solved only after investigation of all available existing type specimens of this
form. If var. striata is shown to be conspecific with M. spinosior, then its name has
priority. Consequently, we make no judgement on this issue here beyond considering syn-
onymy of var. striata with M. specioides as provisional. However, we do not agree with the
synonymy of var. striata with M. hellenica proposed by Seifert at al. (2009) that was based
on the examination of a single syntype specimen: in our experience workers of var. striata
have much more extended frontal lobes (FLI ca. 1.30 vs. < 1.20 in M. hellenica) and much
more developed lobe at the scape bend.

288
 We also studied the holotypes (males) of M. puerilis and M. puerilis ab. dolens, but un-
fortunately not workers associated with these forms; the males we examined fit within the
range of variation that conforms to the features of M. specioides. We have no doubt that the
16 syntypes workers of M. balcanica belong to M. specioides.
The single worker, kindly sent us from PRAGUE and labelled as "Kylesovice, Sil., Czh.,
4.9.1937, leg. J. Palásek", "Cotype", "M. balcanica var. scabrinodoides Sadil (det. 1951)",
"Mus. Nat. Pragae Inv. 3062", "Myrmica puerilis Stärcke, det. С A. Collingwood", in fact
is M. rugulosa. Therefore, we designated as lectotype of var. scabrinodoides a worker, pre­
served in WARSAW (for details see Material examined, above); the lectotype and 4 para-
lectotype (2 workers and 2 males) belong to M. specioides.
Arnoldi (1976) included to the type series of M. sancta tshuliensis the holotype (w) and
18 paratype workers and "allotype" male. However, we found in MOSCOW and PETERS­
BURG the holotype, and 33 workers, 16 males and even 8 queens, all having the same la­
bels as holotype and without doubt belonging to the same nest sample. Most probably, when
Arnoldi described this form he had not mounted all the specimens, the remainder plus the
queens were probably mounted later and so were not listed in the type series. We believe
that at least all workers and males mentioned above, should be considered as paratypes,
although the queens can not belong to the type series because Arnoldi originally described
this subspecies based only on workers and males.
Very recently, Gusten et al. (2006) in a revision of genus Tetramorium investigated the
type series of T. silvestrianum from Spain, and surprisingly showed that it is in fact M. spe­
cioides. We have not checked these types but based on their description and the figure in
Emery's (1924) paper it is fairly obvious that T. silvestrianum is in fact a Myrmica species.
M. specioides was regarded as the senior synonym of M. scabrinodis ahngeri and M. ru­
gulosoides. However we consider the last two names to be junior synonyms of M. scabri­
nodis (see Notes to M. scabrinodis, above), thus we do not confirm this synonymy.
Ecology. M. specioides has a lifestyle very similar to that of M. scabrinodis but prefers
distinctly warmer and drier microclimates, co-existing with M. schencki in many grassland
habitats. Based on personal observations, M. specioides colonies average several hundreds
of workers with one or two queens. The nesting preferences of M. specioides living
in several habitats have been described: on coastal dunes (Bonte et al. 2003), in forest
clearings and forest edges (Du Merle et al. 1978) and from Touraune, France (Meudec and
Lenoir 1973). The chemical secretions of the workers were outlined by Jackson et al. (1989).
Thus, despite its widespread occurrence in Europe, there has been very few ecological or
laboratory studies made of this species (under any of its earlier names), though quite
possibly some of the earlier studies of M. scabrinodis might have been made on
M. specioides (see notes on taxonomic confusions above).

Myrmica spinosior Santschi, 1931

(Figs 267-269)

Myrmica sabuleti var. spinosior Santschi, 193la: 3, w, q, m, France, Spain and Italy (also described as
new by Santschi, 1931b: 346); Weber 1948: 296.
Junior synonym of M. sabuleti: Seifert 1988: 31; Radchenko 1994d: 80; Bolton 1995: 283; not
confirmed here.
Myrmica spinosior: Seifert, 2005: 7.

289
 Type localities and type specimens. "Pyrénées-Orientales. Irun (Lindberg). Corse, Poggiolo.
Espagne, Italie. Le 1 male type est de Castelnovo, Vénétie".
Material examined. Lectotype (designated on the specimen by Radchenko and Elmes 2002;
published by Seifert 2005), w, "Typus", "Pyren. Occ. Irun, 25-26.3.1926, Lindberg", "M. sabuleti var.
spinosior Sant.", "Sammlung Dr. F. Santschi Kairouan", "Lectotype M. sabuleti spinosior Santschi,
design. Radchenko et Elmes, 2002" (BASLE); non-type material: > 20 nest scries from France, Spain,
Portugal and Italy (> 200 w, > 20 q and > 20 m).
Distribution (Map 115). West Mediterranean species (Iberian Peninsula, southern
France, Italy); record for Asia Minor (Seifert 2005) seems doubtful.
Etymology. M. spinosior: from the Latin word spinosus (thorn) with the adjectival com-
parative suffix ior giving spinosior = thornier, to describe the relatively longer propodeal
spines of this species.
Notes. M. spinosior belongs to the sabuleti-comp\ex of the scabrinodis species group
and most resembles M. sabuleti from which it well differs by its distinctly wider frons
(similar to or even wider than in M. scabrinodis (mean FI 0.37 vs. 0.33 in M. sabuleti).
However, M. spinosior differs from M. scabrinodis by the less extended frontal lobes (mean
FU 1.33 vs. 1.42 respectively).
Seifert (1988) synonymised M. spinosior with M. sabuleti but noted that workers of
"West-Mediterranean populations" have a much wider frons. More recently he decided the
differences between the two forms were sufficient to revive var. spinosior from synonymy
and raise it to the species level (Seifert 2005). We have studied the lectotype of var. spino-
sior and a considerable amount of additional material from the West-Mediterranean region
and fully agree with Seifert's proposition.
Ecology. From our observations to date the ecology and biology of M. spinosior is very
similar to that of M. sabuleti. It usually builds nests under stones or in the soil, in moun-
tain grassland pastures at about 1000-2000 m. a.s.l. Colonies can vary from a few hundred
to a thousand or more workers with several functional queens. Sexuais are in the nest in Au-
gust and nuptial flights probably take place at the same time as those as M. sabuleti. In our

Map 115. Distribution of M. spinosior.

29O
experience there is no reason to think that M. spinosior is an ecological variant of M. sabu­
leti (e.g. M. lonae) that is more thermophilic than M. sabuleti, we are more inclined to the
opinion that it is derived from a separate lineage isolated during one of the more recent
glaciations with a considerable overlap of range and possible hybridisation (see notes to
M. lobicornis and M. obscura).

Myrmica stangeana Ruzsky, 1902

(Figs 270-272)

Myrmica stangeana Ruzsky, 1902a: 234, w, Russia and Kazakhstan (also described as new by Ruzsky,
1902b: 474); Arnoldi 1970: 1841, m; Arnoldi, Dlussky 1978: 533; Seifert 1988: 23, q; Radchenko
1994b: 42, 1994c: 144, 1994d: 78; Bolton 1995: 283; Radchenko and Elmes 2009b: 509.
Myrmica bergi subsp. stangeana: Ruzsky 1903: 314, 1905: 678.
Myrmica scabrinodis subsp. stangeana: Emery 1908a: 178, 1921: 4; Weber 1950: 210.
Myrmica stangeana medvedevi Arnoldi, 1970: 1841, w, Ukraine. Synonymy by Seifert 1988: 23; Rad­
chenko 1994 e: 78; Bolton 1995: 281; Radchenko and Elmes 2009b: 509; confirmed here.
Myrmica scabrinodis var. lacustris Ruzsky, 1905: 686, w, Russia and Kazakhstan; Emery 1908a: 177,
1921: 40, Weber 1950: 206, nec Radchenko 1994b: 44, 1994c: 140, 1994c: 77; Bolton 1995: 280;
Seifert 2003: 150; Radchenko et al. 2005: 168; Radchenko, Elmes and Alicata 2006: 516. Synonymy
by Radchenko and Elmes 2009b: 509; confirmed here (see also Notes to M. deplanata, above).

Type localities and type specimens. M. stangeana: "Найдена была эта мирмика в окрестностях Иргиза
и Тургая, на глинистой полынной степи" [This Myrmica was found in the vicinities of Irgiz and Turgai,
in the clayish Artemisia steppe] (Ruzsky 1902a); "Gefunden von Herrn G. Stange auf harten Lehm­
boden, das mit Artemisia beweschen, in der Turgaischen Kirgisetsteppe. Lebt in Gängen im Boden"
(Ruzsky 1902b). M. stangeana medvedevi: "Описан по 12 ww. Голотип w - Аскания Нова, 11 паратипов"
[Described from 12 ww. Holotype w - Askania-Nova, 11 paratypes]. M. scabrinodis var. lacustris:
"Мсстонахожд.: Тобольская губ., солонцы ок. оз. Горького на пути от ст. Бсловской в г. Петропавловск,
6. VII. 96; окр. оз. Бол. Невидим, бл. с. Лопатинскаго, Курганск. окр, 27. VI. 96; солончаки у оз. Сладкого,
с Сладковское Ишимск. окр.. 28. VII. 96" [Localities: Tobolskaya Gubernia, salted marshes near the
Lake Gor'koe, on the way from st. Belovskaya to Petropavlovsk, 6.VÜ.96; vicinity of the Lake Bol'shoy
Nevidim, near vil. Lopatinskoe, Kurgansky Okrug, 27.vi.96; salted marshes near the Lake Sladkoe, vil.
Sladkovskoe, Ishimsky Okrug, 28.vii.96"] (now - Pavlodar Province of Kazakhstan, and Kurgan
Province and north-west of Altaisky Krai of Russia).
Material examined. Lectotype of M. stangeana (designated by Radchenko and Elmes 2009b), w,
"Myrmica scabrinodis stangeana Ruz., w, Киргизская степь, гор. Мал. Богдо, 4.VI.1902, М. Рузский"
[Myrmica scabrinodis stangeana Ruz., w, Kirgizskaya step', gor. Mal. Bogdo, ,4.vi.l902, M. Ruzsky]
(PETERSBURG) (see also Notes below); paralectotypes: 1 w (PETERSBURG) and 1 w (MOSCOW)
with the same label as lectotype; 1 w, "Turgai-Steppe No 26, Myrm. Stangeana m. w 2 ex.", "Paralec-
totypus des. Radchenko" (GENOA); holotype of M. stangeana medvedevi, w, "Аскания-Нова, Украина,
А 6496, V.1927, Medvedev" [Askania-Nova, Ukraine, а 6496, v.1927, Medvedev], "Myrmica stangeana
medvedevi К. Arn. Holotyp" (MOSCOW); paratypes: 6 w with the same labels as holotype (MOS­
COW); lectotype of M. scabrinodis var. lacustris (designated by Radchenko and Elmes 2009b), w
(specimen, stuck on the tip of carton triangle), "Myr. scabrinodis Nyl. var. lacustris R., [w] Am Über
Salzsee Gorjkoe im Gouv. Tobolsk (West-Sibirien) 96. M. Ruzsky" (GENOA); paralectotype: 1 w on
the same triangle as lectotype (GENOA); non-type material: about 40 workers, 2 queens and 2 males
from Russia and Ukraine.
Distribution (Map 116). Steppes from the lower reaches of the Dnepr to the eastern
Kazakhstan, to the north - until Samara Province of Russia.

291
 Map 116. Distribution of M. stangeana.

Etymology. M. stangeana: named for the collector, the Russian naturalist G. V. Stange
(Shtange), who provided material for M. D. Ruzsky from different places in European Rus-
sia and West Siberia. M. medvedevi: named for the collector, the famous Ukrainian ento-
mologist and painter, Prof. Sergey Ivanovich Mevedev (1899-1979) of Kharkov University.
M. lacustris: from the Latin word lacus = lake, pond with the adjectival suffix ustris = ori-
gin or habitat, to indicate that it was found near a lake.
Notes. We placed M. stangeana to the specioides complex of the scabrinodis species
group (Radchenko and Elmes 2004). It is characterized by a combination of features that
distinguish it from M. specioides: the antennal scape at its base only has a narrow horizon-
tal ridge or at most a very small carina; the propodeal spines are rather short, thin and not
widened at the base; the alitrunk with relatively coarse, regular, almost straight longitudi-
nal rugae; the body a rather dark, brownish-red. Additionally, the males of M. stangeana
have a very short scape (SI1 < 0.33).
Arnoldi (1970: 1841) designated as the lectotype a worker (MOSCOW), collected by
Ruzsky from Sarepta near Volgograd. Although this specimen is M. stangeana, it does not
belong to the type series and has no taxonomic value because its label-data does not cor-
respond with the type localities given by Ruzsky (see above). Therefore, we designated as
the lectotype (Radchenko and Elmes 2009b) a worker, collected by Ruzsky in "Kirgizskaya
step', gor. Mal. Bogdo" (for details see above); this locality at least does not contradict
Ruzsky's (1902a, b) data.
We examined the holotype and paratypes of M. stangeana medvedevi, as well as non-type
material, collected in the south of Ukraine. They slightly differ from M. stangeana s. str. by
having a somewhat more developed, but still very narrow, carina at the scape base, and by
other minor features such as colour, character of sculpture, shape of propodeal spines and peti-
ole. However, as the range of morphological variation of M. stangeana is not yet fully appre-
ciated we believe that such minor differences are insufficient to establish a separate subspecies.
The taxonomic history of M. scabrinodis var. lacustris, recently synonymised with
M. stangeana, is much more complicated. For many years the types of this form have been

292
considered to be lost. Furthermore, the original description (Ruzsky 1905: 686) is very brief,
incomplete and at least partly ambiguous: "Workers. Anterior clypeal margin with the small
notch medially. Propodeal spines are short, equal to 1/2 or at most 2/3 of the length of dor­
sal surface of propodeum. Frontal triangle striated distally. Antennal scape at the base is
with a small, inclined dentiform lobe. Metanotal groove weak, not deep, as a result the
mesonotum and propodeal dorsum are more flattened (in typical M. scabrinodis they are
more convex). Body rugosity is finer. Body colour is as in the type [i.e. M. scabrinodis],
but the whole gaster, except for its tip, is dark brown" (our translation to English). Much
of this description could apply to many Eurasian Myrmica species, but the most important
characters are barely understandable, especially the shape and direction (horizontal or ver­
tical) of the lobe at the base'of antennal scape. This feature is very important in separating
scabrinodis-group from lobicornis- or schencki-group species, but Ruzsky's description
could apply to either. On the other hand, Ruzsky clearly indicated a medially-notched
clypeus, which fits to the lobicornis- and schencki-group species.
In the course of the revision of the Asian Myrmica, Radchenko (1994e) found a queen
in MOSCOW that had an original Ruzsky label "H.-Петровский, Акмолинский уезд,
M. scabrinodis var. lacustris Ruzs" [New-Petrovsky, Akmolinslky uezd] (now - Akmola
Province of Kazakhstan). While this specimen can not belong to the type series of var. la­
custris (which was described only from workers), it had been identified by Ruzsky himself
and in the absence of any other material and under the assumption that Ruzsky had deter­
mined his "own" species correctly, Radchenko (1994e) designated this queen as the neo­
type of var. lacustris. However, this neotype queen is conspecific with another taxon,
M. deplanata that was originally described by Ruzsky (1905) as a quadrinomen (i.e. un­
available name). Consequently, the name lacustris was given priority (see notes to M. de­
planata and Radchenko 1994e; see also Seifert 2003; Radchenko, Elmes and Alicata 2006).
The discovery in GENOA of two specimens (workers) belonging to the Ruzsky's origi­
nal type series of M. lacustris negated the above approach because they clearly belong to the
scabrinodis-group of species, having a scape that is angled at the base with a narrow hori­
zontal carina. Therefore, M. lacustris can not be synonym of with M. deplanata (see Notes
to M. deplanata). We designated a lectotype worker of M. lacustris (for labels' details see
above) and redescribed the species (Radchenko and Elmes, 2009b). However, these type
workers are practically indistinguishable from M. stangeana, we could find no real distin­
guishing features, including morphometries, other than their rather lighter colour. Thus we
concluded that M. lacustris Ruzsky, 1905 is a junior synonym of M. stangeana Ruzsky, 1902.
Ecology. Although M. stangeana is a widely distributed species, its poor representation
in collections suggests that it is quite rare throughout its range. Consequently not much is
known about its ecology. Despite M. stangeana appearing to be a species of the Steppe
Zone, it inhabits only intrazonal, wet, usually halophytous habitats. Nests are built in soil,
sometimes with small mounds, but very often without any visible constructions. Colonies
are quite small, up to several hundred of workers, mono- or oligogynous.

Myrmica sulcinodis Nylander, 1846

(Figs 273-275)

Myrmica sulcinodis Nylander, 1846a: 934, w, q, Finland; Mayr 1855: 409, m; Smith F. 1855: 120,
1858: 116; Saunders 1880: 215; André 1883: 317; Dalla Torre 1893: 116; Ruzsky 1895: 31, 1896:

293
 73, 1905: 703; Emery 1908a .166; Bondroit 1912: 351; Forel 1915: 29; Donisthorpe 1915a: 132;
Emery 1916: 119; Bondroit 1918: 104; Donisthorpe 1919: 1; Emery 1921: 42; Müller G. 1923: 42;
Soudek 1923: 111; Ruzsky 1925: 46; Finzi 1926: 86; Karawajew 1927a: 283; Santschi 1931b: 351;
Arnoldi 1934: 157; Stitz 1939: 85; Weber 1948: 267; Sadil 1952: 242; Collingwood 1958b: 70;
Hauschteck 1965: 323; Bernard 1967: 121; Pisarski 1969a: 227, 1969b: 296; Arnoldi 1970: 1839;
Baroni Urbani 1971: 36; Kutter 1977: 70; Dmitricnko, Petrenko 1976: 18; Arnoldi, Dlussky 1978:
531; Collingwood 1979: 57; Kupyanskaya 1979: 128; Espadaler 1981b: 189; Kupyanskaya 1986b:
94; Agosti and Collingwood 1987a: 53, 1987b: 267; Seifert 1988: 7; Atanassov, Dlussky 1992: 87;
Radchenko 1994b: 43, 1994c: 139, 1994f: 81, 1994g: 109; Seifert, 1994: 13; Wu and Wang 1995:
94; Bolton 1995: 283; Seifert 1996: 230; Radchenko et al. 1997: 485; Wei С et al. 2001: 561;
Czechowski et al. 2002: 21; Radchenko and Elmes 2003a: 239; Radchenko, Czechowska and
Czechowski 2004: 47; Radchenko 2005: 144; Seifert 2007: 203; Pfeiffer et al. 2007: 5; Radchenko
2007a: 28, nec Collingwood 1976: 301; Kim B.-J. 1996: 179; 2003: 2, misidentifications.
Myrmica rubra r. (subsp.) sulcinodis: Forel 1874: 76; Emery and Forel 1879: 460; Wheeler W.
M. 1908b: 406.
Myrmica perelegans Curtis, 1854: 214, w, q, m, England. Synonymy by Mayr 1855: 408; Smith F. 1855:
120; Radchenko 1994f: 81; Bolton 1995: 281; confirmed here.
Myrmica sulcinodis var. nigripes Ruzsky, 1895: 32, w, q, m, Russia (also described as new by Ruzsky,
1896: 73); 1905: 707; Finzi 1926: 88, m; Ruzsky 1936: 95; Weber 1948: 268; Sadil 1952: 244;
Arnoldi 1970: 1839. Synonymy by Seifert 1988: 7 {part.); Radchenko 1994f: 81; Bolton 1995: 281;
confirmed here.
Myrmica sulcinodoscabrinodis Ruzsky, 1895: 32, w, Russia (also described as new by Ruzsky, 1896:
73), syn. nov.
Myrmica myrmecophila Wasmann, 1910: 516, q, Austria; Stitz 1939: 88; Weber 1948: 270; Bernard
1967: 123. Synonymy by van Boven 1970: 130; Seifert 1988: 8; Bolton 1995: 281; Radchenko and
Elmes 2003a: 239; confirmed here.
Myrmica sulcinodis var. sulcinodoscabrinodis Forel, 1915: 29, w, Switzerland (Myrmica rubra var.
sucinodoscabrinodis Forel, 1874: 77, nomen nudum) (unresolved junior primary homonym of Myr­
mica sulcinodoscabrinodis Ruzsky, 1895, above); Soudek 1923: 111; Weber 1948: 270. Synonymy
by Bernard 1967: 121; Radchenko 1994f: 81; Bolton 1995: 284; confirmed here.
Myrmica ruginodis var. sulcinodoruginodis Donisthorpe, 1915a: 121, w, England (Myrmica rubra var.
sulcinodoruginodis Emery and Forel, 1879: 461; André 1883: 319; Myrmica ruginodis var. sulci-
nodoruginodis: Dalla Torre 1893: 116; Myrmica sulcinodoruginodis: Emery 1908a: 174; Myrmica
sulcinodis var. sulcinodoruginodis: Finzi 1926: 88; Weber 1948: 268, nomina nuda). Synonymy by
Bernard 1967: 121; Bolton 1995: 284 (see Notes below).
Myrmica sulcinodis derzhavini Ruzsky, 1920: 76, q, Russia. Synonymy by Kupyanskaya 1986b: 94;
Radchenko 1994f: 81; Bolton 1995: 278; confirmed here.
Myrmica sulcinodis eximia Kupyanskaya, 1990: 104, w, q, m, Russia; Bolton 1995: 283. Synonymy by
Radchenko 1994f: 81; confirmed here.
Myrmica sulcinodis vicaria Kuznetsov-ugamsky, 1928: 33, w, Russia; Weber 1948: 270; Radchenko
1994f: 9 1 ; Bolton 1995: 284, syn. nov. (provisionai).
Junior synonym of M. gallienii: Bernard 1967: 121; not confirmed, see Notes to M. gallienii, above.

Type localities and type specimens. M. sulcinodis: "Ad Helsingfors hanc speciem passim inventi".
M. perelegans: "This new and elegant species I found in July 1850, under a stone, on a heath near
Bournemouth in Hampshire". M. sulcinodis var. nigripes: "Найдена на хребте Б. Кумач" [Found on the
Range В. Kumach] (Ruzsky 1895); "Höhenrücken Kumatsch im orenburgischen Ural" (Ruzsky 1896).
M. sulcinodoscabrinodis Ruzsky: "Найден на хр. Ум-Таш; Пермская губерния и уезд, окр. Полазнснского
зав., на гипсовых утесах р. Камы" [Found on the Range Ui-Tash; Permskaya Prov., Permsky Distr., near
Polaznensky, on the gypsum rocks ok Kama river] (Ruzsky 1895); "Bergrücken Ui-Tasch (Orenbu-
rgischer Ural); Gouvernement Perm." (Ruzsky 1896). M. myrmecophila: "... anormale Wiebchenform

294
aufmerksam zu machen, die ich bei Myrmica sulcinodis am 27 August 1891 auf der Höhe des Arl-
bergpasses an der Grenze von Tirol und Vorarlberg (1800 m)". M. sulcinodis var. sulcinodoscabri­
nodis Forel: "Maloja, Swiss"'. M. ruginodis var. sulcinodoruginodis: "I have seen workers from the
colony, discovered at Leith Hills by Arnold on April 18th, 1909, which may be said to represent this va­
riety" [e.g. sulcinodoruginodis]. M. sulcinodis derzhavini: "... одна бескрылая самка, добытая А.
Державиным на берегу р. Еловки, в 30 в. ниже с. Еловки (28.VIII.1908 г.)" [... one dealate queen, col­
lected by A. Derzhavin on the bank of the riv. Elovka, 30 km lower vil. Elovka (28.viii.1908)].
M. sulcinodis vicaria: "Местонахождение: Сучан" [Locality: Suchan].M. sulcinodis eximia: "Материал.
Голотип - рабочий, Хабаровский край, Советская Гавань, 10 км от noe. Тулучи, 24.VIII 1972 (Купянская),
паратипы - 35 рабочих, из той же серии; Чсгдомын, 16.VII 1984 (Купянская), 13 рабочих, 6 Ç, 1 $;
Амурская обл., Климоуцы, 11.VIII 1977 (Купянская), 19 рабочих; Бурятская АССР, Боярский, 6.VIII 1977
(Купянская), 31 рабочий, 1 Ç, 10 S; Култушнос, 24 рабочих; Кяхта, 27.VII 1977, 27.VIII 1979
(Купянская), 52 рабочих, 4 ?, 9 ¿; Сахалин, Оха 12.VII 1981 (Купянская), 1 рабочий" [Material. Holo­
type -worker, Khabarovsky Krai, Sovetskaya Gavan', 10 km from Tulucha, 24.viii.1972, leg. Kupyan­
skaya; paratypes - 35 w from the same nest series; Chegdomyn, 16.vii.1984, leg. Kupyanskaya, 13 w,
6 q, 1 m; Amurskaya Prov., Klimoutsy, ll.viii.1977, leg. Kupyanskaya, 19 w; Buryatskaya ASSR, Bo-
yarsky Distr., 6.viii.l977, leg. Kupyanskaya, 31 w, 1 q, 10 m; Kultushnoe, 24 w; Kyakhta, 27.vii.1977,
27.viii.1979, leg. Kupyanskaya, 52 w, 4 q, 9 m; Sakhalin, Okha, 12.vii.1981, leg. Kupyanskaya, 1 w].
Material examined. Lectotype of M. sulcinodis (designated by Radchenko 2007a), w (bottom
specimen on the pin with 3 w), "H:fors", "W. Nyland.", "Coll. Nyland.", "Mus.-Zool. H: fors Spec. typ.
No. 5054 Myrmica sulcinodis Nyl." (HELSINKI); paralectotypes: 2 w on the pin with lectotype; 3 w
with the same labels as lectotype, but type No. 5055 (HELSINKI); syntypes of M. perelegans, 4 w, 2
q, 2 m, "perelegans Curt." (MELBOURNE); syntypes of M. sulcinodis var. nigripes, 1 m, "M. sulci­
nodis Nyl. v. nigripes R. $ xp. Б. Кумач, 23.VII.94, M. Рузский" [M. sulcinodis Nyl. v. nigripes R. m range
В. Kumach, 23.vii.94, M. Ruzsky] (MOSCOW);'2 w with the same label (PETERSBURG); 1 m, 1 w,
"Ural, Uj-Tash No 29 M. sulcinodis v. nigripes, m." (GENOA); holotype of M. myrmecophila, w -
("ergatoid queen" sensu Wasmann), "Arlberg 27.8.91 G. E. Hol.", "Myrmica myrmecophila Wasm.
sp., п.", "b. Myrmica sulcinodis Nyl.", "Type", "= Ergatoide ouvrier de M. sulcinodis Nyl. det. Van
Boven, 1969", "ergatoide femelle?" (MAASTRICHT); lectotype of M. sulcinodis var. sulcin­
odoscabrinodis Forel (designated here), w (upper specimen on the pin with 3 w), "Typus", "M. sulci­
nodis scabrinodoides [Sicì] w, Malogyia", "M. sulcinodoscabrinodis Forel", "Coli. Forel" (GENEVA);
paralectotypes: 2 w on the pin with lectotype; 2 w with the same labels (GENEVA); holotype of
M. sulcinodis eximia, w, "Хабар, кр., Совгав. р-н, 10 км от п. Тулучи, лиственничник, пр. пл. 8-72, поляна,
под камн. 24.VIII.72, Купянская А. Н." [Khabarovsky Krai, Sovetskaya Gavan', 10 km from vil. Tu­
lucha, Larix forest, plot 8-72, glade, under stone, 24.viii.72, leg. A. N. Kupyanskaya (MOSCOW);
paratypes: several tens of w, q and m from Khabarovsk and Amur Regions, and Buryatia (MOSCOW,
VLADIVOSTOK); non-type material: several hundred of w/about 100 q and m from the whole area..
Distribution (Map 117). Boreal species, distributed from the Atlantic Ocean till the
Pacific Ocean; to the north reaches forest-tundra zone, to the south on plains goes to the
southern limit of the taiga zone, in the more southern regions lives only in mountains; is
absent in Japan.
Etymology. M. sulcinodis: from a combination of the Latin words sulcus = ridge or fur­
row, and nodus = knot or lump, to describe the "deeply wrinkled" sculpture of the petiole
and postpetiole. M. perelegans: from a combination of the Latin words per = extra or very,
and elegans = elegant, to describe the generally "very elegant" appearance of this species.
M. nigripes: from a combination of the Latin words niger = black and pes = foot, probably
to describe the dark patches or "footmarks" present on this species. M. sulcinodoscabri­
nodis: a combination of the names sulcinodis and scabrinodis, to indicate this species ap­
pears intermediate between M. sulcinodis and M. scabrinodis. M. myrmecophila: from the
genitive of the ancient Greek for ant (/№/)/«/£),./.трцпкос {murmëkos) = pertaining to ants

295
combined with phila = lover, derived from the Greek <piloç (philos), to indicate that the au-
thor believed this species to be an "ant-lover" or parasite. M. sulcinodoruginodis: a combi-
nation of the names sulcinodis and ruginodis, to indicate this species appears intermediate
between M. sulcinodis and M. ruginodis. M. derthavini: named for the collector, the fa-
mous Russian zoologist, ichthyologist, hydrobiologist Alexander Nikolaevich Derzhavin
(sometimes spelt Derjavin) (1878-1963). M. eximia: from the Latin word eximia = excel-
lent, to describe the striking or excellent appearance of this species. M. vicaria: from the
word vicarious in the sense of occurring in an abnormal place (derived from the Latin word
vicis = change), probably to indicate that the author considered the location unusual for
this species in the biogeographic sense of vicariance.
Notes. M. sulcinodis belongs to the lobicornis species group. It is a very distinct, obvi-
ous and attractive species as the synonymous names perelegens and eximia suggest; usu-
ally it is a rather large species (for Europe) that often appears "glossy", being distinctly
bicoloured with a dark head, gaster, tibiae and tarsi, and a dark brick-red thorax, waist and
usually femurs. More formally, M. sulcinodis is characterised by its coarse regular longitu-
dinal rugosity on the alitrunk and waist, short petiole without peduncle and with a steep and
straight anterior face, and by the shape of its scape that is strongly, but usually gradually
curved at the base (at most being slightly angulate) but without vertical dent or lobe (at
most with a slight longitudinal ridge on its foot). M. sulcinodis most resembles
M. ademonia, but differs from the latter by having shorter propodeal spines and coarser ru-
gosity on the petiole and postpetiole.
Based on the investigation of the syntypes (workers, gynes and males) of M. perelegans,
preserved in Curtis' collection in MELBOURNE, we confirm earlier synonymy of this
species with M. sulcinodis.
The syntype workers of var. nigripes have blackish patches on the alitrunk. While the
colouration is usually distinctly two-toned as outlined above, specimens (even whole
populations) can be found where the ants are more or less dusky black or uniformly some

Map 117. Distribution of M. sulcinodis.

296
shade of dark red or brown and frequently specimens with black patches on the body can be
found. Therefore we consider variation in coloration to be useless as the basis of the descrip-
tion of a separate form and confirm the synonymy of subsp. nigripes with M. sulcinodis.
The holotype worker (not "ergatoid queen"!) of M. myrmecophila has a grossly enlarged
gaster and minute central ocellum; by all other features it is indistinguishable from M. sul-
cinodis. We are almost certain that it is a worker of M. sulcinodis, infested by Mermis
parasite (see also Radchenko and Elmes 2003a; Czechowski, Czechowska and Radchenko
2007; Czechowski, Radchenko and Czechowska 2007).
We investigated lectotype and 4 paralectotype workers of M. sulcinodis var. sulci-
nodoscabrinodis Forel (see above) and we have no doubt that they are M. sulcinodis. They
are distinctly bicoloured with a brownish-red alitrunk and waist, and brown head dorsum;
also their scape is distinctly angled at the base. We also examined 5 workers from Bon-
droiti collection (BRUSSELS), labelled as: "Engadine Coll. Forel Coll. Emery", "Myrmica
sulcinodoscabrinodis Forel type", "type", "R. I. Sc. N. B. I. G. 21.400", "sulcinodis var. sul-
cinodoscabrinodis For. Engadine cotype Emery, 1919"; "Piz Surlei, Engadino", "R. I. Sc.
N. B. I. G. 21.400". They are M. sulcinodis, but despite they have "type" labels, they can
not belong to the type series, because their locality data do not correspond with that given
by Forel in the original description (see above).
The use of the name "sulcinodoruginodis" in the literature is quite misleading: Emery
and Forel (1879), André (1883), Dalla Torre (1893), Emery (1908a), Finzi (1926), and
Weber (1948) attributed it to Forel (1874). However, in that monograph, this name was
never used formally, Forel only wrote: "Certaines variétés de la M. sulcinodis sont si rap-
prochées de la M. ruginodis que je suis presque certain qu'il existe des intermédiaires entre
ces deux races" (loc. cit., p. 77). The reasons why Emery and Forel (1879) used the name
are not clear, but probably all later authors simply followed them. Nevertheless, all these
records should be considered as nomina nuda.
Donisthorpe (1915a) also attributed this name to Forel (1874) but he actually provided
the first description of M. ruginodis var. sulcinodoruginodis, thus he is the author of this
form: "In these specimens the frontal area is slightly longitudinally striate (which is more
pronounced in some individuals than in others), and the scape of the antennae is slightly
more bent at the base than in typical ruginodis, otherwise the shape, colour, and general
appearance is that of the latter, to which species they are most nearly related" (loc. cit.,
p. 122). Bernard (1967) synonymised the name sulcinodoruginodis with M. sulcinodis,
but he also attributed this variety to Forel (1874), e.g. his "sulcinodoruginodis" is also
nomen nudum. However, based on the brief Donisthorpe's description (see above), his
"sulcinodoruginodis" is a synonym of M. ruginodis rather than of M. sulcinodis, but this
question could be definitely resolved only after finding the type specimens described by
Donisthorpe.
Kupyanskaya (1990) described a new subspecies, M. sulcinodis eximia, from the
eastern part of the area of M. sulcinodis. This form differs from the "typical" M. sulcinodis
in having thicker and inwards-curved propodeal spines in the workers. However, this fea-
ture is quite variable and in Siberia, even in the same nest, some workers have straight
spines and others have curved ones. Individuals with curved spines can also be found
occasionally in west Europe. We can see no reason to divide M. sulcinodis into subspecies
on this character and therefore, confirm the synonymy.
Ruzsky (1895: 32) described from workers from Ural Region "intermediate form"
M. sulcinodoscabrinodis (Myrm. sulcinodis Nyl. f. intrm. M. sulcinodoscabrinodis m.);

297
this name was completely forgotten and ignored by all subsequent authors. Types of this
form are most probably lost and the description is very brief:
"1) posterior surface of propodeum with sparse transversal rugae; scape at the bend
with small oblique lobe. By the other features - as M. sulcinodis. Found on the Range
Ui-Tash.
2) surface of propodeum between spines transversally rugulose; by the other features -
as M. sulcinodis. Permskaya Prov., Permsky Distr., on the gypsum rocks ok Kama river"
(our translation from Russian).
Based on the description, the specimens from Perm almost certainly belong to
M. sulcinodis, but, if their description is correct, workers from Ui-Tash have small vertical
lobe at the scape bend, and may belong to M. lobicornis or even to M. deplanata. Conse-
quently, we consider synonymy of M. sulcinodoscabrinodis with M. sulcinodis as provi-
sional until such time as the types are located.
Ruzsky (1920: 76) described M. sulcinodis derthavini based on the single queen from
Kamchatka. This specimen is lost. It apparently differed from "typical" M. sulcinodis by:
"propodeal spines very short and wide at the base. Scape at the base strongly curved. Colour
darker" (our translation from Russian). Most probably this queen was simply a somewhat
aberrant specimen of M. sulcinodis and the features described are not sufficient to distin-
guish a different subspecies.
Kuznetsov-ugamsky (1928: 33) described M. sulcinodis vicaria from workers from Pri-
morsky Krai of Russia. The types of this "subspecies" are lost, and its description is almost
useless: "Propodeal spines shorter than its dorsal surface, narrowed at the base and sharp
at the tips. Body length smaller than in typical form, 4-4.5 mm (vs. 4.8-6 mm). Posterior
surface of propodeum between spines smooth and shiny. Propodeal lobes directed upwards
and sharp. Body brownish-red. Head and gaster darker" (our translation from Russian).
Based on this description, we exclude synonymy of this species with M. ademonia, which
has longer propodeal spines, but can not exclude the possibility that M. sulcinodis vicaria
is conspecific with some other bicoloured species, e.g. M. kamtschatica or M. displicentia.
Therefore, until such time as the type specimens may be found, we consider synonymy of
subsp. vicaria with M. sulcinodis as provisional.
Of two specimens from Sam-zi-yan (North Korea) determined by Collingwood (1976)
as M. sulcinodis, one is M. ruginodis and the other is M. ademonia (material is in
BUDAPEST, examined).
Ecology. M. sulcinodis has been comparatively well studied. In general it favours habi-
tats that have a coarse sandy subsoil that holds a certain amount of moisture and does not
completely dry out in dry periods. In northern habitats it is especially common on areas of
moorlands and heathlands having the correct soil type, and is also common in open coni-
ferous forests providing the soil is suitable. In the far north it sometimes nests underwood
but in southern moorlands it normally builds its nest directly into the soil with few above-
ground signs except in early spring when colonies may make small solaria. It is very com-
mon in certain sites in alpine meadows on southern mountains where it nests under stones.
Many populations are highly polygynous and several queens can often be seen when a stone
is turned over. Sexuais are produced in early summer and nuptial flights occur in August
and September. Few flights have been observed and we suspect that generally these ants do
not fly far but join small very local swarms.
In one of the first detailed population studies of this species Elmes (1974c) showed that
nests of a large population living in moorland in Scotland, averaged 440 workers (mean of

298
40 nests) and the nests were effectively monogynous with half having no queen. Males were
produced by about half the nests (average of 28 per nest) and gynes were produced by the
20% largest nests (average 65 per nest) giving an overall population sex-ratio of about 1:1.
In a later study of populations on heathlands in southern England, Elmes (1987a, b) showed
that M. sulcinodis are not always monogynous but can be highly polygynous with the num-
ber of queens varying from year to year. This data collected in peculiar ecological circum-
stances following a severe fire, showed that queen numbers cycled over a 5 year period in
a regular fashion and that the cycle persisted for many years (Elmes and Keller 1993). In
these populations nests were smaller averaging 163 workers and 3.7 queens per colony,
weather seemed to play a part in year to year variation in worker numbers but not in queen
numbers. The suggestion was that almost all the new queens are produced by a small num-
ber of large nests but almost all are randomly recruited back into all the nests of the popu-
lation. Many more colonies reared males so that the sex-ratio in these populations were
heavily male biased. These populations produced a great many gynandromorph mosaics
with different proportions of the body being male. The only other species where we have
found such a high proportion of such intercastes is M. gallienii (see ecological notes to that
species).
These studies were taken up and extended to a similar population living on heathland
in Denmark, by Prof. ]. J. Boomsma and co-workers using molecular genetical techniques
(Boomsma 1993; Pedersen and Boomsma 1998, 1999a, b). Seppä et al. (1995) studied
facultative polygyny in relation to habitat succession.
In the laboratory there have been a few studies of the glandular secretions of
M. sulcinodis (e.g. Attygalle et al. 1983; Cammaerts et al. 1983) and cuticular hydrocarbons
(Elmes et al. 2002). The chromosome number of M. sulcinodis is 56, somewhat more than
most other Myrmica species (Hauschteck 1965). Both workers and queens were shown to
lay trophic eggs (Wardlaw and Elmes 1995,1998) and queens can suppress the development
of sexual larvae in the same way as M. rubra (Elmes and Wardlaw 1983b).

Myrmica symbiotica (Menozzi, 1925)

(Fig. 276)

Sommimyrma symbiotica Menozzi, 1925: 25, ergatoid q (?), Italy; Arnoldi 1930: 270; Baroni Urbani
1971: 74; Kutter 1973:263.
Myrmica symbiotica: Bolton 1988: 3; 1995: 284; Radchenko and Elmes 2003a: 224.

Type localities and type specimens. "Monte Calvanella, poco al disopra della strada rotabile. Fanano
Pievepelago. Ho raccolto questa caratteristica nuova formica in uno solo esemplarte e dentro ad un nido
di Myrmica rubra subsp. laevinodis".
Material examined. Holotype "gynomorph worker", "Sestola (Emilia)", "C. Menozzi", "Typus",
"Sommimyrma symbiotica Menoz., Typus" (BOLOGNA).
Distribution (Map 118). It is known only from the holotype from northern Italy.
Etymology. M. symbiotica: from the Greek syn (ovv) = together with, and Mosis
{jiioaid) = way of life, to indicate that the author believed this species to be a symbiont of
Myrmica.
Notes. The combination of features, which distinguish M. symbiotica from other
Myrmica species are: head with 3 minute ocelli; clypeus with big foveolate impression;

299
antennal scape angulated at the base, with a narrow but distinct longitudinal lobe (similar
to those of some M. scabrinodis or M. specioides Bondroit); curved frontal carinae with
relatively narrow frons, but somewhat wider than in M. scabrinodis (FI 0.38, FLI 1.31);
promesonotal dorsum saddle-shape (seen in profile), with the trace of scutum; metanotal
groove very deep and abrupt; petiole and postpetiole without ventral lobes; petiole with
distinct dorsal plate (similar to that of M. scabrinodis); postpetiole wide (PPI4 0.60);
gaster somewhat enlarged; middle and hind tibial spurs strongly reduced, very short, thick,
not pectinate; not a hairy species, petiole with only б long standing hairs. Based on these
characters we placed this species to the scabrinodis-group (Radchenko and Elmes
2003a).
Bolton (1988) made several suggestions regarding the unusual holotype specimen of
M. symbiotica: it is an ergatoid queen or gynecoid worker; perhaps an aberrant specimen
of M. rubra; or possibly it is an ergatoid female of another Myrmica species that was caught
by workers of M. rubra and taken to their nest as prey.
Based of the examination of the holotype, we suggested that it is a pseudogyne worker
rather than ergatoid queen. The shape of its alitrunk and the presence of minute ocelli make
it most similar that of the gynomorphic workers of M hirsuta, so that it could be the worker
form of a social parasite with unknown queens. It certainly is not an aberrant specimen of
M. rubra. The most peculiar feature, never seen in any other Myrmica so far, is the big fove-
olate impression on the clypeus, but with only a single specimen this feature could simply
be some type of aberration. If it is an aberrant specimen of a scabrinodis-group species
(most probably M. scabrinodis), then it was probably created by Mermis parasite infestation.
If so, how did it get into a M. rubra colony, one of the most aggressive Myrmica species in
the Paiaearctic? Bolton's (1988) suggestion, that it could be specimen taken as prey is an
attractive possibility (see also Radchenko and Elmes 2003a).
Thus, the taxonomic and "biological" position of M. symbiotica remains unclear.
Ecology is unknown, except the holotype was found in a nest of M. rubra.

Map 118. Distribution of M. symbiotica.

300
 Myrmica taibaensis Wei, Zhou et Liu, 2001
(Figs 277, 278)

Myrmica taibaensis Wei С, Zhou et Liu, 2001: 562, w, q, China; Radchenko and Elmes 2009a: 69.

Type localities and type specimens. "Holotype worker, Mt. Taibai 2200 m, Taibai Co., Shaanxi
Prov., nested in rotten trunk of tree, in the forest, 19 Aug. 1997, Wei Cong. Paratypes: 35 workers, 5
females, Mt. Taibai 2 000 m, in rotten root in the forest, 27 Aug. 1997, Wei Cong".
Material examined. Paratypes, 3 w, 1 q, "Taibai Mountain, Shaanxi Prov., July 19 1997, Cong Wei
leg.", "Myrmica taibaensis Wei et al., Cong Wei det. (N.W. Sci.-Tech. Univ. Agri. Forest.)" (XI-
ANYANG).
Distribution (Map 119). It is known only from the type series from the Shaanxi Prov.
of China.
Etymology. M. taibaensis: a combination of the name Taibai with the Latin suffix ensis
= place of origin, to indicate that it was found on Mt. Taibai, the main peak of the Qinling
Mountain, Shaanxi Province, China.
Notes. We placed M. taibaensis to the pachei species group and redescribed workers
and queen (Radchenko and Elmes 2009a). Wei С et al. (2001, p. 563) noticed: "The new
species is close to M. inezae. It differs from the later by the shape and number of mandible
teeth, the shape of propodeal spines, and the colour of body". Although in some respects
there are similarities between the inezae- and pac/iei-groups (see Radchenko and Elmes
2001b), the authors were certainly referring to its similarity to M. zveii, which they had
misidentified as M. inezae (see also Radchenko et al. 2008).
The mesonotal and propodeal dorsum of M. taibaensis is transversally rugose, while the
pronotal dorsum is reticulate; the occipital and lateral margins of the head above the eyes
have numerous short subdecumbent hairs; the propodeal spines are long, but massive,
strongly widened at the base.

Map 119. Distribution of M. taibaensis.

301
 Ecology is poorly known. Wei С et al. (2001) noted that two samples were collected
from the nests in a rotten tree trunk and rotten [tree] root, in mountain forest at an altitude
between 2000 and 2200 m a.s.l.

Myrmica tamarae Elmes, Radchenko et Aktac, 2002

(Figs 279, 280)

Myrmica tamarae Elmes, Radchenko et Aktac, 2002: 165, w, q, Turkey.

Type localities and type specimens. Turkey, for details see material examined.
Material examined. Holotype, w, "Turkey, Erzurum Region, Palandöken dag, alt. 2250 m, No.
2140, 25.ix.1979, leg. N. Aktaç" (EDIRNE); paratypes: 15 w, 1 q from the nest of holotype; 16 w, the
same locality, date and collector, alt. 2800 m, No. 2142 (EDIRNE, WARSAW, KIEV, ELMES).
Distribution (Map 120). It is known only from the type series from the Erzurum Prov.
of Turkey (East Anatolia).
Etymology. M. tamarae: this species was dedicated to Tamara Radchenko, wife of
Alexander Radchenko.
Notes. M. tamarae belongs to the excelsa-complex of the lobicornis species group, and
most resembles M. transsibirica differing from it by its somewhat coarser rugosity on the
frons, having not more than 10 sinuous rugae between the frontal carinae level with the
eyes vs. > 13 in M. transsibirica.
Ecology. Little is known about its ecology. M. tamarae is probably quite localised in its
distribution having being found only on high mountain meadows (2250 and 2800 m) in
East Anatolia.

Map 120. Distribution of M. tamarae.

302
 Myrmica tenuispina Ruzsky, 1905
(Figs 281-283)

Myrmica laevinodis var. tenuispina Ruzsky, 1905: 670 (first available use of Myrmica rubra r. laevi­
nodis var. tenuispina Forel, 1904a: 374, w {part.), Uzbekistan, Tadjikistan; Emery 1908a: 170,
Emery 1921: 39; Kuznetsov-ugamsky 1927: 193, unavailable names); Weber 1947: 455.
Myrmica tenuispina: Arnoldi 1976: 551, m; Radchenko 1994b: 42, 1994c: 142, 1994d: 81; Bolton
1995: 284; Radchenko and Elmes 2001b: 258, 2002: 35, q; Schultz et al. 2006: 206, nec Tarbinsky
1976: 25; Dlussky et al. 1990: 184, misidentifications.

Type localities and type specimens. "Ferghana (fi. Kugart, 6-8000' h., 6 w, 5.VIII.1896. Korzin-
skij!); Buchara orient. (Karategin, Kala-i-choit, 2 w, 21.VII; Tabi-dara - Zagyrdescht, 15 w, 17.VI;
Darvas, Tash-Kurgan, 18 w, 22.VIII.1897. Kaznakov!)" (see also Notes below).
Material examined. Lectotype (designated by Radchenko and Elmes 2001b), w (uppers specimen
on the pin with 4 w), "Таби-дара - Загырдсшт, в. Бухара, 17.VI.97, Казнаков" [Tabi-dara - Zagyrdesht,
E. Bukhara, 17.vi.97, leg. Kaznakov], "Myrmica rubra L. r. laevinodis var. tenuispina Forel det, w,
type" (MOSCOW); paralectotypes: 3 w on the pin with lectotype; 3 w with the same labels
(MOSCOW); 8 w with the same labels (PETERSBURG); 1 w with the same labels (GENOA); 2 w,
"Кала и Хонт, Каратсгин, в. Бухара, Казнаков, 21.VI.97" [Kala i Khont, Karategin, E Bukhara, leg. Kaz­
nakov, 21.vi.97] "Myrmica rubra L. r. laevinodis var. tenuispina Forel det, w, type" (PETERSBURG)
(see also Notes below); non-type material: 35 w, 4 q, 3 m from Uzbekistan, Kyrgyzstan, Tadjikistan,
and Afghanistan.
Distribution (Map 121). Tien-Shan, Alai, Pamir, NE Afghanistan.
Etymology. M. tenuispina : from a combination of the Latin words tenuis = thin or slen­
der, and spina = thorn, to describe the shape of the propodeal spines.
Notes. Radchenko (1994b) placed M. tenuispina in the scabrinodis species group, but
more recently we revised this opinion and transferred it to the tibetana-gvoup (Radchenko
and Elmes 2001b). M. tenuispina most resembles M. tibetana but differs from it by its much
longer propodeal spines.
Superficially, workers and queens of M. tenuispina are similar to the species of rubra-
or smythiesii species groups, but well differs from them by the shape of the frontal carinae
that are strongly curved outwards in their anterior third, to form distinctive wide, subsquare
frontal lobes; as a result the frons is relatively wide (FI is subequal to that of the species from
rubra- or smythiesii- species groups) but FLI is distinctly larger (> 1.25 vs. < 1.18). On the
other hand, the males clearly differ from these groups by having a relatively short scape (SIj
< 0.40) (see also Notes to M. tibetana, below).
Forel described var. tenuispina based on workers from four nest samples, collected in
the modern Uzbekistan ("Fergana") and Tadjikistan ("Buchara orient") (see above). We
examined workers from all the type series, preserved in MOSCOW and PETERSBURG,
and found that in fact they included a second species (M. ferganensis). To avoid further
misunderstanding, we designated as the lectotype of M. tenuispina a worker from Tajikistan
(Tabidara - Zagyrdesht, see above). Material from Kala i Khont also belongs to M. tenui­
spina, and we considered specimens with corresponding labels as paralectotypes.
The following "type" specimens had frontal lobes that were much less extended than
those of the lectotype and paralectotypes and by this and other features they correspond to
M. ferganensis, thus we exclude them from the type series of M. tenuispina. Most proba­
bly, the erroneous treatment of "M. tenuispina" by Tarbinsky (1976) and Dlussky et al.
(1990) was provoked by study of these "type" specimens from Tash-Kurgan and Kugart:

303
 Map 121. Distribution of M. tenuispina.

- 13 w, "Таш-Курган, Дарваз, в. Бухара, 22.VIII.97, Казнаков" [Tash-Kurgan, Darvaz,

E Bukhara, 22.viii.97, leg. Kaznakov], "Myrmica rubra L. r. laevinodis var. tenuispina Forel
det, w, type" (MOSCOW, PETERSBURG);
- 6 w, "Кугарт, 6-8000', Ферганской обл., Коржин., 5.VIII.95" [Kugart,
6-8000', Fergana obi., leg. Korzhinsky, 5.viii.95], "Myrmica rubra L. r. laevinodis var.
tenuispina Forel det, w, type" (PETERSBURG).
In Arnoldi's (1976: 551) paper there is a printing error: the symbol "worker" is printed
as "queen", which led Bolton (1995) to note that queen was described by Arnoldi. In fact,
he redescribed workers (based on the type series from Tabi-Dara, loc. cit., p. 552) and de­
scribed males for the first time; queens were described first by Radchenko and Elmes (2002).
Ecology. M. tenuispina is associated with the mountains of Middle Asia and NE
Afghanistan, where lives on subalpine meadows at altitudes between 2700 and 4100 m and
nests in the soil and under stones.

Myrmica tibetana Mayr, 1889

(Figs 284, 285)

Myrmica tibetana Mayr, 1889: 279, w, Tibet; Dalla Torre 1893: 118; Forel 1903: 697; Ruzsky 1905:
661; Emery 1908a: 181; Ruzsky 1915:310, m (?-sec Notes below); Emery 1921: 42; Weber 1947:
465; Radchenko 1994b: 41; Bolton 1995: 284; Radchenko and Elmes 2001b: 257, 2003b: 3, nec
Collingwood 1961: 56, misidentification.

Type localities and type specimens "Aprii 1884, Jumel-Kuku-Gebirge; Mai-Juni 1884, Tibet
septentr."
Material examined. Lectotype (designated by Radchenko and Elmes 2001b), w, "Tibet" (previ­
ously labelled as paratype) (BERLIN); non-type material: 15 w, 5 m (?) from Tibet.
Distribution (Map 122). Tibet.

304
 Etymology. M. tibetana: named for Tibet, the region where the type material was
collected.
Notes. M. tibetana typifies the tibetana species group (Radchenko 1994b erroneously
placed it to the ruftra-group). It has, like M. tenuispina, frontal carinae that are strongly
curved outwards in their anterior third, and wide, subsquare frontal lobes, but differs from
the latter species by its short propodeal spines. The modem treatment of M. tibetana is con­
firmed by the investigation of the lectotype specimen (see above).
We investigated the material (9 w, 3 m, including "androtype"), determined by Ruzsky
as M. tibetana (MOSCOW, PETERSBURG) and labelled as ""ущ. Хату, сев. Бурхан Будда,
10,800' Козлов, кон. vii.01" [gorge Khatu, N of Burkhan Budda, 10,800', leg. Kozlov, end
of vii.01] (these data correspond with those given by Ruzsky 1915). The workers fully cor­
respond with the lectotype of M. tibetana.
We placed M. tibetana and M. tenuispina in the same group because the workers of
both species have uniquely shaped frontal carinae and frontal lobes from among the Old
World Myrmica species (Radchenko and Elmes 2001b). However the males of these two
species have quite different length scapes, those of M. tibetana are distinctly longer than
those of M. tenuispina (SIj > 0.75 versus < 0.40, respectively). However, the males of both
species were described later than the original description of the species (by Ruzsky 1915 and
Arnoldi 1976 respectively). From the published data and data-labels we are not sure whether
they belong to the same nest samples as the conspecific workers, or were just collected
separately in the same localities. We suspect at least one series of males do not belong to
the species to which they were ascribed because it would be most unusual for the males of
two species that have such similar workers to have such different length of scape. This prob­
lem will be resolved when somebody collects the males and workers of M. tibetana and
M. tenuispina from the same nest.
Collingwood's record of this species for Afghanistan in fact belongs to M. afghanica
(see Radchenko and Elmes 2003b and above).

Map 122. Distribution of M. tibetana.

305
 Radchenko (1994b) synonymised var. furva with M. tibetana, but now we consider it
as a junior synonym of M. bactriana (see above).
Queens are unknown.
Ecology is unknown, except that is was found at subalpine meadows at altitudes over
3000 m a.s.l.

Myrmica titanica Radchenko et Elmes, 2001

(Fig. 286)

Myrmica titanica Radchenko et Elmes, 2001a: 222, w, Vietnam; Radchenko, Elmes and Viet 2006: 39.

Type localities and type specimens. Northern Vietnam, for details see material examined.
Material examined. Holotype, w, "Northern Vietnam, Lao Cai, Sa Pa, Fan Sipan, alt. 2020 m, Aprii
1998, leg. B. T. Viet" (KAGOSHIMA); paratype, 1 w with the same label, probably from the same nest
(ELMES); non-type material: Northern Vietnam, 1 w, Lao Chau province, western slopes of Mt. Fan
Si Pan (W Cong Troi), 2100-2200 m, 6.V.2002, leg. K. Eguchi; 11 w, CongTroi, Sa Pa district, Lao Cai
province, 2000-2200 m, 28-iv-2.v.2002, leg. K. Eguchi.
Distribution (Map 123). Northern Vietnam.
Etymology. M. titanica: from the word titanic = colossal or gigantic derived from the
Titans, the "Older Gods" of Ancient Greece, used here to describe the extraordinary size
of this species compared to other Myrmica.
Notes. M. titanica belongs to the ntee-complex of the ritae species group. Except for
M. angulata, it differs from all other species of the ritae-complex by the punctated surfaces
between the rugae of the petiole and postpetiole. In addition, it is the largest Myrmica
species in the World so far discovered, having HW > 1.80 mm, HL > 2.10 mm and AL >
3.30 mm (for details see Radchenko and Elmes 2001a; Radchenko, Elmes and Viet 2006).
Queens and males are unknown.

Map 123. Distribution of M. titanica.

306
 Ecology is little known. Almost all specimens have been collected foraging on the ground
or on fallen trees in well-developed forest. K. Eguchi (pers, comm.) collected workers emerg­
ing from a natural cavity in a tree cut down for timber; unfortunately, he had no tools to cut
into the wood to "excavate" the colony. This led us to suppose that M. titanica might be
a truly arboreal species living and foraging mainly in the canopy, in which case it might be
endangered by excessive logging activities. Specimens were found at altitudes 2000-2200 m.

Myrmica tobiasi Radchenko et Elmes, 2004

(Figs 287-289)

Myrmica tobiasi Radchenko et Elmes, 2004: 224, w, q, m, Kazakhstan, Kyrgyzstan.

Type localities and type specimens. Kazakhstan and Kyrgyzstan, for details see material examined.
Material examined. Holotype, w, "Казахстан, Алма-Атинский заповедник, Талгар, № 749, VIII—1968,
В. Анциферов" [Kazakhstan, Alma-Atinsky Natural Reserve, Talgar, No. 749, viii—1968, leg. V.
Antsyferov] (MOSCOW); paratypes: 2 w, 1 m from the nest of holotype; 12 w, " Казахстан, Алма-
Атинская обл., Зашшйский Алатау, 17.VIII.1972, В. Анциферов" [Kazakhstan, Alma-Ata Region, range
Zailiysky Alatau, 17.viii.1972, leg. V. Antsyferov]; 6 w, "Казахстан, Алма-Атинский заповедник, В.
Анциферов" [Kazakhstan, Alma-Atinsky Natural Reserve, leg. V. Antsyferov]; 7 w, 2 m, "Киргизия,
Чон-Ксмин, с. Новороссийка, 6.VIII.1966, Ю. Тарбинский" [Kyrgyzstan, Chon-Kemin, vili. Novoros-
siyka, 6.VÍÜ.1966, leg. Yu. Tarbinsky]; 1 q, 2 m, "Киргизия, Чон-Ксмин, Калмак-аксу, 7.VIII.1966, Ю.
Тарбинский" [Kyrgyzstan, Chon-Kemin, Kalmakaksu, 7.viii.1966, leg. Yu. Tarbinsky]; 2 w, "Киргизия,
Сары-Чслск, Аркит, Ходжа-Ата No. 63-215, 25.V. 1963, Г. Длусский и Ю. Тарбинский" [Kyrgyzstan,
Sary-Chelek Nature Reserve, Arkit, Khodzha-Ata, No. 63-215, 25.V.1963, leg. G. Dlussky and Yu.
Tarbinsky]; 1 q, "Киргизия, Иссык-Кульская обл., Хаджи-сау, 20.VI1I.1997" [Kirgizia, Issyk-КиГ Prov.,
Khadzhi-sau, 20.viii.1997] (MOSCOW, KIEV).
Distribution (Map 124). Tien-Shan (southern Kazakhstan and Kyrgyzstan).
Etymology. M. tobiasi: this species was dedicated to Prof. Vladimir Ivanovich Tobias, the
well-known Russian hymenopterologist from the Zoological Institute of the Russian
Academy of Sciences, St. Petersburg.
Notes. M. tobiasi belongs to the spec/o/des-complex of the scabrinodis species group. Its
female castes share most features with M. specioides, but also resemble M. scabrinodis in
some respects. The size and shape of the lobe at the base of scape and the extended frontal
lobes are similar to M. scabrinodis but they clearly differ from that species and resemble
M. specioides by having a wider frons (mean FI 0.40 vs. 0.36 for M. scabrinodis), shorter
propodeal spines (mean ESLI 0.35 vs. 0.41) and by a more rounded petiole (M. scabrinodis
has a horizontal or slightly declined dorsal plate that makes a sharp edge with the anterior
surface). Compared to M. specioides the workers are clearly separated by their more
developed and extended frontal lobes (mean FLI 1.40 vs. 1.32) and even wider frons (mean
FI 0.40 vs. 0.38). Males of M. tobiasi clearly differ from those of M. scabrinodis by having
much shorter sub-erect hairs on the tibiae and tarsi (compared to very long, curved erect
hairs), however without female castes it is almost impossible to separate M. tobiasi males
from those of M. specioides.
Ecology. Gennady Dlussky (pers, comm.) collected M. tobiasi in the valley of river
Khodzha-Ata at an altitude of 1400-1500 m. Nests were under stones on meadows and
sandy banks with tall grass and shrubs among open forest, comprising willow and apple
trees. Other species collected included Tetramorium spp., Tapinoma (cf. erraticum),

307
 Map 124. Distribution of M. tobiasi.

Messor structor (Latr.), Lasius sp. (alienus-group), Cataglyphis aenescens (Nyl.) and
Crematogaster sordidula (Nyl.).

Myrmica transsibirica Radchenko, 1994

(Figs 290-292)

Myrmica transsibirica Radchenko, 1994a: 212, w, q, m, Russia; 1994b: 43, 1994c: 139, 1994f: 90,
1994g: 109; Bolton 1995: 284.
Myrmica taediosa Bolton, 1995: 284 (replacement name for Myrmica carinata Kupyanskaya, 1990:
114); Elmes et al. 2002: 167; Radchenko 2005: 145; Pfeiffer et al. 2007: 5; Yamane 2008: 29,
syn. nov.
Myrmica carinata Kupyanskaya, 1990: 114, w, q, m, Russia (South Siberia and Far East) [junior pri-
mary homonym of Myrmica carinata Smith F., 1859: 148 (transferred to Tetramorium Mayr)];
Radchenko 1994b: 43, 1994c: 144, 1994f: 90.

Type localities and type specimens. M. transsibirica: "Holotype worker: Russia, vicinity of Kras-
noyarsk, Pogorelsky bor, No. 29, 1964 (V. Dmitrienko). Paratypes: 11 workers from the same nest; 65
workers, 1 female, Aitai, Nos. 4, 14, 119, 125, 130, 131, 237, 415, 506, 1988 (L. Omelchenko); 13
workers, Aitai, 1968 (Z. Zhigulskaya); 2 females, 2 males, Altai, Kuraiskaya kotlovina, No. 441,
10 VIII 1963 (Z. Zhigulskaya); 6 workers, Aitai, Chuiskaya kotlovina, Nos. 537, 549, 18 and 27 VI
1964 (Z. Zhigulskaya); 1 worker, Aitai, Unzezia on river Katun', 15 VIII 1909 (Gorchakovsky);
1 worker, Chakassia, V 1974; 2 workers, Minusinsk, No. 1286, V 1974; 6 workers, E Siberia, river
Bogdashka, near river Angara, 21 VI 1873 (Gortung); 4 workers, 1 female, 1 male, Ulan-Ude, Nos.
1118, 1124, 1 VIII 1972; 3 workers, Irkutsky distr., village of Zun-Murino, No. 295, 10 VIII 1965
(A. Plcshanov); 10 workers, Chitinskaya obi., Sretensky distr., Ust'-Dunaevo, Pad' Kulinda, 10 IX
1957 (G. Dlussky); 195 workers, Evreiskaya AO, Mt. Obluchje, 40 km S of village of Pashkovo, 2-16
VI 1978 (Toms); 20 workers, 1 female, Amurskaya obi., river M. Para - river B. Ergcl, 5 IX 1958
(Zinoviev); Primorsky Krai: 9 workers, Khankaisky distr., Sintuche, 1 V 1972 (A. Kupyanskaya);

308
9 workers, Barabach-Levada, 9 V 1972 (A. Kupyanskaya); 23 workers, 4 males, Kedrovaya Pad', 5 V
1971, 18 IX 1973, 16 VI 1975 (A. Kupyan-skaya); 2 workers, 1 male, same locality, 6 IX 1964
(G. Dlussky); 8 workers, Tetuchinsky distr., rivers Iman and Krasnaya, 4 IX 1971 (A. Kupyanskaya);
13 workers, Dalnegorsky distr., Tetuche-Pristan', lake Vas'kovka, 26 VI 1972 (A. Kupyanskaya);
7 workers, Pejshula, Zmeinaya Gorka, 21 VII 1969 (A. Kupyanskaya); 11 workers, Sudzuche, buhta
Kut, 5 VIII 1946 (Sharov); Suvorovo, 24 VI 1972 (Zherychin); 4 workers, vicinity of Vladivostok,
5 V 1989 (L. Rybalov); 1 queen, lake Hanka, 2 IX 1986 (A. Kotenko); 4 workers, Sakhalin, vicinity of
Juzhno-Sakhalinsk, 25 VII 1975 (A. Kupyanskaya); 2 workers, Sakhalin, village of Matrosovka, 9 VIII
1986 (S. Basarukin); Kunashir: 3 workers, Alechino, 23 VIII 1975 (Tichomirova); 3 workers, Tretia-
kovo, 21 VIII 1975 (Tichomorova); 11 females, 2 males, lake Goriachee, volcano Golovina, 9 VIII
1970 (Tichomirova)". M. carinata: "Материал. Голотнп - рабочий, Приморский крап, Рудная Пристань,
берег моря, под камнем, 26.VI 1972 (Купянская); паратипы: 55 рабочих из той же серии. Кроме того,
изучено около 800 рабочих, 44 Ç и 27 S из Приморского края (Кедровая Падь, Приморская, Славянка,
Владивосток, Анисимовка, вершина горы Крнничная, Тигровой, Шкотовскос плато, Чандолаз, Лазовский
заповедник, 8 км С Бровнич, Уссурийский заповедник, Таежное, Рсшстниково), Хабаровского края (Хаба­
ровск, 12 км Ю Тумнина, Тулучи, Комсомольск-на-Амурс, Березовый); Амурской обл. (Климоуцы);
Читинской обл. (Александровский Завод); Сахалина (Ново-Алсксандровск, Южно-Сахалинск, Стародубскос,
Озерный, Тымовскос, Алсксандровск- Сахалинский, Ноглики), Курильских островов (о-в Кунашир:
Алсхино, Трстьяково) [Material: Holotype - worker, Primorsky Krai, Rudnaya Pristan', see shore, under
stone, 26.vi.1972, leg. Kupyanskaya; paratypes: 55 workers from the same nest series. Additionally, in­
vestigated about 800 workers, 44 queens and 27 males from Primorsky Krai (Kedrovaya Pad', Pri-
morskaya, Slavyanka, Vladivostok, Anisimovka, top of the Mt. Krinichnaya, Tigrovaya, Shkotovskoe
plateau, Chandolaz, Lazovsky Natural Reserve, 8 km N Brovnich, Ussuriysky Natural Reserve, Taezh-
noe, Reshetnikovo); Khabarovsky Krai (Khabarovsk, 12 km S Tumchin, Tulucha, Komsomolsk-on-
Amur, Berezovyi); Amurskaya Prov. (Klimoutsy); Chitinskaya Prov. (Aleksandrobsky Zavod);
Sakhalin (Novo-Aleksandrovsk, Yuzhno- Sakhalinsk, Starodubskoe, Ozernyi, Tymovskoe, Aleksan-
drovsk-Sakhalinsky, Nogliki); Kurily Islands (isl. Kunashir: Alekhino, Tretiakovo)".
Material examined. Holotype of M. transsibirica: "окрестности Красноярска, Погорельский бор, No.
29, 1964 (В. Дмитриснко)" [vicinity of Krasnoyarsk, Pogorelsky bor, No. 29, 1964 (V. Dmitrienko)]
(MOSCOW); paratypes: > 300 w, 15 q, 9 m (see type localities, above) (MOSCOW, KIEV); holotype
of M. carinata, w, "Прим. Кр., Рудная Пристань, берег моря, под камнем, 26.vi.1972, Купянская" [Pri­
morsky Kari, Rudnaya Pristan', 26.vi.1972, leg. A. Kupyanskaya] (MOSCOW); paratypes: several tens
of w, q and m from Russian Far East (MOSCOW, VLADIVOSTOK); non-type material: about 100 w,
several tens of q and m from South Siberia, Mongolia, Russian Far East, China and North Korea.
Distribution (Map 125). South Siberia (to the west until Kuznetsky Alatau), Russian
Far East, north-east of China, Korean Peninsula, Japan (Hokkaido).
Etymology. M. transsibirica: a combination of the Latin word trans = across or over,
with the name Siberia, to indicate it is widespread throughout Siberia. M. taediosa: from the
Latin word taedios = boredom or weariness, probably Bolton was weary of having to find
replacement names for species. M. carinata: from the Latin word carinata = shell-shaped,
like a keel, to describe the shape of the lateral portions of clypeus that are raised into
sharp ridge in front of antennal insertions and form subvertical carinae (similar to that of
Tetramorium).
Notes. M. transsibirica belongs to the ежсе/sa-complex of the lobicornis species group.
It has a feature characteristic for genus Tetramorium but very rare in genus Myrmica: the
lateral portion of clypeus is raised into a sharp ridge in front of the antennal insertions, so that
antennal sockets are distinctly separated from the clypeal surface. Only two other Myrmica
species have this feature, M. excelsa and M. tamarae. The shape of scape of M. transsibirica
is quite different from that of M. excelsa being distinctly narrower at the bend than on its
midlength (seen in profile), and the vertical (shorter) part has no longitudinal groove and

309
 Map 125. Distribution of M. transsibirica.

lateral ridges (seen in front). It is most similar to M. tamarae by the shape of its head, frontal
lobes and antennal scape, but differs from it by less coarser rugosity on the frons, having
> 13 rugae level with the eyes vs. not more 10 sinuous rugae in M. tamarae.
When Radchenko (1994a) prepared a paper with description of the new Myrmica species
including M. transsibirica, he had not seen Kupyanskaya's (1990) monograph and had not
seen the types of M. carinata Kupyanskaya, for which Bolton (1995) proposed the re­
placement name M. taediosa because the name M. carinata is preoccupied. After examina­
tion of Kupyanskaya's types and additional non-type material, we found that M. trans­
sibirica is conspecific with M. carinata and therefore, had priority over the replacement
nameM. taediosa.
Ecology. As its name implies, this species is widespread species in Siberia, living at
a wide range of altitudes from sea level to 2000 m a.s.l. Usually it lives in various kinds of
forests, but prefers light, sparse and warm ones; it also can be found among shrubs and in
steppe-like habitats. Nests are usually built in the soil, often under stones. Nuptial flights
occur August-September.

Myrmica tschekanovskii Radchenko, 1994

(Figs 293, 294)

Myrmica tschekanovskii Radchenko, 1994a: 215, w, m, Russia; 1994b: 44, 1994c: 13, 1994e: 77, 1994g:
105; Bolton 1995: 284.

Type localities and type specimens. Russia (East Siberia), for details see material examined.
Material examined. Holotype, m, "В. Сибирь, I-I. Тунгуска возле Енисея, 64° 40' с. ш. 5.VIII.1873,
Чскановский" [East Siberia, riv. Nizhniaya Tunguska near riv. Enisei, 64°40'N., 5.viii.1873, leg.
Tshekanovsky] (MOSCOW); paratype: 1 w with the same label (MOSCOW); non-type material: 1 m,
"Якутск (Москвин), VIII.927" [Yakutsk, leg. Moskvin, viii.927] (MOSCOW).

ЗЮ
 Distribution (Map 126). East Siberia.
Etymology M. tschekanovskii: this species was named for the collector, the Russian
(originally Polish) geographer and naturalist Dr. Alexander Lavrentievich Czekanowski
(alternative spelling Tschekanovski) (1833-1876).
Notes. M. tschekanovskii is characterized by the 12-segmented antennae of its males.
Based on this feature, Radchenko (1994a) placed it to the arnoldii species group (= former
subgenus Dodecamyrmica). However, M. tschekanovskii males have a relatively long scape
(SI, > 0.70), while males of M. arnoldii have short scape (SI, < 0.40) and under our more
recent approach to the species group concept in genus Myrmica, species with such large
differences in the male morphology probably do not belong to the same group. Thus the
taxononlic position of M. tschekanovskii is somewhat uncertain, perhaps, it should typify
its own separate species group.
Queens and ecology are unknown.

Map 126. Distribution of M.. tschekanovskii.

Myrmica tulinae Elmes, Radchenko et Aktac, 2002

(Figs 295-297)

Myrmica tulinae Elmes, Radchenko et Aktac, 2002: 158, w, q, m, Turkey; Radchenko et al. 2003: 54,
2004: 50; Radchenko and Elmes 2004: 232.

Type localities and type specimens. Turkey, for details see material examined.
Material examined. Holotype, w, "Turkey, Thrace, Kirklareli District, near Bulgarian border, alt.
.380 m, 24.viii.1999, TR-59, leg. J. E. and G. W. Elmes" (EDIRNE); paratypes: 49 w, 3 q, 18 m from the
nest of holotype; 10 w, same locality, date and collectors, TR-60; 24 w, 3 m, same locality and date, No
99/1721, leg. K. Kiran; 17 w, same locality and date, No. 99/1722, leg. K. Kiran; 30 w, 1 q, 1 m, same
locality and date, No. 99/1723, leg. K. Kiran; 20 w, 3 m, "Turkey, Thrace, 53 km east Kirklareli, Vize
- Sergen Köyü, Об.ix.1988, leg. N. Aktac"; 9 w, "Turkey, Thrace, Kirklareli, Dercköy, 18.iv.1989,
No. 89/16, leg. N. Aktac; 1 w, 1 q, "Turkey, Thrace, Kirklarcli, Pehlivanköy-Ku?tepe köyü, Ol.x.1996,

311
No. 96/124, leg. N. Aktac; 60 w, 15 q, 2 m, "NE Turkey, south edge of Artvin Region, high pass ca. 10
km N of Ardahan, alt. 2150 m, 15.viii.2000, TR-123, leg. G. W. Elmes" (EDIRNE, KIEV, WARSAW,
ELMES); non-type material: > 150 w, about 20 q and 20 m from France, Italy, Netherlands, Poland,
and Georgia.
Distribution (Map 127). Turkey, Georgia, France, Italy, Netherlands and Poland. Be-
cause based only on the female castes M. tulinae can be confused with M. sabuleti, this
species in fact may be widespread in Euro-Caucasian region.
Etymology. M. tulinae: this species was dedicated to Prof. Tulin Aktac of Trakya Uni-
versity, Turkey, the wife of our friend and colleague Prof. Nihat Aktac of Trakya University.
Notes. M. tulinae belongs to the scabrinodis-comp\ex of the scabrinodis species group.
The female castes of M. tulinae most resemble those of M. sabuleti and are barely distin-
guishable from them. In contrast, males of M. tulinae well differ from those of M. sabuleti
by their much shorter scape (SI, < 0.40 vs. > 0.50), but by the length of the scape and by
the character of pilosity on the scape and tibiae they are very similar to M. scabrinodis males
and could easily be confused with them if found without female castes. Thus, one needs
both workers and males to be sure of correct identification of M. tulinae (see also Elmes et
al. 2002).
Ecology. M. tulinae lives in a quite wide range of altitudes, from about sea level up to
1800 m a.s.l., were it inhabits moderately humid sites or even wet and boggy places, but
never lives in dry habitats (see also Elmes et al. 2002; Radchenko et al. 2003).

Map 127. Distribution of M. tulinae.

Myrmica turcica Santschi, 1931

(Figs 298-300)

Myrmica scabrinodis var. turcica Santschi, 193lb: 343, w, q, Turkey; Weber 1950: 210.
Myrmica turcica: Seifert 1988: 18 (part., nec m and material from Kazakhstan); Radchenko 1994b: 42,
1994d: 79; Bolton 1995: 284; Radchenko and Elmes 2004: 230.

312
Senior synonym of Myrmica georgica: Seifert 1988: 18. Revived from synonymy: Radchenko and Elmes
2004: 230 (see also Notes to M. georgica, above).

Type localities and type specimens. "Ankora (Gadeau de Kcrville), vv, q, types, Moldavie; Val du
Berlad (Montandon), w, q" (see also Notes below).
Material examined. Syntypes, 1 w, 1 q on the same pin, "Type", "Angora, Asie Min. (Kerville)",
"223", "M. scabrinodis v. turcica Sant q vv type", "Sammlung Dr. F. Santschi Kairouan"; 2 w on the
same pin, "Type", "Angora (Kerville)", "17", "M. scabrinodis v. turcica Sant q w type", "Sammlung
Dr. F. Santschi Kairouan"; 2 w on the same pin with the same labels, but without number (BASLE);
non-type material: > 300 w, > 50 q, > 100 m from Turkey.
Distribution (Map 128). Asia Minor.
Etymology. M. turcica: name derived from the medieval Latin name Turcicum — Turkey,
to indicate its place of origin.
Notes. M. turcica belongs to the specioides-comp\ex of the scabrinodis species group. Its
workers resemble those of M. specioides by a similarly shaped petiolar node, relatively short
propodeal spines (mean ESLI 0.36 in both species), but differ by the more extended frontal
lobes (mean FLI 1.43 vs. 1.32). Despite the fact that Santschi (1931b: 343) wrote "Le coude
du scape fait un angle droit devant et un peu plus distant de la base que chez le type [e.g.
M. scabrinodis], le bord supérieur presque aussi lobule que chez sabuleti", in the syntype
workers and numerous non-type specimens of M. turcica that we examined, the shape and
size of the scape lobe is much more similar to that of M. scabrinodis rather than M. sabu-
leti. Additionally, workers of M. turcica are very similar to M. scabrinodis by the shape of
the frontal carinae and lobes (mean FI 0.36 in both species), but well differ from the latter
by their much shorter propodeal spines (mean ESLI 0.36 vs. 1.41). Combination of these
features well separate M. turcica from M. kozakorum (see Notes to that species). While
the males of M. turcica and M specioides are very similar and hard to separate, they are
easily distinguished from M. scabrinodis by much shorter, more decumbent hairs on the
tibiae and tarsi. If there were a greater geographical overlap of M. turcica with M. specioides

Map 128. Distribution of M. turcica.

313
and M. scabrinodis the taxonomy and discrimination of all three species would be very
difficult.
When Santschi described var. turcica, he referred material from Moldova to this form,
but did not include those workers and queens to the type series (see above). We examined
one worker from BASLE, labelled as "Moldavie Val du Berlad Montandon", "M. scabrino-
dis v. turcica Sant.", "Sammlung Dr. F. Santschi Kairouan" and determined it as M. scabri-
nodis.
Since males were not described, we describe them for the first time (based on about 100
males from various regions of Turkey).
First description of males (Fig. 300).
Head distinctly longer than broad, with convex sides and occipital margin, and rounded
occipital corners; anterior clypeal margin widely rounded, not-notched medially. Mandibles
with 6 teeth. Scape short (Slj < 0.45, SI2 < 0.50), very smoothly curved at the base, anten-
nae 13-segmented; second funicular segment only slightly longer than the third one. Alitrunk
relatively long, scutum slightly convex, scutellum does not project dorsally above scutum
when seen in profile. Propodeum with relatively long triangular denticles. Petiole longer than
height, with almost straight anterior face and narrowly rounded dorsum; postpetiole sub-
globular.
Head dorsum (excluding clypeus) densely punctated. Sculpture of alitrunk quite vari-
able: scutum and scutellum from almost completely smooth to finely longitudinally striated;
sides of pronotum and mesopleura from smooth to striated; sides of propodeum finely rugu-
losa Petiolar node longitudinally ruguloso-striated, postpetiole smooth and shiny.
Head margins and mandibles with numerous suberect hairs. Alitrunk, waist and gaster
with numerous fine hairs. Scape, tibiae and tarsi with not long hairs: those on the external
tibial margin longer than the half of the maximal width of tibia, those on the external margin
of basitarsus longer that its width. Body colour brownish.
Ecology. In Turkey M. turcica generally lives in grassland at altitudes 1000-2000 m a.s.l.
but also can be found in anthropogenic environments such as gardens, orchards and poplar
plantations. It prefers slightly damper places near to streams (even if dry in summer), where
it nests in the soil and under stones. M. turcica has an ecology that is superficially similar to
that of both M. scabrinodis and M. specioides. The most unusual feature of its behaviour was
that we observed workers foraging at midday in very hot conditions, however they were
foraging in the shade of trees and shrubs rather than in direct sunlight. We have not seen any
other Myrmica species in "Greater Europe" foraging at this time of day in such hot conditions.

Myrmica urbanii Radchenko et Elmes, 1998

(Fig. 301).

Myrmica urbanii Radchenko et Elmes, 1998: 16, w. India; 2001b: 242.

Type localities and type specimens. India (Megalaya), for details see material examined.
Material examined. Holotype, w, "India, Shillong, Mawphlang, East Khasi, 3200 ft., 2.05.86, leg.
V. Darlong" (LONDON); paratypes: 86 w from the same nest as holotype; 2 w, "Mawphlang, Mega-
laya, 1850 m, 15.05.1976, leg. Wittmer, Baroni Urbani"; 6 w, "Upper Shillong, Megalaya, 1900 m,
13.05.1976, leg. Wittmer, Baroni Urbani"; 3 w, "Cherranukh, India, 1961, leg. Korovin" (LONDON,
ELMES, BASLE, KIEV, MOSCOW).

314
 Distribution (Map 129). It is known only from the type localities in NE India (Mega-
laya).
Etymology. M. urbanii: this species was dedicated to our friend, the distinguished
Italian myrmecologist Prof. Cesare Baroni-Urbani.
Notes. M. urbanii belongs to the ritae-complex of the ritae species group. It is charac-
terised by the not-very-coarse sinuous rugosity of the frons (having not less than 6 rugae be-
tween frontal carinae level with the eyes), by the coarsely reticulated rear third of the head
dorsum, and by the presence of abundant erect hairs on the head margins. Queens and
males are unknown.
Ecologyi s poorly known; it lives at altitudes between 1000 and 2000 m a.s.l., one nest
was found in soil beneath a log in oak forest.

Map 129. Distribution of M. urbanii.

Myrmica vandeli Bondroit, 1920

(Figs 302-304)

Myrmica vandeli Bondroit, 1920a: 148, q, m, France (also described as new by Bondroit, 1920b: 301);
Finzi 1926: 115; Santschi 1931b: 347; Stitz 1939: 97; Bernard 1967: 122; Kutter 1977: 71, w;
Elmes and Thomas 1985: 97; Espadaler 1986: 49; Agosti and Collingwood 1987a: 53, 1987b: 268;
Seifert 1988: 33; 1994: 12; Radchenko 1994b: 42; Bolton 1995: 284; Seifert 1996: 226; Schlick-
Steiner and Steiner 2000: 113; Elmes at al. 2003: 145; Radchenko et al. 2003: 50; Radchenko and
Elmes 2003a: 230; Czechowska et al. 2004: 111; Radchenko et al. 2004: 48; Radchenko and Elmes
2004: 232; Stankiewicz and Antonova 2005: 123; Seifert 2007: 197; Espadaler 2008: 40; Rad-
chenko 2009: 70.
Myrmica scabrinodis var. vandeli: Weber 1950: 211.

Type localities and type specimens. "Environs de Pontarlier: Les Granges et environs, Ste-
Colombe, juillet et août" (Bondroit 1920a); "Doubs: Les Granges, près de Pontarlier, vers 820 mètres;
vol nuptial vers la fin d'août (A. Vandel)" (Bondroit 1920b).

315
 Material examined. Lectotype (designated by Radchenko and Elmes 2003a), q (upper specimen
on the pin with 2 q and 2 m, in copula with m), "Les Ganges, Jura", "Coll. R. I. Sc. N. B. France",
"R. U. Sc. N. B. I. G. 21.400", "Type", "Myrmica vandeli Bondr. Type" (BRUSSELS); paralectotypes:
1 q, 2 m on the pin with lectotype; 2 q, б m with the same labels as lectotype; 1 q, 2 m, "Les Ganges
près Pontarlier", "Coll. R. I. Sc. N. B. France", "R. U. Sc. N. B. I. G. 21.400", "Type", "Myrmica van-
deli Bondr. Type"; 1 m (without waist and gaster), "Pontarlier, A. Vandel", "Les Ganges, fin A ut
1918", "Coll. R. I. Sc. N. B. France", "R. U. Sc. N. B. I. G. 21.400", "Type", "Myrmica vandeli Bondr.
Type"; 2 m, "St. Colombe, Doubs, 2.ix.1919", "Coll. R. I. Sc. N. B. France", "R. U. Sc. N. B. I. G.
21.400", "Myrmica vandeli Bondr. Type" ; 1 m, "Gangcs-Marboz, Doubs, 3.ix.1919", "Coll. R. I. Sc.
N. B. France", "R. U. Sc. N. B. I. G. 21.400", "Myrmica vandeli Bondr. Type" (BRUSSELS); non-
type material: about 100 w, > 40 q and m from Great Britain, Switzerland, France, Poland, Czech
Republic, Bulgaria, Romania and Ukraine.
Distribution (Map 130). France, Spain, Great Britain (England and Wales), Sweden,
Germany, Switzerland, Austria, Poland, Czech Republic, Slovakia, Rumania, former
Yugoslavia, Bulgaria and Ukraine; record of Agosti and Collingwood (1987a) for Turkey
(Trace) needs confirmation.
Etymology. M. vandeli: this species was named for the French biologist Prof. Albert
Vandel (1894-1980) who collected the type specimens.
Notes. M. vandeli belongs to the sabuleti-complex of the scabrinodis species group.
It was originally described from queens and males from France and was known only from
the type series for more than 50 years, until Kutter (1977) described workers of this species
from Switzerland. Elmes and Thomas (1985) showed that it is a quite common free-living
species in France that can easily be discriminated from M. scabrinodis (all castes) by
morphometries.
M. vandeli possess several "socially parasitic" features, such as reduced spurs on the
mid- and hind tibiae and hairy body, and by this characters is similar to M. bibikoffi and
M. hirsuta. On the other hand, workers of M. vandeli appear very similar to those of
M. scabrinodis by the shape of alitrunk and waist, by the size and shape of the lobe at the
bent of the scape, by the shape of frontal carinae and frontal lobes (including FI and FLI),
but differ from the latter by more abundant standing hairs on the body (petiole with > 10,
often > 20 hairs vs. < 10, usually < 8 hairs), by the presence of at least shallow medial
notch on the anterior clypeal margin, by the less coarse, almost straight longitudinal ru-
gosity on the alitrunk dorsum (vs. strong reticulation in M. scabrinodis) and by the reduced
(to various extents) sculpture of the petiolar and postpetiolar dorsum, and finally, by its re-
duced tibial spurs.
Males of M. vandeli have relatively long scapes and could be confused with M. sabuleti,
but they well differ from it by the presence of abundant long standing hairs on the scape and
head margins, and by the much shorter hairs on the mid- and hind tibiae.
Queens of M. vandeli differ from those of all species of the scabrinodis-group by their
large size, very dark, almost black body colour, and especially by the characteristic longi-
tudinally-concentric rugosity on the petiolar node dorsum.
Ecology. M. vandeli inhabits almost exclusively rather warm but humid sites, such as
wet meadows, marshes and bogs, where it builds nests usually in moss pads, tussocks of
grass and sedge, rarely in the soil under stones. Almost always M. vandeli coexists
with M. scabrinodis, though it is generally a much less abundant and widespread species.
The "parasitic" morphological features of the queens and males led Bondroit (1920a: 148)
to suggest a socially-parasitic style of life for M. vandeli: "Il est probable que le M. Vandeli
est une espèce parasite". We have found both "pure" colonies and colonies of mixed

316
M. scabrinodis and M. vandeli (Elmes et al. 2003; Radchenko et al. 2003; Radchenko 2009)
in the centrai part of its range, where environmental conditions probably are optimal for
M. vandeli, we hardly ever located mixed colonies, but at the edges of its distribution (e.g.
in Great Britain, Poland or Ukraine) M. vandeli often coexists with M. scabrinodis. This led
us to suggest that in "poor" or new habitat queens of M. vandeli might resort to facultative
social parasitism to found colonies, but in optimum habitats it may be able to establish
colonies by independent foundation or by colony division.

Map 130. Distribution of M. vandeli.

Myrmica varisculpta Radchenko et Rigato, 2009

(Fig. 305)

Myrmica varisculpta Radchenko et Rigato, 2009: 76, w, India (in Radchenko et al. 2009a).

Type localities and type specimens. India (Kashmir), for details see material examined.
Material examined. Holotype, w, "India, Kashmir, Leh, viii.1986, leg. С Canepari" (MILAN).
Distribution (Map 131). It is known only from the type locality India (Kashmir).
Etymology. M. varisculpta: from a combination of the Latin words varius = different
and sculptus = sculptured, to describe the various combinations of sculpture of the alitrunk.
Notes. M. varisculpta was described from a unique holotype worker, it belongs to the pa­
chei species group. Recently this group was considerably enlarged with the discovery of
a number of new species from China (Radchenko and Elmes 2009a). It is the third such
species discovered on the Indian side of the Himalaya and the mixture of sculpture on its
alitrunk dorsum clearly separates it from the others, M. pachei and M. villosa, which have
only transverse striations. It is most similar to the Chinese species M. phalacra and
M. hlavaci but has a distinctly longer head than these two (for the separation from all
related species see Key, Chapter 5.6, 5.7).
Queens, males and ecology are unknown.

317
 Map 131. Distribution of M. varisculpta.

Myrmica villosa Radchenko et Elmes, 1999

(Figs 306, 307)

Myrmica villosa Radchenko et Elmes, 1999a: 31, w, q, Bhutan; 2001b: 257, 2009a: 69.

Type localities and type specimens. Bhutan, for details see material examined.
Material examined. Holotype, w, "Dechli Рака, 5 km O Pélela, 3300 m, 19-20.06.1972 (Natural-
History Museum Basel Bhutan Expedition, 1972)" (BASLE); paratypes: 15 w, 1 q with the same label
(BASLE, LONDON, KIEV, ELMES).
Distribution (Map 132). It is known only from the type locality in Bhutan.

Map 132. Distribution of M. villosa.

318
 Etymology. M. villosa: from the adjective villose derived from the Latin word villus =
shaggy hair, to describe the pilosity of the body.
Notes. M. villosa belongs to the pachei species group, and is closely related to M. pachei,
differing from the latter mainly by a denser sculpture of the head dorsum, by its more abun-
dant, long, thin, fine, curved body hairs and longer hairs on legs, and by the lighter,
brownish red colour of the alitrunk, waist and gaster. These two species differ from all other
known species of the pachei-group by having only transversal striations on the alitrunk
dorsum. Males are unknown.
Ecology is unknown except that the type series were found at the altitude 3300 m a.s.l.

Myrmica vittata Radchenko et Elmes, 1999

(Fig. 308)

Myrmica vittata Radchenko et Elmes, 1999a: 33, w, Pakistan; 2001b: 260.

Type localities and type specimens. Pakistan, for details see material examined.
Material examined. Holotype, w, "Pakistan, Bumburet, 24.05.83, leg. M. Brancucci (LONDON);
paratypes: 2 w with the same label; 2 w, "Pakistan, Kalam, 12.08.1994, leg. S. Dakatra" (LONDON,
MILAN, KIEV, ELMES).
Distribution (Map 133). It is known only from the type localities in Pakistan.
Etymology. M. vittata: from the Latin word vitta - band or ribbon with the adjectival suf-
fix - ata, to describe the band of straight longitudinal rugae on the head and alitrunk dorsum.
Notes. The taxonomic position of M. vittata is unclear; we can not place it to any species
group. It differs from all the other Eurasian Myrmica species, which have a weak curve at the
base scape, by the absence of rugae, surrounding the antennal sockets, combined with the coarse,
regular longitudinal rugosity of the head, alitrunk (including its dorsum) and petiolar nodes.
Queens, males and ecology are unknown.

Map 133. Distribution of M. vittata.

319
 Myrmica wardi Radchenko et Elmes, 1999
(Figs 309-311)

Myrmica wardi Radchenko et Elmes, 1999a: 38, w, q, m, India, Pakistan; 2001b: 252.

Type localities and type specimens. India and Pakistan, for details see material examined.
Material examined. Holotype, w, "Kashmir, Ladakh, Leh, 34°11'N, 77°35'E, 3450 m, No. 3094,
21.08.1978, leg P.Ward" (LONDON); paratypes: б w with the same label; 6w, 1 q, "Kashmir, Pahal-
gam, 34°02'N, 75°19'E, 2190 m, No. 3003, 27.07.1978"; about 100 w, 2 m, "Kashmir, Lidderwat,
34°09'N, 75°15'E, 2700 m, No. 3015, 3018c, 3023, 3029, 30.07.1978"; 1 w, 1 q, "Kashmir, Kulan,
34°16'N, 75°09'E, 2100 m, No. 3050-10, 3050-12, 08.08.1978"; 29 w, 1 q, "Kashmir, Ladakh,
Panikhar, Suru R., 34°07'N, 75°57'E, No. 3077, 3078, 17.08.1978"; 3 w, "Kashmir, Ladakh, Leh,
3411'N, 77°35'E, 3450 m, No. 3090, 21.08.1978"; 4 w, "Kashmir, Sonamarg, 34°18'N, 75°18'E, 2700
m, No. 3102" (ali leg. P. Ward); 8 w, Pakistan, "Chitral Valley, between Dir and Lawari Pass, 2400 m,
11.08.1994, leg. S. Dacatra"; 3 w, "Indien, Himahal Pradesh, vie. Theong, 25 km E Shimla, 2400 m,
No 394, 29.09.1996, leg. A. Schulz and K. Vock"; 3 w, 2 q, "India, Himahal Pradesh, Kullu valley,
5-7 km SW Rothang La Pass, 2500-2900 m, No. 432, 01.10.1996, leg. A. Schulz and K. Vock"
(LONDON, BASLE, MILAN, SCHULZ, WARD, KIEV, ELMES, MEI); non-type material: 2 w from
Pakistan (see also Notes below).
Distribution (Map 134). It is known only from the type localities in India (Kashmir and
Himahal Pradesh States) and Pakistan.
Etymology. M. wardi: this species was dedicated to our friend, the distinguished
American ant taxonomist, Prof. Philip S. Ward of the University of California, who
collected the types of this and several other species described by us.
Notes. M. wardi belongs to the cachmiriensis-complex of the rugosa species group.
It most resembles M. cachmiriensis, its workers differ by the much denser longitudinal rugu-
losity on the head dorsum with densely punctated surface between the rugae, by the darker
colour of the body, by the presence of much longer, well visible subdecumbent hairs

Map 134. Distribution of M. wardi.

320
on head sides, etc. Males differ by their distinctly higher petiole (PIj = 1.33-1.36 vs.
1.40-1.64), and by the reduced rugulosity and presence of punctures on the sides of the
alitrunk (those of M. cachmiriensis are densely but not coarsely rugulose). Based on the
identifications made by some of the earlier myrmecologists, we guess that this species often
has been confused with other Himalayan Myrmica species.
We determined as M. 'war di the "paratype" worker, collected in Pakistan and attributed
to M. aimonissabaudiae (see Notes to the that species, above).
Ecology. Philip Ward (pers, comm.) collected this species in Kashmir at altitudes of
2190-3450 m a.s.l. in the same regions, but at lower altitudes than M. rhytida (mean 2700 m
vs. 3600 m). It lives mainly in open woodland, such as meadows with sparse trees (willow,
pear, maple and fir), woody shrubs with Rosa spp., and stony grasslands and pastures
having a low sward. A few nests were found in the soil, but most specimens were collected
while foraging, which suggests that this species does not often use stones as nest sites (many
ants are collected by people turning-over stones). Foragers freely climbed over low vegeta-
tion, perhaps being indicative of foraging habits similar to the European M. rubra.

Myrmica weberi Elmes et Radchenko, 2009

(Figs 312, 313)

Myrmica weberi Elmes et Radchenko, 2009: 112, w, q, Nepal, India and Bhutan.
Myrmica indica: Radchenko and Elmes 1998: 11, w (part.), 2001b: 241 (part.).

Type localities and type specimens. Nepal, India and Bhutan, for details see material examined.
Material examined. Holotype, w, "NPL28 (No. 11 - Elmes coli, label), Nepal, Sankhawalava
Maghang, Kharka, Makalu Barun Conservation Area, 27°36'18.5"N 87°7'30"E, 2634 m, 7.XÌ.2005, leg.
Alpert, Alonso and Subedi, CK-3, yak meadow under rocks, under stones" (HARVARD); paratypes:
15 w, 1 q with the same label as holotype; 7 w, "NPL25, Nepal, Sankhawalava Maghang, Kharka,
Makalu Barun Conservation Area, 27°35'36.6"N 87°7'20.7'E, 2548 m, 5.xi.2005, leg, D. Emmett and
Subedi, MK 21-283, Winkler trap"; 3 w, "NPL 31, Nepal, Sankhawalava Makalu Barun Conservation
Arca, MK-4, 27°35'24.8"N, 87°7'18.7"E, 2563 m, 3.XÌ.2005, leg. G. Alpert, on rock under moss, large
colony with 2 q"; 3 w, "NPL 32, Nepal, Sankhawalava Makalu Barun Conservation Area, MK-6,
27°35'24.8"N 87°7'18.7"E, 2563 m, 3.xi.2005, leg. G. Alpert, open meadow, under logs"; 7 w, "NPL
33, Nepal, Sankhawalava, Makalu Barun Conservation Area, MK W17, 7.xi.05, Winkler trap"; 1 w,
"India, Darjeeling Distr., Tiger Hill, 2500 m, 27.V.1975, leg. W. Wittmcr", "M. indica Weber det. Rad-
chenko and Elmes"; 1 w, "Kosi, Chauki, 2°11-12'N 87°27-28'E 2000-3000 m, 22-24.vi.01, NHMB
Exped. Npl. 2001"; 1 w, "Bhutan, Nolding, 41 km О Wangdi, 2800 m, Nat.-Hist. Museum Basel -
Bhutan Expedition, 1972", "M. indica Weber det Radchenko and Elmes"; 1 w, "NPL2.1, Nepal, Rigmo,
H. Tabata, 7.VÌ.1978" "M. indica Weber det Radchenko and Elmes" (BASLE, KIEV, ELMES, HAR-
VARD); non-type material: 10 w, "Bhutan, Sampa-Kotoka, 1400-2600 m, 9.06, Nat.-Hist. Museum
Basel - Bhutan Expedition, 1972", "M. indica Weber det Radchenko and Elmes".
Distribution (Map 135). Nepal, India (Darjeeling Distr.) and Bhutan.
Etymology. M. weberi: this species was dedicated to the well-known American myrme-
cologist, Prof. Neil A. Weber, who wrote the first monographic review on genus Myrmica
(see biography in Chapter 1.6).
Notes. M. weberi belongs to the ritae-complex of the ritae species group. It most re-
sembles M. indica and M. alperti, but differs from them by the distinctly smaller size,
the thinner and longer propodeal spines, by the more developed reticulation on the head

321
 Map 135. Distribution of M. weberi.

dorsum, etc. Additionally, it differs from M. indica by the shape of frontal carinae that curve
outwards to merge with the rugae, which surround antennal sockets, and from M. alperti -
by the sharply pointed apically propodeal lobes.
Ecology. This species is probably restricted to the southern slopes of the Central Hi­
malaya (Nepal, India and Bhutan), where it lives between 2000 and 3000 m a.s.l. Alpert's
collection data suggests that it might be a rather reclusive species that nests under moss,
dead wood and rocks, and forages close to the ground and in litter.

Myrmica weii R a d c h e n k o et Z h o u , 2 0 0 8
(Fig. 314)

Myrmica weii Radchenko et Zhou, 2008: 779, w, China (in Radchenko et al. 2008); Radchenko and
Elmes 2009a: 72.
Myrmica inezae: Wei Z. et al. 1999: 60; Wei С et al. 2001: 561, misidentifications.

Type localities and type specimens. China, for details see material examined.
Material examined. Holotype, w, "China, Shaanxi Prov., Houzhenzi, July 25 1997, Cong Wei leg."
(GU1LIN).
Distribution (Map 136). It is known only from the type locality in the Shaanxi Prov.,
China.
Etymology. M. weii: named for the collector, the Chinese entomologist Prof. Cong Wei
of The Northwest Agriculture and Forestry University, Yangling, China.
Notes. M. weii belongs to the pachei species group. It well differs from M. pachei and
M. villosa by the much coarser transverse rugosity on the propodeal and mesonotal dorsum
and reticulated pronotal dorsum. Its head margins have numerous long suberect hairs,
and by this feature it is similar to M. pleiorhytida, M. multiplex, M. yunnanensis and

322
 Map 136. Distribution of M. weii.

M. heterorhytida. It differs from the first species by its distinctly longer head (CI > 1.20 vs.
< 1.15), it has a much lower petiole than M. multiplex (PIj 1.68 vs. < 1.55), and has coarser
transversal rugosity on the alitrunk dorsum than M. yunnanensis and M. heterorhytida
(number of the rugae on this area < 20 vs. > 25).
In some respects M. weii resembles M. inezae for which it was mistaken when first col-
lected. It well differs from that species by its reticulated pronotal dorsum whereas the whole
alitrunk dorsum of M. inezae is transversely rugose, and especially by the shape of petiole
and propodeal spines.
Queens, males and ecology unknown.

Myrmica wesmaeli Bondroit, 1918

(Figs 315-317)

Myrmica wesmaeli Bondroit, 1918:. 106, w, France; 1920a: 151; Arnoldi 1934: 158; Seifert 1988: 39,
q; Bolton 1995: 284; Elmes et al. 2002: 169; Seifert 2005: 4; Radchenko et al. 2008: 53, nec Finzi
1926: 97, w, q, m; Novak and Sadil 1941: 78; Chang and He 2001: 26, misidentifications.
Myrmica sulcinodis var. wesmaeli: Emery 1921: 42; Santschi 1931b: 340.
Myrmica scabrinodis subsp. wesmaeli: Weber 1950: 211.
Senior synonym of Myrmica sulcinodis var. sulcinodoscabrinodis Forel: Emery 1921: 42; not con-
firmed, see Notes to M. sulcinodis, above.
Junior synonym of M. lobicornis: Bernard 1967: 122; not confirmed, see Notes below.

Type localities and type specimens. "Pyrénées-Orientales: Canigou, vers 1.800 m".
Material examined. Lectotype (designated by Seifert 1988), w, "Canigou", "Coli. R. I. Sc. N. B.
France, Pyr. Orie. Mont Canigou", "Myrmica Wesmaeli Bondr. Type", "Lectotype, det. Seifert 1987"
(BRUSSELS); paralectotypes: 7 w.with the.same labels (BRUSSELS); non-type material: 45 w, 5 q,
4 m from Spain.

323
 Distribution (Map 137). Iberian Peninsula, France (Pyrenees).
Etymology. M. wesmaeli: named for the Belgian entomologist Constantin Wesmael
(1798-1872).
Notes. M. wesmaeli belongs to the lobicornis species group (this was stressed by Bon­
droit 1920a). It differs from the sympatric M. lobicornis and M. lobulicornis by its wider
frons and especially by the shape of petiole, which has developed peduncle and concave, not-
steep anterior surface that meets the dorsal one to form a rounded blunt angle (seen in pro­
file). It well differs from the sympatric M. xavieri by its notched medially anterior clypeal
margin and by the its wider frons and less extended frontal lobes.
For many years, M. wesmaeli was a somewhat enigmatic taxon, especially after Finzi's
(1926) erroneous treatment of this species. He redescribed workers and described queens
and males for the first time (the "types" of queen and male from Pyrenees) of "M. wesmaeli"
based on the material from Iberian Peninsula, Italy, Croatia ("Dalmazia"), and Czech Re­
public. He wrote about workers: "Scapo piegato ad angolo alla base, senza alcun lobo о in­
grossamento nel punto della piegatura", and about males: "Scapo ... lungo come i primi tre
articoli del funicolo" (loc. cit., p. 97-98). Based on these features, the described specimens
can not belong to M. wesmaeli, whose workers and queens have a small vertical lobe or
dent at the base of the scape, and males have the much longer antennal scape. Possibly,
Finzi's Iberian material belongs to M. aloba, but specimens from the other regions may rep­
resent different species.
Not surprisingly, some of the subsequent authors considered M. wesmaeli as infraspe-
cific form of M. sulcinodis or M. scabrinodis (Emery 1921; Santschi 1931b; Weber 1950),
but Bernard (1967) treated it as a junior synonym of M. lobicornis.
Novak and Sadil (1941) under the name "M. wesmaeli" include species from the scabri-
nodis-group, probably M. specioides, M. hellenica or even M. constricta: "Scape at the base only
angularly curved, not widened at the bent and without longitudinal lobe" (loc. cit., p. 78, trans­
lation from Czech). Records of M. wesmaeli for China (Chang and He 2001) can not belong
to this species and most probably should be referred to some other eastern lobicornis-group
species.
Seifert (1988) made the first correct treatment of M. wesmaeli since its first description,
he redescribed workers and made the first description of queens, but did not describe the
males. Since the males of "M. wesmaeli" described by Finzi (1926) belong to another species,
we describe males below (2 males from Spain, Areo, Lérida, leg. X. Espadaler, July 1975).
First description of males (Fig. 317).
Head longer than broad, with convex sides and occipital margin, and rounded occipital
corners; anterior clypeal margin widely rounded and notched medially. Scape long (SIj 0.80,
SI2 0.85), quite strongly but not angularly curved at the base, antennae 13-segmented;
second funicular segment only slightly longer than the third one. Alitrunk relatively long,
scutum slightly convex, scutellum does not project dorsally above scutum when seen in
profile. Propodeum with short blunt denticles. Petiole relatively long, with slightly concave
and not steep anterior surface and rounded node dorsum; postpetiole higher than long, with
convex dorsum.
Head dorsum finely longitudinally striato-rugulose, surface between rugulae densely
punctated, appears dull, while clypeus smooth and shiny. Scutum smooth, only with very
fine superficial striation posteriorly, scutellum with semiconcentric rugulosity. Sides of
pronotum and mesopleura smooth, only catepistern posteriorly with short rugulae, sides of
propodeum rugulose. Petiole and postpetiole smooth and shiny.

324
 Head margins and mandibles with numerous, relatively long suberect hairs.
Alitrunk, waist and gaster with numerous fine hairs. Scape, tibiae and tarsi with quite
dense subdecumbent to suberect pilosity. Body colour blackish-brown, appendages
brownish.
Ecology. There have been no detailed studies of the ecology of M. wesmaeli. It usually
is found in the south-eastern part of the Pyrenean mountains where it is most common at
altitudes of 1500-1800 m. a.s.l. in subalpine pastures. It often nests under stones. Sexuais
are present in the nests in July and August so nuptial flights probably occur from August to
September.

Map 137. Distribution of M. wesmaeli.

Myrmica williamsi Radchenko et Elmes, 1999

(Fig. 318)

Myrmica williamsi Radchenko et Elmes, 1999a: 34, w, India; 2001b: 260.

Type localities and type specimens. India, for details see material examined.
Material examined. Holotype, w, "India, Kashmir, Pantitop, 2000 m, 6.09.86, leg. P. Williams"
(LONDON).
Distribution (Map 138). It is known only from the type locality in the Kashmir State in
India.
Etymology. M. williamsi: named for the collector, Dr. Paul H. Williams of the Natural
History Museum, London.
Notes. M. williamsi was described only from a single holotype worker because this
specimen so well differs from any other Myrmica species that have antennal scapes
weakly curved at their base and frontal carinae that curve outwards to merge with the

325
rugae which surround antennal sockets. The main distinguishing features are the very
short, dentiform but sharp propodeal spines and unusual sculpture of the alitrunk: the
promesonotal dorsum is coarsely reticulate and very finely punctated, sides of the
pronotum are sinuously rugulose; while the remaining parts of the alitrunk, petiole and
postpetiole have no rugae but are densely punctated. The taxonomic position of this
species is uncertain.
Queens, males and ecology are unknown.

Map 138. Distribution of M. williamsi.

Myrmica wittmeri Radchenko et Elmes, 1999

(Fig. 319)

Myrmica wittmeri Radchenko et Elmes, 1999a: 38, w, India, Pakistan; 2001b: 255.

Type localities and type specimens. India and Pakistan, for details see material examined.
Material examined. Holotype, w, "Indien, Him. Prad. [Himahal Pradesh], Mahri, 3000-3200 m,
15.05.1977, leg. Wittmer et Brancucci" (BASLE); paratypes: 3 w (one without head) with the same
label; 3 w, "Pakistan, Kalam, 2300 m, 12.07.94, leg. S. Dakatra"; 6 w, "India, Himahal Pradesh, Kullu
valley, La Pass, 3400-3700 m, 02.10.1996, No. 420, 422, leg. A. Schulz and K. Vock" (LONDON,
MILAN, SCHULZ, KIEV, ELMES).
Distribution (Map 139). It is known only from the type localities in India (Himahal
Pradesh State) and Pakistan.
Etymology. M. wittmeri: named for the collector, the coleopterologist Dr. Walter
Wittmer (1915-1998) of the Natural History Museum, Basle, Switzerland.
Notes. M. wittmeri belongs to the smythiesii species group and distinctly differs from the
other species of this group by its much darker colour and reduced sculpture on the alitrunk.
Queens, males and ecology are unknown.

326
 Map 139. Distribution of M. wittmeri.

Myrmica xavieri R a d c h e n k o , Elmes et Savolainen, 2 0 0 8

(Figs 320-322)

Myrmica xavieri Radchenko, Elmes et Savolainen, 2008: 51, w, q, m, Spain.

Type localities and type specimens. Spain, for details see material examined.
Material examined. Holotype, w, "Spain, Aragón, Province of Teruel, Sierra de Albarracín, Fuente del
Canto, ca. 3 km WSW of Bronchales, 40°30'28"N, 1°37'08"W, 1630 m a.s.l., l.viii.2000, No. 334/2000 (se-
ries E-186 in ELMES), leg. R. Savolainen and K. Vepsäläinen" (LONDON); paratypes: 9 w, 30 q, 35 m
from the nest of holotype; 10 w, "Spain, Castilla y Léon, Prov. Soria, Sierra de Urbion, ca. 25 km N Soria,
42°N, 2°30'W, 1500 m a.s.l., 7.VÜ.1990, E-117, leg. G. W. Elmes (KIEV, ELMES, ESPADALER).
Distribution (Map 140). It is known only from the type localities in Spain.
Etymology. M. xavieri: this species was dedicated to our friend and colleague, the
Spanish myrmecologist Prof. Xavier Espadaler of Barcelona.
Notes. We tentatively placed M. xavieri in the lobicornis-group based on the shape of
scape of the female castes (it has a small vertical lobe at the bend) and on the long scape of
the males. On the other hand, the shape of scape of the workers and queens is similar to that
of the species of schencki-group (e.g. shield-like process presents on its short, vertical part).
Moreover, all castes have a not-notched medially anterior clypeal margin (in contrast, the
species of the lobicornis- and schencki-groups have distinctly notched clypeus). Finally, the
scape of the male is even longer than that of other lobicornis-group species, in length it is
similar to the raòra-group species but it is distinctly angled at its base. We suspect that
M. xavieri is an Iberian endemic being a relict of a fauna isolated in Spain during recent ice
ages and may typify its own separate species group. In the recent molecular phylogeny of
Jansen et al. (2010) M. xavieri is most related to M. sulcinodis (see Fig. 332 in Chapter 4).
Ecology. Nothing is known of its ecology other than the two nests were fairly typical for
lobicornis-group species, being in soil under stones. The nest of the holotype was found at
1630 m in open pine forest (Pinus sylvestris) with moist soil and the other nest was in well-
trampled grass on a mountain pass c.1500 m near pine forest.

327
 Map 140. Distribution of M. xavieri.

Myrmica yamanei Radchenko et Elmes, 2001

(Fig. 323)

Myrmica yamanei Radchenko et Elmes, 2001a: 223, w, Vietnam; Radchenko, Elmes and Viet 2006: 41.

Type localities and type specimens. Northern Vietnam, for details see material examined.
Material examined. Holotype, w, "Northern Vietnam, Lao Cai, Sa Pa, Xeo Mi Ti, alt. 1680 m,
29-30.vii.1998, leg. B. T. Viet" (YAMANE); non-type material: 1 w, Northern Vietnam, Lai Chau
Prov., western slopes of Mt. Fan Si Pan (W. Cong Troi), 2100-2200 m, 6.V.2002, leg. K. Eguchi; 15 w,
Vientam. Lao Cai, Sa Pa, Cong Troi, 1800-2000 m, 8-14.X.2006, leg. K. Eguchi.
Distribution (Map 141). Northern Vietnam.

Map 141. Distribution of M. yamanei.

328
 Etymology. M. yamanei: this species was dedicated to our friend and colleague, the
Japanese myrmecologist Prof. Seiki Yamane from Kagoshima University.
Notes. We have recently placed M. yamanei in the newly established draco-complex of
the ritae species group (Radchenko, Elmes and Viet 2006) ; the body sculpture of the species
belonging to this complex is intermediate between that of the ritae- and boltoni-complexes.
M. yamanei well differs from M. draco and M. schoedli by the much coarser sculpture on
the head dorsum with the well developed reticulation on its rear part.
Queens and males are unknown.
Ecology. Nothing is known about this species except that it lives in forests at altitudes
between 1680 and 2200 m.

Myrmica yunnanensis Radchenko et Elmes, 2009

(Fig. 324)

Myrmica yunnanensis Radchenko et Elmes, 2009a: 82, w, China.

Type localities and type specimens. China, for details see material examined.
Material examined. Holotype, w, "China, N Yunnan Prov., Zhongdian, 3300 m, 6-8.vii.2002, leg.
S. Murzin and I. Shokhin" (PETERSBURG); paratype: 1 w with the same label as holotype (KIEV).
Distribution (Map 142). It is known only from the type locality in the Yunnan Prov, of China.
Etymology. M. yunnanensis: a combination of the name Yunnan and the Latin suffix
ensis = place of origin, to indicate that it was found in Yunnan Province of China.
Notes. M. yunnanensis belongs to the pachei species group. It is characterized by the fol-
lowing Combination of the main diagnostic features, which distinguish it from the other
species of this group: pronotum with dense though not coarse reticulation; mesonotal and
propodeal dorsum with quite dense though not coarse, transversal sinuous rugae numbering
> 25; the head margins with numerous long erect to suberect hairs; the head distinctly

Map 142. Distribution of M. yunnanensis.

329
elongate (CI > 1.20), suboval, with barely marked occipital corners; the frontal carinae do not
curve outwards and do not merge with the rugae that surround antennal sockets; surface of
head dorsum between rugae appears shiny, only with very fine superficial micropunctures.
Queens and males are unknown.
Ecology is unknown except that the two workers were found on mountain meadow at
the altitude 3300 m a.s.l.

Myrmica zojae Radchenko, 1994

(Figs 325-327)

Myrmica zojae Radchenko, 1994a: 210, w, q, m, Russia; 1994b: 43, 1994c: 141, 1994Í: 90, 1994g: 107;
Bolton 1995: 284.

Type localities and type specimens. Russia, for details see material examined.
Material examined. Holotype, w, "Ю-В Алтай, Курайская котловина, № 572, 9.VI.1964, 3. Жигульская"
["SE Altai, Kuraiskayakotlovina, No. 572, 9.VÌ.1964, leg. Z. Zhigulskaya] (MOSCOW); paratypes: 19 w,
1 q, 1 m with the same label, Nos. 546, 572, 575, 617, 638, 657, vi-vii. 1964, leg. Z. Zhigulskaya
(MOSCOW, KIEV); non-type material: 8 w from Altai Mts. and 7 w from Southern Ural (Uzun-Kul).
Distribution (Map 143). Russia (SE Altai and S Ural).
Etymology. M. zojae: this species was dedicated to the collector, our friend and col-
league, the Russian myrmecologist Dr. Zoja Alexandrovna Zhigulskaya from Magadan.
Notes. M. zojae belongs to the lobicornis species group and differs from M. lobicornis
by its wider frons, by the shape of petiole, by the shorter propodeal spines, by the shape of
the lobe at the base of scape, which is directed upward and somewhat forward, etc. From
other sympatric species from the lobicornis-group it also well differs by the shape and
bigger size of the scape lobe.
Ecology. M. zojae is a rather xerophylous species, inhabiting steppes where it builds
nests in soil.

Map 143. Distribution of M. zojae.

330
3.3.2. Drawings of the details of structure of extant Myrmica species

Figure 5. Details of structure of M. ademonia (worker, paratype). A, head, frontal view; B, scape, lateral
view; C, base of scape, anterior view ("foot of scape"); D, alitrunk and waist, lateral view; E, alitrunk
and waist, dorsal view; F, hind tibia. Scale bars = 1 mm.

331
Figure 6. Details of structure of M. ademonia (queen, paratype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

332
•Figure 7. Details of structure of M. ademonia (male, paratype). A, head, frontal view; B, scape and
three basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view;
E, hind tibia. Scale bar = 1 mm.

333
Figure 8. Details of structure ofM.afghanica (worker, holotype). A, head, frontal view; B, scape, lateral
view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view;
F, hind tibia and basitarsus. Scale bars = 1 mm.

334
Figure 9. Details of structure of M afghanica (queen, paratype). A, head, frontal view; B, scape, lateral
view; С, scape, dorsal view; D, alitrunk and petiole, lateral view; E, alitrunk and petiole, dorsal view;
F, hind tibia and basitarsus. Scale bars = 1 nini. ,

335
Figure 10. Details of structure of M. aimonissabaudiae (worker, lectotype). A, head, frontal view;
B, scape, lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind
tibia. Scale bar = 1 mm.

336
Figure 11. Details of structure of M. aimonissabaudiae (queen, paralectotype). A,.head, frontal view;
B, scape/lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind
tibia. Scale bar = 1 mm.

337
Figure 12. Details of structure of M. aimonissabaudiae (male). A, head, frontal view; B, scape and
three basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view;
E, hind tibia. Scale bar = 1 mm.

338
Figure 13. Details of structure of M. aloba (worker, lectotype). A, head, frontal view; B, scape, lateral
view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view;
F, apex of hind tibia. Scale bars = 1 mm.

339
Figure 14. Details of structure of M. aloba (queen, paralectotype). A, head, frontal, view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

340
Figure 15. Details of structure of M. aloba (male). A, head, frontal view; B, scape and three basal
funicular segments; C, alitrunk and waist, lateral view; D,alitrunk and waist,.dorsal view; E, hind tibia.
Scale bar = l.mm.

341
Figure 16. Details.of structure of M. alperti (worker, holotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

342
Figure 17. Details of structure of M. anatolica (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal
view; F, hind tibia. Scale bar = 1 mm. I

343
Figure 18. Details of structure of M. anatolica (queen, paratype). A, head, frontal view; B, scape, lateral
view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view;
F, hind tibia. Scale bar = 1 mm.

344
Figure 19. Details of structure of M. anatolica (male, paratype). A, head, frontal view; B, antenna;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

345 t
Figure 20. Details of structure of M. angulata (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

346
Figure 21. Details of structure of M. angulata (queen, paratype). A, head, frontal view; В, scape, lateral
view; С, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

347
Figure 22. Details of structure of M. angulinodis (worker, neotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
:
Scale bar = 1 mm.

348
Figure 23. Details of structure of M. angulinodis (queen). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm. •

349
Figure 24. Details of structure of M. angulinodis (male). A, head, frontal view; B, scape and three basal
funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

350
Figure 25. Details of structure of M. arisana (worker, lectotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm. .

351
Figure 26. Details of structure of M. arisana (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral .view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

352
Figure 27. Details of structure of M. arisana (male). A, head, frontal view; B, scape and three basal
funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

353
Figure 28. Details of structure of M. arnoldii (worker, paratype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, apex of hind tibia;
F, apex of middle tibia. Scale bars = 1 mm.

354
Figure 29. Details of structure of M. arnoldii (queen). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
. bars = 1 mm.

355
Figure 30. Details of structure of M. arnoldii (male). A, head, frontal view; B, antenna; C, alitrunk
and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

356
Figure 31. Details of structure of M. bactriana (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

357
Figure 32. Details of structure of M. bactriana (male, paralectotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

358
Figure 33. Details of structure of M. bakurianica (worker, neotype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal
view; F, hind tibia. Scale bars = 1 mm.

359
Figure 34. Details of structure of M.. bakurianica (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

36О
Figure 35. Details of structure of M. bakurianica (male). A, head, frontal view; B, scape and four basal
funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm. t

361
Figure 36. Details of structure of M. bergi (worker, lectotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bars = 1 mm.

362
Figure 37. Details of structure of M. bergi (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

363
Figure 38. Details of structure of M. bergi (male). A, head, frontal view; B, antenna; C, alitrunk
and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

364
Figure 39. Details of structure of M. bibikoffi (worker, paratype). A, head, frontal view; B, scape, lateral
view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view;
F, apex of middle tibia; G, apex of hind tibia. Scale bars = 1 mm.

365
Figure 40. Details of structure of M. bibikoffi (queen, paratype). A, head, frontal view; B, scape, lateral
view; С, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view;
F, apex of middle tibia; G, apex of hind tibia. Scale bars = 1 mm.

366
Figure 41. Details of structure of M. bibikoffi (male, paratype). A, head, frontal view; B, antenna;
С, alitrunk and waist, lateral view; D, alitrunk and waist, E, hind tibia; F, apex of middle tibia; G, apex
of hind tibia. Scale bars = 1 mm.

367
Figure 42. Details of structure of M. boltoni (worker, holotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

368
Figure 43. Details of structure of M. brancuccii (worker, holotype). A, head; frontal view; B, scape,
lateral view; C, alitrunk arid waist, lateral view; D, alitrunk waist and base of first gastral tergite, dorsal
view; E, hind tibia. Scale bar = 1 mm.

369
Figure 44. Details of structure of M. cachmiriensis (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

370
Figure 45. Details of structure of M. cachmiriensis (queen). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

371
Figure 46. Details of structure of M. cachmiriensis (male). A, head, frontal view; B, scape and five
basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind
tibia. Scale bar = 1 mm.

372
Figure 47. Details of structure of M.. cagnianti (worker, paratype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bars = 1 mm.

373
Figure 48. Details of structure of M. cagnianti (queen, paratype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

374
Figure 49. Details of structure of M. caucasicola (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, base of scape, anterior view ("foot of scape"); E, alitrunk and
waist, lateral view; F, alitrunk and waist, dorsal view; G, hind tibia. Scale bars = 1 mm.

375
Figure 50. Details of structure of M. caucasicola (queen). A, head, frontal view; B, scape, lateral view;
C, scape, dorsal view; D, base of scape, anterior view ("foot of scape"); E, alitrunk and waist, lateral
view; F, alitrunk and waist, dorsal view; G, hind tibia. Scale bars = 1 mm.

376
Figure 51. Details of structure of M. caucasicola (male, paralectotype). A, head, frontal view; B, scape
and three^basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal
view; E, hind tibia. Scale bars = 1 mm.

377
Figure 52. Details of structure of M. collingwoodi (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

37S
Figure 53. Details of structure of M. collingwoodi (queen, paratype). A, head, frontal view; B, alitrunk
and waist, lateral view; C, propodeal spines and waist, dorsal view. Scale bar = 1 mm.

379
Figure 54. Details of structure of M. commarginata (worker, neotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

38О
Figure 55. Details of structure of M. constricta (worker, syntype). A, head, frontal view; B, scape, lateral
view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view;
F, hind tibia. Scale bars = 1 mm.

381
Figure 56. Details of structure of M. constricta (queen, syntype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

382
Figure 57. Details of structure of M. constricta (male, syntype). A, head, frontal view; B, scape and
three basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view;
E, hind tibia. Scale bar = 1 mm.

383
Figure 58. Details of structure of M. curiosa (worker, holotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

384
Figure 59. Details of structure of M. curiosa (queen, paratype) .A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E; hind tibia. Scale
bar = 1 mm.

385
 Л. F, С, H
В, С, I), К

Figure 60. Details of structure of M. deplanata (worker, neotype). A, head, dorsal view; B, scape,
posterolateral view; C, scape, dorsal view; D, base of scape, anterolateral view; E, base of scape, anterior
view ("foot of scape"); F, alitrunk and waist, lateral view; G, alitrunk and waist, dorsal view; H, hind
tibia. Scale bars = 1 mm.

386
Figure 61. Details of structure of M. deplanata (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

387
 А, С, D, E

Figure 62. Details of structure of M. deplanata (males). A, head, frontal view; B, scape and three basal
funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind
tibia. Scale bars = 1 mm.

388
Figure 63. Details of structure of M. displicentia (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bars = 1 mm.

389
Figure 64. Details of structure of M. displicentia (queen, paratype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

390
Figure 65. Details of structure of M. displicentia (male, paratype). A, head, frontal view; B, scape and
three basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view;
E, hind tibia. Scale bar = 1 mm.

391
Figure 66. Details of structure of M. divergens (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bars = 1 mm.

392
Figure 67. Details of structure of M. divergens (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

393
Figure 68. Details of structure of M. divergens (male). A, head, frontal view; B, antenna; C, head
(sculpture omitted), alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

394
Figure 69. Details of structure of M. draco (worker, holotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

395
Figure 70. Details of structure of M. draco (queen, paratype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

396
Figure 71. Details of structure of M. draco (male, paratype). A, head, frontal view; B, antenna;
C, head, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

397
Figure 72. Details of structure ofM.dshungarica (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

398
Figure 73. Details of structure of M. dshungarica (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

399
Figure 74. Details of structure of M. dshungarica (male). A, head, frontal view; B, scape and three
basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view;
E, hind tibia. Scale bar = 1 mm.

400
Figure 75. Details of structure of M..eidmanni (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal
view; F, hind tibia. Scalebars = 1 mm.

401
Figure 76. Details of structure of M. eidmanni (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

402
Figure 77. Details of structure of M. eidmanni (male). A, head, frontal view; B, scape and three basal
funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

403
Figure 78. Details of structureofM . emeryi (worker, holotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

404
Figure 79. Details of structure of M. ereptrix (queen, holotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, apex of hind tibia;
F, apex of middle tibia. Scale bars = 1 mm.

405
Figure 80. Details of structure of M. excelsa (worker, paratype). A, head, frontal view; B, scape, lateral
view; C, base of scape, anterior view ("foot of scape"); D, alitrunk and waist, lateral view; E, alitrunk
and waist, dorsal view; F, hind tibia. Scale bars = 1 mm.

406
Figure 81. Details of structure of M. excelsa (queen, paratype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and: waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

407
Figure 82. Details of structure of M. excelsa (male, paratype). A, head, frontal view; B, scape and three
basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view;
E, hind tibia. Scale bar = 1 mm.

408
Figure 83. Details of structure of M. ferganensis (worker, syntype): A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

409
Figure 84. Details of structure of M. forcipata (worker, syntype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bars = 1 mm.

410
Figure 85. Details of structure of M. forcipata (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

4U
Figure 86. Details of structure of M. forcipata (male). A, head, frontal view; B, scape and three basal
funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

412
Figure 87. Details of structure of M. foreliana (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

413
Figure 88. Details of structure of M. fortior (worker, lectotypc). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

414
Figure 89. Details of structure of M. fortior (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

415
Figure 90. Details of structure of M. fortior (male). A, head, frontal view; B, antenna; C, alitrunk and
waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bars = 1 mm.

416
Figure 91. Details of structure of M. gallienii (worker, lectotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bars = 1 mm.

417
Figure 92. Details of structure of M. gallienii (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

418
Figure 93. Details of structure of M. gallienii (male). A, head, frontal view; B, scape and three basal
funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

419
Figure 94. Details of structure of M. georgica (worker, paratype). A, head, frontal view; B, scape, lateral
view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view;
F, hind tibia. Scale bars = 1 mm.

420
Figure 95. Details of structure of M. gigantea (worker, holotype). A, head, frontal view; B, scape, lateral
view; С, propodeum and waist, lateral view; D, apex of hind tibia; E, spur of middle tibia. Scale
bars = 1 mm.

421
Figure 96. Details of structure of M. hecate (worker, lectotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

422
Figure 97. Details of structure of M. hecate (queen). A, head,, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

423
Figure 98. Details of structure of M. hecate (male). A, head, frontal view; B, scape and three basal
funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

424
Figure 99. Details of structure of M. hellenica (worker, syntype). A, head, frontal view; B, scape, lateral
view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view;
F, hind tibia. Scale bars = 1 mm.

425
Figure 100. Details of structure of M. hellenica (queen) (paralectotype of M rugulososcabrinodis).
A, head, frontal view; B, scape, lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist,
dorsal view; E, hind tibia. Scale bar = 1 mm.

426
Figure 101. Details of structure of M. hellenica (male) (paralectotype of M. rugulososcabrinodis).
A, head, frontal view; B, scape and four basal funicular segments; C, alitrunk and waist, lateral view;
D, alitrunk and waist, dorsal.view; E, hind tibia. Scale bar = 1 mm.

427
Figure 102. Details of structure of M. heterorhytida (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

428
Figure 103. Details of structure of M. hirsuta (worker). A, head, frontal view; B, scape, lateral view;
C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view; F, distal
part of hind tibia; G, distal part of middle tibia. Scale bars = 1 mm.

429
Figure 104. Details of structure of M. hirsuta (queen, paratype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

430
Figure 105. Details of structure of M. hirsuta (male, paratype). A, head, frontal view; B, scape and four
basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view;
E, hind tibia. Scale bar = 1 mm.

431
Figure 106. Details of structure of M. hlavaci (worker, holotype). A, head, frontal view; B, scape, lateral
view; G, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

432
Figure 107. Details of structure of M. indica (worker, lectotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

433
Figure 108. Details of structure of M. indica (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

434
Figure 109. Details of structure of M. indica (male). A,, head, frontal view; B, antenna; C, alitrunk and
waist, lateral view; D, hind tibia. Scale bar = 1 mm.

435
Figure 110. Details of structure of M. inezae (worker, holotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.Scale
bar = 1 mm.

436
Figure 111. Details of structure of M. inezae (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

437
Figure 112. Details of structure of M. inucta (worker, holotype). A, head, dorsal view; B, scape,
posterolateral view; C, scape, dorsal view; D, base of scape, anterolateral view; E, base of scape, anterior
view ("foot of scape"); F, alitrunk and waist, lateral view; G, alitrunk and waist, dorsal view;
H, hind tibia. Scale bars = 1 mm.

438
Figure 113. Details of structure of M. jennyae (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

439
Figure 114. Details of structure of M. jennyae (queen, paratype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

440
Figure 115. Details of structure of M. jennyae (male, paratype). A, head, frontal view; B, antenna;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

441
Figure 116. Details of structure of M. jessensis (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bars = 1 mm.

442
Figure 117. Details of structure of M. jessensis (queen). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

443
Figure 118. Details of structure of M. jessensis (male). A, head, frontal view; B, scape and three basal
funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

444
Figure 119. Details of structure of M. juglandeti (worker, paratype). A, head, frontal view; B, scape,
lateral view; C; alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

445
Figure 120. Details of structure of M. juglandeti (male, paratype). A, head, frontal view; B, scape and
three basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view;
E, hind tibia. Scale bar = 1 mm.

446
Figure 121. Details of structure of M. kabylica (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, apex of hind tibia; F, apex of
middle tibia. Scale bars = 1 mm.

447
Figure 122. Details of structure of M. kabylica (male). A, head, frontal view; В, antenna; С, alitrunk
and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia; F, distal part of hind tibia;
G, distal part of middle tibia. Scale bars = 1 mm.

448
Figure 123. Details of structure of M. kamtschatica (worker, paratype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bars = 1 mm.

449
Figure 124. Details of structure of M. kamtschatica (queen, paratype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

450
Figure 125. Details of structure of M. kamtschatica (male, paratype). A, Head, frontal view; B, scape
and three basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal
view; E, hind tibia. Scale bar = 1 mm.

451
 А, С, D
В, E, F

Figure 126. Details of structure of M. karavajevi (queen, syntype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, apex of hind
tibia; F, apex of middle tibia. Scale bars = 1 mm.

452
Figure 127. Details of structure of M. karavajevi (male, syntype). A, head, frontal view; B, antenna;
С, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia; F, distal part of hind
tibia; G, distal part of middle tibia. Scale bars = 1 mm.

453
Figure 128. Details of structure of M. kasczenkoi (worker, neotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bars = 1 mm.

454
Figure 129. Details of structure of M. kasczenkoi (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, hind tibia. Scale bar = 1 mm.

455
Figure 130. Details of structure of M. kasczenkoi (male). A, head, frontal view; B, scape and three
basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view;
E, hind tibia. Scale bar = 1 mm.

456
Figure 131. Details of structure of M. kirghisorum (worker, paratype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunkand waist, dorsal
view; F, hind tibia. Scale bars = 1 mm.

457
Figure 132. Details of structure of M. kirghisorum (queen, paratype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

458
Figure 133. Details of structure of M. kirghisorum (male, paratype). A, head, frontal view; B, scape and
four basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view;
E, hind tibia. Scale bar = 1 mm.

459
Figure 134. Details of structure of M. koreana (worker, holotype). A, head, dorsal view; B, scape,
posterolateral view; C, scape, dorsal view; D, base of scape, anterolateral view; E, base of scape, anterior
view ("foot of scape"); F, alitrunk and waist, lateral view; G, alitrunk and waist, dorsal view; H, hind
tibia. Scale bars = 1 mm.

46O
Figure 135. Details of structure of M. koreana (queen). A, head, frontal view; B, scape, lateral view;
C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view; F, hind
tibia. Scale bars = 1 mm.

461
Figure 136. Details of structure of M. koreana (male). A, head, frontal view; В, antenna; С, scape and
three basal funicular segment; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view;
E, hind tibia and basitarsus. Scale bars = 1 mm.

462
Figure 137. Details of structure of M. kotokui (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

463
Figure 138. Details of structure of M. kotokui (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

464
Figure.139. Details of structure of M. kotokui (male). A, head, frontal view; B, scape and three basal
funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

465
Figure 140. Details of structure of M. kozakorum (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal
view; F, hind tibia; G, apex of hind tibia. Scale bars = 1 mm.

466
Figure 141. Details of structure of M. kozakorum (queen, paratype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal
view; F, hind tibia. Scale bar = 1 mm.

467
Figure 142. Details of structure of M. kozakorum (male, paratype). A, head, frontal view; B, scape and
four basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view;
E, hind tibia and basitarsus. Scale bars = 1 mm.

468
Figure 143. Details of structure of M. kozlovi (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

469
Figure 144. Details of structure of M. kiyzhanovskii (worker, paratype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

470
Figure 145. Details of structure of M. kurokii (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

471
Figure 146. Details of structure of M. kurokii (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

472
Figure 147. Details of structure of M. kurokii (male). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D,.alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

473
 A, I), E, F, G, H

В, С

Figure 148. Details of structure of M. laurae (queen; A-E, H, holotype; F, G, paratype of M. samnitica).
A, head, frontal view; В, scape, lateral view; С, scape, dorsal view; D, alitrunk and petiole, lateral view;
E, alitrunk and petiole, dorsal view; F, waist, lateral view; G, waist, dorsal view; H, central part of
forewing. Scale bars =1 mm.

474
Figure 149. Details of structure of M. laurae (male) (paratype of M. samnitica). A, head, frontal view;
В, antenna; С, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar =1 mm.

475
Figure 150. Details of structure of M. lemasnei (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, distal part of hind tibia;
F, distal part of middle tibia. Scale bars =1 mm.

476
Figure 151. Details of structure of M. lemasnei (male). A, head frontal view; В, antenna; С, alitrunk
and waist, lateral view; D, alitrunk and waist, dorsal view; E, distal part of hind tibia; F, distal part
of middle tibia. Scale bars =1 mm.

477
Figure 152. Details of structure of M. lobicornis (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, base of scape, anterior view ("foot of scape"); E, alitrunk
and waist, lateral view; F, alitrunk and waist, dorsal view; G, hind tibia. Scale bars = 1 mm.

478
Figure 153. Details of structure of M. lobicornis (queen, paralectotype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal
view; F, hind tibia. Scale bars = 1 mm.

479

\
Figure 154. Details of structure of M. lobicornis (male). A, head, frontal view; B, antenna; C, alitrunk
and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

48O
Figure 155. Details of structure of M. lobulicornis (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal
view; F, hind tibia. Scale bars = 1 mm.

481
Figure 156. Details of structure of M. lobulicornis (queen). A, head, frontal view; B, scape, lateral view;
C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view; F, hind
tibia. Scale bar = 1 mm.

482
Figure 157. Details of structure of M. lobulicornis (male). A, head, frontal view; B, scape and four
basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind
tibia. Scale bar = 1 mm.

483
Figure 158. Details of structure of M. lonae (worker, syntype). A, head, frontal view; B, scape, lateral
view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view;
F, hind tibia. Scale bar = 1 mm.

484
Figure 159. Details of structure of M. lonae (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

485
Figure 160. Details of structure of M. lonae (male). A, head, frontal view; B, scape and four basal
funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

486
Figure 161. Details of structure of M. luteola (worker, paratype). A, head, frontal view; B, scape, lateral
view; С, alitrunk, waist and base of first gastral tergite, lateral view; D, alitrunk, waist and base of first
gastral tergite, dorsal view; E, distal part of hind tibia; F, distal part of middle tibia. Scale bars = 1 mm.

487
Figure 162. Details of structure of M. luteola (queen, paratype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bars = 1 mm.

488
Figure 163. Details of structure of M. luteola (male, paratype). A, head, frontal view; B, antenna;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

.489
Figure 164. Details of structure of M. margaritae (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

490
Figure 165. Details of structure of M. martensi (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

491
Figuré 166. Details of structure of M. martensi (queen, paratype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view. Scale bar = 1 mm.

492
Figure 167. Details of structure of M. mirabilis (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view (sculpture omitted) ; D, alitrunk and waist, dorsal view;
E, hind tibia. Scale bar = 1 mm.

493
Figure 168. Details of structure of M. mixta (queen, holotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

494
Figure 169. Details of structure of M. multiplex (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

495
Figure 170. Details of structure of M. multiplex (queen, paratype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

496
Figure 171. Details of structure of M. myrmicoxena (A-G, queen, lectotype; H, male) (male, after
Kutter 1977). A, H, head, frontal view; В, scape, lateral view; C, alitrunk and waist, lateral view;
D, alitrunk and waist, dorsal view; E, hind tibia; F, distal part of hind tibia; G, distal part of middle tibia.
Scale bars = 1 mm.

497
Figure 172. Details of structure of M. nitida (worker, holotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

498
Figure 173. Details of structure of M. nitida (queen, paratype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

499
Figure 174. Details of structure of M. nitida (male, paratype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

500
Figure 175. Details of structure of M. obscura (worker). A, head, frontal view; B, scape, lateral view;
C, scape, dorsal view; D, base of scape, anterior view ("foot of scape"); E, alitrunk and waist, lateral
view; F, alitrunk and waist, dorsal view; G, hind tibia. Scale bars = 1 mm.

501
Figure 176. Details of structure of M. obscura (queen). A, head, frontal view; B, scape, lateral view;
C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view; F, hind
tibia. Scale bars = 1 mm.

502
Figure 177. Details of structure of M. obscura (male). A, head, frontal view; В, antenna; С, scape and
three basal funicular segments; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view;
F, hind tibia. Scale bars = 1 mm.

503
Figure 178. Details of structure of M. onoyamai (worker, holotype). A, head, dorsal view; B, scape,
posterolateral view; C, scape, dorsal view; D, base of scape, anterolateral view; E, base of scape, anterior
view ("foot of scape"); F, alitrunk and waist, lateral view; G, alitrunk and waist, dorsal view; H, hind
tibia. Scale bars = 1 mm.

504
Figure 179. Details of structure of M. onoyamai (queen, paratype). A, head, dorsal view; B, scape,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal
view; F, hind tibia. Scale bars = 1 mm.

505
Figure 180. Details of structure of M. onoyamai (male, paratype). A, head, frontal view; B, antenna;
С, scape and three basal funicular segments; D, alitrunk and waist, lateral view; E, alitrunk and waist,
dorsal view; F, hind tibia and basitarsus. Scale bars = 1 mm.

506
Figure 181. Details of structure of M. ordinaria (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

507
Figure 182. Details of structure of M. ordinaria (queen, paratype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

508
Figure 183. Details of structure of M. orthostyla (worker, paratype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

509
Figure 184. Details of structure of M. orthostyla (queen, paratype). A, head, frontal view; B, scape,
lateral view; С, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

510
Figure 185. Details of structure of M. orthostyla (male, paratype). A, head, frontal view; B, scape and
three basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view;
E, hind tibia. Scale bar = 1 mm.

5"
Figure 186. Details of structure of M. pachei (worker, lectotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

512
Figure 187. Details of structure of M. pachei (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 ram.

51З
Figure 188. Details of structure of M. pachei (male, paralectotype). A, head, frontal view; B, scape and
three basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view;
E, hind tibia. Scale bar = 1 mm.

514
Figure 189. Details of structure of M. pararitae (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

.515
Figure 190. Details of structure of M. pararitae (queen, paratype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

516
Figure 191. Details of structure of M. pelops (worker, paratype). A, head, dorsal view; B, scape,
posterolateral view; C, scape, dorsal view; D, base of scape, anterolateral view; E, base of scape, anterior
view ("foot of scape"); F, alitrunk and waist, lateral view; G, alitrunk and waist, dorsal view; H, hind
tibia. Scale bars = 1 mm.

517
Figure 192. Details of structure of M. petita (queen, holotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm

518
Figure 193. Details of structure of M. phalacra (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

519
Figure 194. Details of structure of M. phalacra (queen, paratype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

520
Figure 195. Details of structure of M. pisarskii (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bars = 1 mm.

521
Figure 196. Details of structure of M. pisarskii (queen, paratype). A, head, frontal view; B,scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

522
Figure 197. Details of structure of M. pisarskii (male, paratype). A, head, frontal view; B, scape and
three basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view;
E, hind tibia. Scale bar = 1 mm.

523
Figure 198. Details of structure of M. pleiorhytida (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

524
Figure 199. Details of structure of M. pleiorhytida (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

525
Figure 200. Details of structure of M. poldii (worker, holotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

526
Figure 201. Details of structure of M. polyglypta (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

527
Figure 202. Details of structure of M. pulchella (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

528
Figure 203. Details of structure of M. pulchella (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

529
Figure 204. Details of structure of M. ravasinii (worker, syntype). A, head, dorsal view; В, scape,
posterolateral view; С, scape, dorsal view; D, base of scape, anterolateral view; E, base of scape, anterior
view ("foot of scape"); F, alitrunk and waist, lateral view; G, alitrunk and waist, dorsal view; H, hind
tibia. Scale bars = 1 mm.

530
Figure 205. Details of structure of M. ravasinii (queen). A¡ head, frontal view; B, scape, lateral view;
C, .alitrunk and waist, lateral view; D, alitrunk, and waist, dorsali view; E, hind tibia. Scale
bar = 1 mm.

531
Figure 206. Details of structure of M. ravasinii (male). A, head, frontal view; В, scape and three basal
funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bars = 1 mm.

532
Figure 207. Details of structure of M. rhytida (worker, holotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

533
Figure 208. Details of structure of M. rhytida (queen, paratype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

534
Figure 209. Details of structure of M. rhytida (male, paratype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

535
Figure 210. Details of structure of M. rigatoi (worker; A-D, F, holotype). A, head, frontal view;
B, scape, lateral view; C, alitrunk and waist, lateral view; D, propodeal spines and waist, dorsal view;
E, alitrunk and waist, dorsal view; F, hind tibia. Scale bar = 1 mm.

536
Figure 211. Details of structure of M. ritae (worker, lcctotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind .tibia. Scale
bar = 1 mm.

537
Figure 212. Details of structure of M. ritae (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk.and waist, lateral view; D, propodeal spines and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

538
Figure 213. Details of structure of M. rubra (worker, lectotypeofM . laevinodis). A, head, frontal view;
B, scape, lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind
tibia. Scale bars = 1 mm.

539
Figure 214. Details of structure of M. rubra (queen, paralectotype of M. laevinodis). A, head, frontal
view; B, scape, lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view;
E, hind tibia. Scale bars = 1 mm.

540
Figure 215. Details of structure of M. rubra (male, paralectotype of M. laevinodis). A, head, frontal
view; B, antenna; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

541
Figure 216. Details of structure of M. ruginodis (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bars = 1 mm.

542
Figure 217. Details of structure of M. ruginodis (queen, paralectotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bars = 1 mm.

543
Figure 218. Details of structure of M. ruginodis (male, paralectotype). A, head, frontal view; B, antenna;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bars = 1 mm.

544
Figure 219. Details of structure of M. rugosa (worker, lectotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

545
Figure 220. Details of structure of M. rugosa (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

546
Figure 221. Details of structure of M. rugosa (male). A, head, frontal view; B, antenna; C, alitrunk and
waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

547
Figure 222. Details of structure of M. rugulosa (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal
view; F, hind tibia. Scale bars = 1 mm.

548
Figure 223. Details of structure of M. rugulosa (queen, paralectotype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal yiew; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal
view; F, hind tibia. Scale bars = 1 mm.

549
Figure 224. Details of structure of M. rugulosa (male). A, head, frontal view; B, scape and three basal
funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

550
Figure 225. Details of structure of M. rupestris (worker, lectotypc). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

551
Figure 226, Details of structure of M. rupestris (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

552
Figure 227. Details of structure of M. rupestris (male). A, head, frontal view; B, antenna; C, alitrunk
and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

553
Figure 228. Details of structure of M. ruzskyana (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

554
Figure 229. Details of structure of M. ruzskyana (male, paralectotype). A, head, frontal view; B, scape
and three basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal
view; E, hind tibia. Scale bar = 1 mm.

555
Figure 230. Details of structure of M. sabuleti (worker, lectotype). A, head, dorsal view; B, scape,
posterolateral view; С, scape, anterolateral view; D, scape, dorsal view; E, alitrunk and waist, lateral
view; F, alitrunk and waist, dorsal view; G, hind tibia. Scale bars = 1 mm.

556
Figure'231. Details of structure of M. sabuleti (queen). A, head, frontal view; B, scape, lateral view;
C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view; F, hind
tibia. Scale bar = 1 mm.

557
Figure 232. Details of structure of M. sabuleti (male, paralectotype). A, head, frontal view; B, antenna;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

558
Figure 233. Details of structure of M. salina (worker, lectotype). A, head, frontal view; B, scape, lateral
view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view;
F, hind tibia. Scale bars = 1 mm.

559
Figure 234. Details of structure of M. salina (queen, paralectotype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal
view; F, hind tibia. Scale bars = 1 mm.

56O
Figure 235. Details of structure of M. salina (male, paralectotype). A, head, frontal view; B, antenna;
С, scape and four basal funicular segments; D, alitrunk and waist, lateral view; E, alitrunk and waist,
dorsal view; F, hind tibia. Scale bar = 1 mm,

561
Figure 236. Details of structure of M. saposhnikovi (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal
view; F, hind tibia. Scale bars = 1 mm.

5Ó2
Figure 237. Details of structure of M. saposhnikovi (queen). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bars = 1 mm.

563
Figure 238. Details of structure of M. saposhnikovi (male). A, head, frontal view; B, scape and four
basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind
tibia. Scale bar = 1 mm.

564
Figure 239. Details of structure of M. scabrinodis (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal
view; F, hind tibia. Scale bars = 1 m.

565
Figure 240. Details of structure of M. scabrinodis (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

566
Figure 241. Details of structure of M. scabrinodis (male). A, head, frontal view; B, scape and three
basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind
tibia. Scale bar = 1 m.

567
Figure 242. Details of structure of M. schencki (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, base of scape, anterior view ("foot of scape"); E, alitrunk and
waist, lateral view; F, alitrunk and waist, dorsal view; G, hind tibia. Scale bars = 1 mm.

568
Figure243. Details of structure of M. schencki (queen, paralcctotype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal
view; F, hind tibia. Scale bars = 1 mm.

569
Figure 244. Details of structure of M. schencki (male, paralectotype). A, head, frontal view; B, antenna;
С, scape and three basal funicular segments; D, alitrunk and waist, lateral view; E, alitrunk and waist,
dorsal view; F, hind tibia. Scale bars = 1 mm.

570
;Figure 246. Details of structure of M. schoedli (worker, holotype). A, head, frontal view; B, scape,
lateral view; C- alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

571
Figure 247. Details of structure of M. schoedli (queen, paratype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

572
Figure 248. Details of structure of M. schoedli (male). A, head, frontal view; B, antenna; C, alitrunk
and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

573
Figure 249. Details of structure of M. schulzi (worker, holotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

574
Figure 250. Details of structure of M. sculptiventris (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk, waist and first gastral tergite, dorsal view;
E, hind tibia. Scale bar = 1 mm.

575
Figure 251. Details of structure of M. serica (worker, lectotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

576
Figure 252. Details of structure of M. serica (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E; hind tibia. Scale
bar = 1 mm.

577
Figure 253. Details of structure of M. serica (male). A, head, frontal view; B, antenna; C, head
(sculpture omitted), alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

578
Figure 254. Details of structure of M. siciliana (worker, holotype). A, head, frontal view; B, scape,
posterolateral view; C, scape, dorsal view; D, base of scape, anterolateral view; E, base of scape, anterior
view ("foot of scape"); F, alitrunk and waist, lateral view; G, alitrunk and waist, dorsal view; H, hind
tibia. Scale bars = ; 1 mm.

579
Figure 255. Details of structure of M. siciliana (queen, paratype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal
view; F, hind tibia. Scale bars = 1 mm.

58О
Figure 256. Details of structure of M. siciliana (male, paratype). A, head, frontal view; B, antenna;
С, scape and three basal funicular segments; D, alitrunk and waist, lateral view; E, alitrunk and waist,
dorsal view; F, hind tibia. Scale bars = 1 mm.

58i
Figure 257. Details of structure of M. sinensis (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

582
Figure 258. Details of structure of M. sinoschencki (worker, holotype). A, head, dorsal view; B, scape,
posterolateral view; C, scape, dorsal view; D, base of scape, anterolateral view; E, base of scape, anterior
view ("foot of scape"); F, alitrunk and waist, lateral view; G, alitrunk and waist, dorsal view; H, hind
tibia. Scale bars = 1 mm.

583
Figure 259. Details of structure of M. slovaca (worker, syntype). A, head, frontal view; B, scape, lateral
view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view;
F, hind tibia. Scale bars = 1 mm.

584
Figure 260. Details of structure of M. slovaca (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

585
Figure 261. Details of structure of M. slovaca (male). A, head, frontal view; B, scape and four basal
funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bars = 1 mm.

586
Figure 262. Details of structure of M. smythiesii (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

587
Figure 263. Details of structure of M. smythiesii (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

588
Figure 264. Details of structure of M. specioides (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view;E, alitrunk and waist, dorsal
view; F, hind tibia. Scale bars = 1 mm.

589
Figure 265. Details of structure of M. specioides (queen, paralectotype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal
view; F, hind tibia. Scale bars = 1 mm.

590
Figure 266. Details of structure of M. specioides (male, paralectotype). A, head, frontal view;
B, antenna; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia and
basitarsus. Scale bar = 1 mm.

591
Figure 267. Details of structure of M. spinosior (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal
view; F, hind tibia. Scale bars = 1 mm.

592
Figure 268. Details of structure of M. spinosior (queen). A, head, frontal view; B", scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.

593
Figure 269. Details of structure of M. spinosior (male). A, head, frontal view; B, scape and four basal
funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

594
Figure 270. Details of structure of M. stangeana (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal
view; F, hind tibia. Scale bars = 1 mm.

595
Figure 271. Details of structure of M. stangeana (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

596
Figure 272. Details of structure of M. stangeana (male). A, head, frontal view; B, antenna; C, alitrunk
and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

597
Figure 273. Details of structure of M. sulcinodis (worker, lectotype). A, head, frontal view; B, scape,
lateral view; O, base of scape, anterior view ("foot of scape"); D, alitrunk and waist, lateral view;
E, alitrunk and waist, dorsal view; F, hind tibia. Scale bars = 1 mm.

598
Figure 274. Details of structure of M. sulcinodis (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

599
Figure 275. Details of structure of M. sulcinodis (male). A, head, frontal view; B, scape and three basal
funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

600
Figure 276. Details of structure of M. symbiotica (worker, holotype). A, head, frontal view; B, antenna,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk, waist and gaster,
dorsal view; F, distal part of hind tibia; G, distal part of middle tibia. Scale bars = 1 mm.

601
Figure 277. Details of structure of M. taibaensis (worker, paratype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1mm.

602
Figure 278. Details of structure of M. taibaensis (queen, paratype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view;D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1mm.

603
Figure 279. Details of structure of M. tamarae (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

604
Figure 280. Details of structure of M. tamarae (queen, paratype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

605
Figure 281. Details of structure of M. tenuispina (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

6C)6
Figure 282. Details of structure of M. tenuispina (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

607
Figure 283. Details of structure of M. tenuispina (male). A, head, frontal view; B, scape and three basal
funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

608
Figure 284. Details of structure of M. tibetana (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. •
Scale bar = 1 mm.

609
Figure 285. Details of structure of M. tibetana (male). A, head, frontal view; B, scape and three basal
funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

Ó10
Figure 286. Details of structure of M. titanica (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

611
Figure 287. Details of structure of M. tobiasi (worker, holotype). X^head, frontal view; B, scape, lateral
view; C, scape, dorsal; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view; F, hind
tibia. Scale bars = 1 mm.

612
Figure 288. Details of structure of M. tobiasi (queen, paratype). A, head, frontal view; B, scape, lateral
view; C, scape, dorsal; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view; F, hind
tibia. Scale bars = 1 mm.

613
Figure 289. Details of structure of M. tobiasi (male, paratype). A, head, frontal view; B, antenna;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

614
Figure 290. Details of structure of M. transsibirica (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, base of scape, anterior view ("foot of scape"); D, alitrunk and waist, lateral view;
E, alitrunk and waist, dorsal view; F, hind tibia. Scale bars = 1 mm.

615
Figure 291. Details of structure of M. transsibirica (queen, paratype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bars = 1 mm.

6l6
Figure 292. Details of structure of M. transsibirica (male, paratype). A, head, frontal view; B, scape and
four basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk.and waist, dorsal view;
E, hind tibia. Scale bar = 1 mm.

617
Figure 293. Details of structure of M. tschekanovskii (worker, paratype). A, head, frontal view;
B, scape, lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind
tibia. Scale bar = 1 mm.

6l8
Figure 294. Details of structure of M. tschekanovskii (male, holotype). A, head, frontal view;
B, antenna; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
' Scale bar = 1 mm.

619
Figure 295. Details of structure of M. tulinae (worker, holotype). A, head, frontal view; B, scape, lateral
view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view;
F, hind tibia; G, apex of hind tibia. Scale bars = 1 mm.

620
Figure 296. Details of structure of M. tulinae (queen, paratype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

621
Figure 297. Details of structure of M. íulinae (male, paratype). A, head, frontal view; B, antenna;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

622
Figure 298. Details of structure of M. turcica (worker, lectotype). A, head, frontal view; B, scape, lateral
view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view;
F, hind tibia. Scale bar = 1 mm.

623
Figure 299..Details of structure of M. turcica (queen). A, head, frontal view; B, scape, lateral view;
C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view; F, hind
tibia. Scale bar = 1 mm.

624
Figure 300. Details of structure of M. turcica (male). A, head, frontal view; B, antenna; C, alitrunk and
waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale bar = 1 mm.

625
Figure 301. Details of structure of M. urbanii (worker, holotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

626
Figure 302. Details of structure of M. vandeli (worker). A, head, frontal view; B, scape, lateral view;
C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view; F, apex of
hind tibia. Scale bars = 1 mm.

627
Figure 303. Details of structure of M. vandeli (queen, lectotype). A, head, frontal view; B, scape, lateral
view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view;
F, hind tibia; G, apex of hind tibia. Scale bars = 1 mm.

628
Figure 304. Details of structure of M. vandeli (male, paralectotype). A, head, frontal view; B, antenna;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia and basitarsus.
Scale bar = ,1 mm.

629
Figure 305. Details of structure of M. varisculpta (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

630
Figure 306. Details of structure of M villosa (worker, holotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

631
Figure 307. Details of structure of M. villosa (queen, paratype). A, head, frontal view; B, alitrunk and
waist, lateral view. Scale bar = 1 mm.

632
Figure 308v Details of structure of M. vittata (worker, holotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

633
Figure 309. Details of structure of M. wardi (worker, holotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view (puntutres omitted);
E, hind tibia. Scale bar = 1 mm.

634
Figure 310. Details of structure of M. wardi (queen, paratype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1mm.

635
Figure 311. Details of structure of M. wardi (male, paratype). A, head, frontal view; B, scape and four
basal funicular segments; C, alitrunk and waist, lateral view; D, hind tibia. Scale bar = 1 mm.

636
Figure 312. Details of structure of M. weberi (worker, holotype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk'and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

637
Figure 313. Details of structure of M. weberi (queen, paratype). A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

638
Figure 314. Details óf structure of M. weii (worker, holotype).-A, head, frontal view; B, scape, lateral
view; C, alitrunk and waist, lateral view; D, alitrunk, and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

639
Figure 315. Details of structure of M. wesmaeli (worker, lectotype). A, head, frontal view; B, scape,
lateral view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal
view; F, hind tibia. Scale bars = 1 mm.

640
Figure 316. Details of structure of M. wesmaeli (queen). A, head, frontal view; B, scape, lateral view;
C, alitrunk and waist," lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

641
Figure 317. Details of structure of M. wesmaeli (male). A, head, frontal view; B, scape and three basal
funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

642
Figure 318. Details of structure of M. williamsi (worker, holotype). A,, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral, view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

643
Figure 319. Details of structure of M. wittmeri (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

644
Figure 320. Details of structure of M. xavieri (worker, holotype). A, head, dorsal view; B, scape,
posterolateral view; C, scape, dorsal view; D, base of scape, anterolateral view; E, base of scape, anterior
view ("foot of scape"); F, alitrunk and waist, lateral view; G, alitrunk and waist, dorsal view; H, hind
tibia. Scale bars = 1 mm..

645
Figure 321. Details of structure of M. xavieri (queen, paratype). A, head, dorsal view; B, scape,
posterolateral view; C, scape, dorsal view; D, base of scape, anterolateral view; E, base of scape, anterior
view ("foot of scape"); F, alitrunk and waist, lateral view; G, alitrunk and waist, dorsal view; H, hind
tibia. Scale bars = 1 mm.

646
Figure 322. Details of structure of M. xavieri (male, paratype). A, head, frontal view; B, antenna;
C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

647
Figure 323. Details of structure of M. yamanei (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1 mm.

648
Figure 324. Details of structure ol M. yunnanensis (worker, holotype). A, head, frontal view; B, scape,
lateral view; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia.
Scale bar = 1/inm.

649
Figure 325. Details of structure of M. zojae (worker, holotype). A, head, frontal view; B, scape, lateral
view; C, scape, dorsal view; D, alitrunk and waist, lateral view; E, alitrunk and waist, dorsal view;
F, hind tibia. Scale bars = 1 mm.

650
Figure 326. Details of structure of M. zojae (queen, paratype). A, head, frontal view; B, scape, lateral
view; C, alitrunk arid waist, lateral view; D, alitrunk and waist, dorsal view; E, hind tibia. Scale
bar = 1 mm.

651
Figure 327. Details of structure ofM.zojae (male, paratype). A, head, frontal view; B, scape and three
basal funicular segments; C, alitrunk and waist, lateral view; D, alitrunk and waist, dorsal view; E, hind
tibia. Scale bar = 1 mm.

652
3.4. Fossil species, undoubtedly belonging to the genus Myrmica

Myrmica eocenica Radchenko, Dlussky et Elmes, 2007

(Fig. 328)

Myrmica eocenica Radchenko, Dlussky et Elmes, 2007: 1497, w, Russia, Baltic Amber.

Material examined. Holotype, w, No PIN, 964/479, Kaliningrad Prov, of Russia, Baltic Amber,
Late Eocene (PIN-MOSCOW).
Etymology. M. eocenica: named for the Eocene Epoch when the type specimen was
trapped in amber, the name Eocene means dawn of the recent period, is derived from the
Greek F.OO (eos) = dawn and KOIVOCJ (koinos) = recent.
Notes. M. eocenica (as all other known Amber Myrmica, except for M. paradoxa) is
very similar morphologically to the modern species from the ritae species group. It differs
from M. longispinosa by the sharply pointed apically propodeal lobes, by the strongly
divergent propodeal spines, and by its somewhat bigger size (> 6 mm vs. ca. 5 mm). Besides
the well-developed pectinate spurs on the middle and hind tibiae, it differs from M. rudis by
the shape of the propodeal spines and petiole: in M. rudis the spines are somewhat curved
and directed backward and upward and its petiole has a much shorter peduncle and sub-
triangular node. M. eocenica most resembles M. intermedia by the shape and sculpture of
the body but differs by its straight and strongly divergent propodeal spines and body length
(> 6 mm vs. 4.7 mm) (see also Key, Chapter 5.8).

Myrmica intermedia (Wheeler, 1915)

(Fig. 329)

Nothomyrmica intermedia Wheeler, 1915: 61, w, East Prussia (now - Kaliningrad Prov, of Russia),
Baltic Amber, Late Eocene; Radchenko 1994b: 48; Bolton 1995: 292.
Myrmica intermedia: Radchenko, Dlussky and Elmes 2007: 1496.

Etymology. M. intermedia: from the word intermediate derived from the Latin prefix
inter - between and media = in the middle; Wheeler (1915: 61) wrote: "Closely resem-
bling N. rudis and almost intermediate between this form and Myrmica longispinosa in
many particulars".
Notes. This species was originally described in the extinct genus Nothomyrmica, dif-
fering from Myrmica by the single character - lack of the pectinate spur on the middle and
hind tibiae. Nowadays this feature is less important in the Myrmica taxonomy than believed
at the beginning of 20th century, and we synonymised Nothomyrmica with Myrmica (Rad-
chenko et al. 2007).
The holotype of M. intermedia is most probably lost (see also Notes to M. longispinosa,
below) and no other specimens are known. It differs from M. rudis by the straight and
directed backward propodeal spines, but is similar to M. longispinosa differing from the
latter mainly by the absence of the pectinate spurs on the middle and hind tibiae (see also
Key, Chapter 5.8).

653
 Myrmica longispinosa Mayr, 1868

Myrmica longispinosa Mayr, 1868: 86, w, East Prussia (now- Kaliningrad Prov, of Russia), Baltic
Amber, Late Eocene; Wheeler 1915: 59; Radchenko 1994b: 48; Bolton 1995: 281; Radchenko,
Dlussky and Elmes 2007: 1495.

Etymology. M. longispinosa : from a combination of the Latin words longus = long and
spinosa (adj. F) = thorned, to described the obviously long propodeal spines.
Notes. Mayr (1868) described this species based on a single worker from the Baltic
Amber. Later Wheeler (1915) studied Mayr's holotype and one additional not-numbered
worker in the Königsberg collection (Germany at that time, nowadays Kaliningrad in Rus-
sia) . At present the holotype of M. longispinosa is absent from the Mayr's collection in
Naturhistorisches Museum Wien (Ponomarenko and Schultz 1988). The Königsberg^
amber collection was apparently mostly lost during World War II, but nowadays a small
part of it is preserved in the collection of the Institut und Museum für Geologie und Paläon-
tologie der Universität Göttingen (Germany). Based on the database of the fossils of
Göttingen's Museum no specimens of M. longispinosa were found there (Perkovsky, pers.
comm. 2004). Hence, we believe that both Mayr's holotype and Wheeler's material, be-
longing to this species, are lost.
Consequently, we have never seen any material belonging to this species. Based on Mayr's
original description and Wheeler's comments this species is characterized by a coarse sculp-
ture, the presence of a pectinate spur on the hind tibiae, a long antennal scape, long
propodeal spines, which are directed backwards and are very feebly divergent, subparallel (in
dorsal view). Unfortunately, Mayr's drawing of this species is too schematic to be really help-
ful (see Mayr 1868, Fig. 86). For the separation of this species from other Amber Myrmica
see also Key, Chapter 5.8.

Myrmica paradoxa Radchenko, Dlussky et Elmes, 2007

(Fig. 330)

Myrmica paradoxa Radchenko, Dlussky et Elmes, 2007: 1496, w, Germany, Saxonian (= Bitterfel-
dian) Amber, Late Eocene.

Material examined. Holotype and two paratypes, w in the same piece of amber, No F-199,
Germany, Saxonian (= Bitterfeldian) Amber, Late Eocene (HAMBURG).
Etymology. M. paradoxa: named from the Greek word napaôofy (paradoxus) = para-
doxical or contrary to expectations, to indicate that this species has an unusual combina-
tion of features compared,to any extant or extinct Myrmica species.
Notes. M. paradoxa possesses very short propodeal denticles: such a shape of propodeal
armature is very peculiar and unknown in any extant or other fossil Myrmica species.
Additionally, while it has a sculpture and shape of the head, pilosity of the body, shape of
petiole and postpetiole and a distinct pectinate spurs on the hind tibiae that are typically
Myrmica, its distinctly three-jointed antennal club and the generally short and robust
alitrunk are also quite atypical for Myrmica. Unfortunately, the maxillary and labial palpes,
which can be diagnostic of certain genera, are invisible. We suppose that M. paradoxa
represents an extinct lineage within the genus (see also Radchenko et al. 2007).

654
 Myrmica rudis (Mayr, 1868)
(Fig. 331)

Macromischa rudis Mayr, 1868: 85, w, East Prussia [now - Kaliningrad Prov., Russia], Baltic Amber,
Late Eocene; Dalla Torre 1893: 120; André 1895: 82; Handlirsch 1907: 875.
Nothomyrmica rudis: Wheeler 1915: 23; Radchenko 1994b: 48; Bolton 1995: 292.
Myrmica rudis: Radchenko, Dlussky and Elmes 2007: 1495.

Material examined. One badly preserved worker, No. 1945/6, Baltic Amber, Poland, Late Eocene
(MZ-WARSAW).
Etymology. M. rudis: from the Latin word rudis, probably in its sense of being "coarse"
or "rough", probably to indicate its very coarse body sculpture.
Notes. The types of M. rudis are most probably lost (see Notes to M. longispinosa, above).
Although the specimen we studied is in quite bad condition and the transparency of the piece
of amber is poor, the general shape of the ant is visible, including the shape of the petiole and
the propodeal spines that are directed upward and backward (characteristic for M. rudis).
However, at this time we think it would be unreasonable to designate this specimen as the
neotype of M. rudis. For its separation from other species see Key, Chapter 5.8.

Figure 328. M. eocenica (worker, holotype). A, body, dorsal view; В, body, dorso-lateral view (sculpture
omitted). Scale bars = 1 mm.

655
 Figure 329. M. intermedia (worker, holotype) (after Wheeler 1915).

Figure 330. M. paradoxa (worker; A, holotype; B, paratype). Scale bars = 1 mm

656
 Figure 331. M. rudis (worker, holotype) (after Wheeler 1915).

3.5. Names incertae sedis in Myrmica

Myrmica bessarabica Nasonov, 1889

Myrmica scabrinodis var. bessarabica Nasonov, 1889: 36, w, Moldova.

Etymology. M. bessarabica: named for Bessarabia where the types were taken, Bessara­
bia was the historical name for the region that more or less equates to modem Moldova.
Notes. Nasonov (1889) described M. scabrinodis var. bessarabica from Bessarabia based
on several workers. The original description is very brief and incomplete: "Antennae quite
weakly bent at the base, forming very blunt angle, above which elevates blunt tooth.
Propodeum between spines smooth and shiny, with one or two transversal impressions.
Frontal area distinctly striated" (loc. cit., p. 36; our translation from Russian).
Ruzsky (1905) noted that he did not see the types of var. bessarabica, but investigated
two workers from Nasonov's collection (MOSCOW) with the labels "Бессараб1я". By his
opinion, these specimens "belong to the typical M. scabrinodis, differing by the only almost
reduced striation between propodeal spines" (loc. cit., p. 688; our translation from Russian).
It is important to note that Ruzsky also considered several modern "good" species, in­
cluding M. specioides and M. sabuleti, under the name M. scabrinodis. Later Emery (1908a,
1921) also regarded this form as a variety of M. scabrinodis, but Weber (1948) and
Sadil (1952) without any comments considered M. bessarabica as junior synonym of
M. sabuleti.

657
 More recently, Dlussky (Dlussky et al. 1990; Atanassov, Dlussky 1992) found a speci­
men (worker) in the collection of Nasonov (MOSCOW) labelled as "Бессараб1я", and be­
lieved that it belongs to the type series of var. bessarabica. As a result, he labelled it as the
"holotype" of var. bessarabica (incorrectly because this specimen at best, maybe considered
as the lectotype or perhaps a neotype) and raised M. bessarbica to the species level. As this
specimen is indistinguishable from M. specioides, Dlussky considered M. bessarabica as
a senior synonym of the latter name (and of its junior synonyms, see Notes to M. specioides,
above). Several authors (e.g. Arakelyan 1994; Radchenko 1994b, e, d; Bolton 1995) ac­
cepted this Dlussky's treatment.
However, Seifert (2002) disagreed with Dlussky's opinion and proposed to consider
M. bessarabica as incertae sedis in the genus Myrmica. He speculated about origin of the la­
bels on the pin with the worker of "M. bessarabica" in Nasonov's collection, and concluded
that the label "Бессарабш" was not written by Nasonov because it is too clean (not dusty).
Of course, we can not be absolutely sure that this was the original Nasonov's label, but it is
obvious from the spelling that it has written before 1918 (the letter "i" was deleted from the
Russian grammar since 1918). So, it is quite possible that this is Nasonov's original label.
However, the second label, "Holotype Myrmica scabrinodis bessarabica Nassonov", was
added by Gennady Dlussky (pers, comm.) at the end of 1980's. There is no doubt that this
"holotype" specimen is conspecific with M. specioides, but the most important question re­
mains: does this specimen belong to the original type series of var. bessarabica, or belong to
"M. scabrinodis" series also recorded by Nasonov from Bessarabia?
The second Seifert's (2002) argument against considering var. bessarabica as
a synonym of M. specioides is based on the original Nasonov's description: "... Antennae
quite weakly bent at the base, forming very blunt angle, above which elevates blunt tooth..."
(our translation from Russian). If one interprets the direction of this "tooth" as being
vertical then obviously the "real" var. bessarabica does not belong to the scabrinodis species
group, but should be placed to the schencki- or fobz'corm's-groups (most probably being
conspecific with M. deplanata). However, in the absence of any additional arguments, we
accept Seifert's proposition that M. bessarabica is best considered as incertae sedis in the
genus Myrmica (see also Radchenko and Elmes 2004).

Myrmica breviceps Smith F., 1878

Myrmica breviceps Smith F., 1878: 12, q, India or China (?).

Etymology. M. breviceps: from a combination of the Latin word brevis = short and
ceps derived from Latin caput = head, to indicate that this species has a relatively short
head.
Notes. The types of this species appear to be lost. No locality and date were given in the
original description, but as the paper, published by the government of India, related to the
scientific results of the Second Yarkand Mission, one might suppose that the specimen was
taken somewhere in the region of Yarkand, now in Xinjiang Province of China (between the
Takla Makan desert and Tadjikistan and Kyrgyzstan) and was a Middle Asian species.
However, this is not certain as other material described in the same paper were taken in
the Punjab and the Pamir mountains en route to Yarkand. Smith (1878, p. 12) states that
"This insect very closely resembles two or three of the British species, particularly Myrmica

658
sulcinodis; its head is, however, shorter than that of the British insects"; otherwise the
description is not very helpful.
The original description (Smith F. 1878, p. 12) reads: "Rufo-ferruginous; the head
strongly longitudinally striated; a small smooth shining space at the base of the clypeus,
which is deeply longitudinally grooved; the mandibles striated. Thorax above with a black
patch on each side of the mesothorax, and another at the anterior portion; the metatho-
racic spines short, stoutand acute. The first node of the abdomen longitudinally rugose,
the second transversely so; the abdomen smooth and shining, ferruginous at the base; the
rest dark rufo-fuscous, nearly black, and with scattered erect pubescence. Female, length
21/2 lines [i.e. ca 5.25 mm].".
Overall this description of M. breviceps does not give any reason to remove this species
from genus Myrmica. Even if we assume it was a Middle Asian species, it is impossible to
relate the original description of M. breviceps with any confidence to any of the species of
Middle Asia, including M. sulcinodis. Therefore in the absence of type specimens, we pro-
pose to consider it as unidentifiable taxon, incertae sedis in the genus Myrmica.

Myrmica cursor Smith F., 1878

Myrmica cursor Smith F., 1878: 11, q, India or China (?).

Etymology. M. cursor: from the Latin cursus = course, probably used in the same sense
as in archeology where it describes long parallel banks or ridges, to indicate the striate body
of this species "capite thoraceque logitudinaliter striatis".
Notes. The situation regarding this species' is exactly the same as that of M. breviceps
(above), no location was given so that it could have been collected anywhere between the
Punjab and Yarkand. The whereabouts of the types are unknown.
The original description of this species (Smith F. 1878, p. 11) follows: "Obscure fer-
ruginous, with the head sometimes nearly black, or with the sides more or less ferruginous;
the antennae with the scape and a few of the nasal joints of the flagellum pale ferruginous,
the rest fusco-ferruginous; the head longitudinally but irregularly striated, the striae at its
sides formed of confluent punctures. Thorax longitudinally striated, oblong-ovate, and
having a longitudinal ferruginous space above, enclosed by a black margin; the sides and be-
neath ferruginous; the metathorax with two short, stout, acute, compressed spines; the legs
pale ferruginous. Abdomen globose, smooth and shining; the first node of the petiole wedge-
shaped when viewed sideways, and coarsely rugose, the second node globose and sub-
rugose. The scape of the antennae in this species is bent and slender at the base as in the
M. ruginodis of Europe, which is closely resembles; it isj however, a rather smaller insect.
Female, length 2 3/4 lines [i.e. ca 5.8 mm].".
Most probably M. cursor belongs to the genus Myrmica, but without types it is uniden-
tifiable, and we propose consider it incertae sedis in the genus Myrmica.

Myrmica granulinodis Nylander, 1846

Myrmica granulinodis Nylander, 1846b: 1060, q, m, Russia.

659
 Etymology. M. granulinodis: name from granular = rough texture or grainy, combined
with the Latin word nodus = knot or lump, to describe the sculpture of the waist region.
Notes. This species was described by Nylander (1846b) from Siberia (no precise local-
ity given). During the examination of the Nylanderi ant collection (HELSINKI) (see Rad-
chenko 2007a) we did not locate the types of M. granulinodis and suppose they are most
probably lost.
The description of this species in very brief and hardly usable since most of the impor-
tant diagnostic features of Myrmica species were not characterized. It is obvious only that
the scape of females is angularly curved ("...scapo antennarum basi geniculato integro..."),
and the male's scape is feebly curved at the base and quite long, equal to the seven
following funicular segments ("...at scapus basi curvatus tertiam partem totius antennae
parum excedens vel longitudinae articulorum septem sequentium...") (loc. cit., p. 1060; full
translation of the original description to English see Weber 1950: 205). Later Nylander
(1856, p. 81) supposed that this species may be conspecific with M. denticornis (junior
synonym of M. lobicornis).
Subsequent authors considered M. granulinodis as a good species (Smith F. 1858; Dalla
Torre 1893; Ruzsky 1905), as a variety/subspecies of M. scabrinodis (André 1883; Weber
1950), or even as a junior synonym of M. sabuleti (Emery 1908a; Sadil 1952), even though
Nylander's name had priority.
Based on the original description we can only place M. granulinodis to the lobicornis
species group, but without types it is impossible to determine which of the Siberian species
it might be; therefore we confirm Radchenko's (1994f) proposal to consider M. granulino-
dis as incertae sedis in the genus Myrmica.

Myrmica ruginodis khamensis Ruzsky, 1915

Myrmica ruginodis var. khamensis Ruzsky, 1915: 441, w, Tibet.

Etymology. M. khamensis: named for Kham (also spelled as Kam) with the Latin suf-
fix ensis = place of origin, to indicate that the type species were taken in modem southeast
Tibet, in the old region of Kham which also included parts of west China.
Notes. The types of var. khamensis are most probably lost, and the original description
Ruzsky 1915, p. 441) is very brief: "Worker. Differs from type [e.g. M. ruginodis] by smooth
and shiny surface of propodeum between spines. Propodeal spines somewhat shorter and
less developed. Dorsal surface of propodeum more flattened. Body sculpture finer
["smaller"], not so sparse and coarse. Colour darker.
Localities: 1) vil. Dzherku on the riv. Tzan-da, Kam, basin of the riv. Yangtze, mid.
viii. 1900; 2) valley of the riv. Tetunga, NE Tibet, Nan-Shan Range, beginning of ix.1901, leg.
Kozlov" (our translation from Russian). It was taken by Kozlov on the Imperial Russian
Geographical Society's 5 th expedition to Kham and Mongolia (1899-1901).
Based on this description, Ruzsky's (1915) figures and on the modern know-
ledge of the Tibetian fauna, M. ruginodis var. khamensis at best can be placed to the
smythiesii species group, but can not be identified to any modern taxon, therefore we pro-
pose to consider it incertae sedis in the genus Myrmica.

66o
 Myrmica scabrinodis intermedia Kuznetsov-ugamsky, 1927

Myrmica scabrinodissubsp.intermedia Kuznetsov-Ugamsky, 1927: 194, w, Kazakhstan [unresolved

junior secondary homonymofMyrmica intermedia (Wheeler, 1915: 61)]. ...

Etymology. M. intermedia: from the word intermediate derived from Latin words inter
= between and médius — in the middle, to show that it shares features of M. scabrinodis and
M. lobicornis (according Kuznetsov-Ugamsky's opinion).
Notes. Types of M. scabrinodis subsp. intermedia are most probably lost, the original de-
scription (Kuznetsov-ugamsky, 1927: 194) follows: "Epinotumdornen lang, scharf
zugespitzt, etwas nach unten gebogen, mehr nach hinten, als nach oben gerichtet. Mesoe-
pinotaleinsenkung scharf und ziemlich tief. Metasternalzähnchen eckig, abgestumpft. Erstes
Petiolusglied wie bei M. scabrinodis lobicornis ziemlich kurz. Fühlerschaft an der geknick-
ten Stelle dornartig vorgezogen. Körperfärbung hellgelb, etwas rötlich, Stirn bräunlich;
Abdomen oben schwärzlich-braun. Bisweilen der Grundton ist dunkler, bräunlich-rot. Kör-
perlange 3,6-4,5 mm.
Окрестности Копала, под камнем (найдена только одна колония П. П. Архангель­
ским) [Vicinity of Kopal, under stone (only one colony was found by P.P. Arkhangelsky)]".
Additionally, in the Key Kuznetsov-ugamsky (loc. cit., p. 190) wrote: "... Fühlerschaft
geknickten Stelle breit lappenförmig ausgezogen...".
It is clear from the description that the scape of workers of this form has a lobe at the
scape base, and therefore it can not belong to M. saposhnikovi (for details see Key, Chap-
ter 5.4). From the original description and Key it is hard to understand whether the lobe
projects horizontally (like in the scabrinodis-group) or vertically (like in the lobicornis-
group). However, its petiole seems similar to that of M. lobicornis ("...Erstes Petiolusglied
wie bei M. scabrinodis lobicornis ziemlich kurz...", loc. cit., p. 194), so on balance we
suppose that subsp. intermedia belongs to the lobicornis-group. If so, it could be conspecific
with M. kirghizorum and the name inermedia would have priority; but, to change the
current taxonomic status of a recognised species based on such unprovable speculation
would be highly undesireable. Therefore until types are found, we consider subsp. interme-
dia as incertae sedis in Myrmica.

Myrmica luctuosa SmithF., 1878

Myrmica luctuosa Smith F., 1878: 12, m, India.

Etymology. M. luctuosa: the word Latin adjective (F) luctuosa = mournful or grievous,
we assume that Smith chose this name because the male wings were particularly pale.
Notes. The original description of the male of this species (Smith F. 1878, p. 12) follows:
"Black; head and thorax longitudinally striated; the metathorax with transverse curved
striae, and with two stout compressed spines, its apex obscurely rufo-piceous; the club of
the antennae paler than rest of the antennae. The wings pale fulvo-hyaline and iridescent;
the stigma and nervures pale ferruginous; the legs long and slender, with the apical joints
of the tarsi pale testaceous. Abdomen smooth and shining, and with a few scattered pale
hairs; the nodes of the abdomen rugose, the first oblong, the second globose. Male, length
2 3/4 lines [i.e. ca 5.8 mm]. Hab.-Muree (Mari), Punjab hills".

661
 The type specimen seems to be lost. Based on its description alone, M. luctuosa could
belong to genus Myrmica but equally to some other Myrmicinae genera. As it is impossible
to guess at any species from the description, we propose to consider it incertae sedis in the
genus Myrmica.

Myrmica rhynchophora Foerster, 1850

Myrmica rhynchophora Foerster, 1850b: 488, m, Algeria. Junior synonym of Myrmica rubida: Mayr
1861: 63 (now-Manica). Incertae sedis in the genus Myrmica: current status.

Etymology. M. rhynchophora: probably named because it resembled a weevil (genus

Rhyncophora, derived from the Greek for snout-headed).
Notes. This species was described from a male ant taken in Algeria and the type speci-
men appears to be lost. Mayr (1861) without giving any reasons synonymised
M. rhynchophora with Myrmica rubida (now - Manica). However, no Manica species have
ever been recorded from Algeria so the proposed synonymy seems incorrect. On the other
hand, there are several Myrmica species known from Algeria and we therefore consider
M. rhynchophora incertae sedis in the genus Myrmica.

Myrmica scabrinodis ussuriensis Kuznetsov-ugamsky, 1928

Myrmica scabrinodissubsp.ussuriensis Kuznetsov-ugamsky, 1928: 36, w, Russia.

Etymology. M. ussuriensis: a combination of the name Ussuri and the Latin suffix ensis
= place of origin, to indicate that it is a resident of Ussuri Region of the Russian Far East.
The Ussuri river forms a boundary with China and flows north to join the Amur river.
Notes. The types of subsp. ussuriensis most probably are lost. Kuznetsov-ugamsky
(1928, p. 36, figs. 20, 21) described this form based on workers from the Russian Far East
(Nikolsk-Ussuryisky). The most important diagnostic features, given in the original de-
scription, are: "By the structure of the scape it resembles M. scabrinodis lobicornis from
Transbaikalia (sensu Ruzsky 1905), particularly scape strongly flattened laterally and with
well developed raised teeth on the bend... Petiole (seen in profile) angulated, propodeal
spines widened at the base, narrowed posteriorly and acute, shorter than dorsal propodeal
surface... Alitrunk dorsum (in profile) straight, with a barely visible metanotal groove. By
this feature it resembles var', deplanata... Body length 4.8-5.0 mm" (our translation from
Russian).
Based on this rather incomplete description and on the quite schematic figures of the
scape, waist and propodeal spines, we could place subsp. ussuriensis equally to the
lobicornis- or schencki species groups. In the first case, it might be a senior synonym of
M. eidmanni and in the second case, a senior synonym of M. koreana. Given the ambigu-
ous description and as we can not find the type specimens of subsp. ussuriensis, we believe
that it is sensible to accept Radchenko's (1994f) proposal to consider this form incertae
sedis in the genus Myrmica.

662
3.6. Nonima nuda in the genus Myrmica

Myrmica scabrinodis var. atlantica Stärcke, 1942: 27, The Netherlands. Nomen nudum.
Myrmica busoni Chapman and Capco, 1951: 125, India. Nomen nudum (attributed to Ruzsky).
Myrmica daurica Motschoulsky, I860: 500, Russian Far East. Nomen nudum. We found worker
specimen in MOSCOW, with an original label reading "Myrmica daurica", and determined it as
M. eidmanni.
Myrmica margaritae subsp. inornata Menozzi, 1941: 15, China (in Eidmann 1941). Nomen nudum.
Myrmica bergi subsp. iranica Arnoldi, 1948: 211, Azerbaijan. Nomen nudum. We examined workers
from Diabar basin (SE Azerbaijan) (MOSCOW), labelled by Arnoldi as "M. bergi iranica", and
determined them as M. bergi.
Myrmica kirbii Stephens, 1829: 356. Nomen nudum.
Myrmica ruginodis (f. intrm.) M. laevinodoruginodis Ruzsky, 1896: 73, Russia. Nomen nudum (at-
tributed to Forel).
Myrmica lobicornis var. lobata Kutter, 1977: 66. Nomen nudum (attributed to Finzi).
Myrmica rubra var. lobicornosulcidnodis Forel, 1874: 77, Switzerland. Nomen nudum.
Myrmica lucidula Smith F. 1857b: 39, Great Britain. Nomen nudum. Referred by Donisthorpe (1915a:
66) to Formicoxenus nitidulus (Nylander).
Myrmica deplanata subsp. lulakeranica Arnoldi, 1948: 211, Azerbaijan. Nomen nudum. We examined
workers from Talysh (Azerbaijan) (MOSCOW), labelled by Arnoldi as "M. deplanata lulakera-
nica", and determined them as M. deplanata.
Myrmica melanocephala Mayr, 1855: 411, Austria. Nomen nudum (attributed to Koch). Referred by
Mayr (1855: 411) to Leptothorax acervorum (Fabricius).
Myrmica neglecta Stärcke, 1942: 27, Netherlands. Nomen nudum. Referred by Stärcke (1942: 27) to
Myrmica puerilis.
Myrmica nodicornis Chappell, 1886: 58, Great Britain. Nomen nudum. Referred by Donisthorpe
(1915a: 134) to Myrmica lobicornis.
Myrmica predella Kutter, 1977: 6, fig. 30e, Switzerland. Nomen nudum.
Myrmica rugosa Mayr, 1855: 411. Nomen nudum (attributed to Koch). Referred by Mayr (1855: 411)
to Myrmica scabrinodis.
Myrmica rugulosa var. scabrinodorugulosa Nasonov, 1892: 9, Poland. Nomen nudum.
Myrmica (f. intrm.) sulcinodolobicornis Ruzsky, 1895: 31. Nomen nudum (attributed to Forel).
Myrmica balcanica var. testaceus Sadil, 1953: 207, Czechoslovakia. Nomen nudum.

3.7. Unavailable names in the genus Myrmica

Myrmica rugosa subsp. dshungarica var. brevispina Kuznetsov-ugamsky, 1927: 193, w, Kyrgyzstan.
Unavailable name, material referred to M. dshungarica by Tarbinsky 1976: 29; Radchenko
1994e: 73.
Myrmica rugulosa limanica nat. chersonensis Arnoldi, 1934: 164, w, m, Ukraine; Karawajew 1934: 76;
Weber 1948: 307. Unavailable name, material referred to M. limanica by Arnoldi 1970: 1840; to
M. gallienii by Radchenko 1994d: 76.
Myrmica rugosa subsp. dshungarica var. dentata Kuznetsov-ugamsky, 1927: 192, m, Kyrgyzstan.
Unavailable name, material referred to M. dshungarica by Radchenko 1994e: 73.
Myrmica scabrinodis subsp. lobicornis var. grandis Kuznetsov-ugamsky, 1927: 196, w Kyrgyzstan.
Unavailable name, material referred to M. lacustris by Radchenko 1994 e: 77 (see also Notes to
M. deplanala, above).
Myrmica rugosa subsp. dshungarica var. infuscata Kuznetsov-ugamsky, 1927: 192, w, q, Kyrgyzstan.
Unavailable name, material referred to M. dshungarica by Tarbinsky 1976: 29; Radchenko
1994e: 73.

663
Myrmica speciolosa Seifert, 1988: 14. Unavailable name.
Myrmica rugulosa limanica nat. strandi Arnoldi, 1934: 164, w, Ukraine; Karawajew 1936: 274, Weber
1948: 307. Unavailable name, material referred to M. limanica by Arnoldi 1970: 1840; to
M. gallienii by Radchenko 1994d: 76.
Myrmica rugosa subsp. dshungarica var. subacuta Kuznetsov-ugamsky, 1927: 192, m, Kyrgyzstan.
Unavailable name, material referred to M. dshungarica by Tarbinsky 1976: 29; Radchenko
1994e: 73.
Myrmica scabrinodis subsp. kasczenkoi var. subaltaica Ruzsky, 1925: 45, w, Russia. Unavailable name,
material referred to M. kasczenkoi by Radchenko 1994f: 82.

3.8. List of taxa originally described in the genus Myrmica, but afterward
transferred to other genera or excluded from Myrmica

Myrmica adelaidae Smith F., 1858: 128, w, Australia. Transferred to Podomyrma Smith F. by Dalla
Torre 1893: 60.
Myrmica agilis Smith F., 1857a: 71, w, Malaysia. Transferred to Pheidole Westwood by Donisthorpe
1932a: 449.
Myrmica algirica Lucas, 1849: 300, w, q, m, Algeria. Junior synonym of Crematogaster scutellaris
Olivier (misspelled asM. algerica): Emery 1891a: 14.
Myrmica aemula Heer, 1849: 161, q, Switzerland (Miocene). Incertae sedis in Myrmicinae (see also
Radchenko et al. 2007).
Myrmica amaurocyclia Foerster, 1850b: 486, w, Algeria. Junior synonym of Formica arenaria Fabri-
cius: Roger 1862: 292 (given as Atta arenaria; now -Messor Forel).
Myrmica angustula Nylander, 1856: 88, w, France. Transferred to Temnothorax Mayr by Bolton
2003:271.
Myrmica archaica Meunier, 1915: 212, m, Germany (Oligocene). Transferred to Aphaenogaster by
Radchenko et al. 2007: 1494.
Myrmica aspersa Smith F., 1865: 72, w, Indonesia. Transferred to Tetramorium by Bolton 1977: 86.
Myrmica atomaria Gerstäcker, 1859: 263, w, Mozambique. Junior synonym olMonomorium ominosa
(Gerstäcker): Roger 1863: 31.
Myrmica atratula Schenck, 1852: 91, q, m, Germany. Transferred to Anergates Forel by Forel 1874: 67.
Myrmica basalis Smith F., 1858: 125, w, Sri Lanka. Transferred to Monomorium by Mayr
1865: 92.
Myrmica beesoni Mukerjee, 1934: 7, w, India. Transferred to Tetramorium by Radchenko and Elmes
2001b: 239.
Myrmica bicolor Heer, 1867: 31, q (?), Yugoslavia (Miocene). Junior synonym of Myrmica tertiaria
Heer: Dalla Torre 1893: 117 (see also Mayr 1867b: 60). Incertae sedis in Myrmicinae (see also
Radchenko et al. 2007).
Myrmica bidens Foerster, 1850a: 50, w, q, m, Germany. Junior synonym of Myrmecina latreillei Cur-
tis: Mayr 1855: 421.
Myrmica bidentata Smith F., 1858: 124, w, India. Transferred to Monomorium by Radchenko and
Elmes 2001b: 238 (see also Mayr 1886: 359).
Myrmica bremii Heer, 1849: 161, q, Yugoslavia (Miocene). Incertae sedis in Myrmicinae (see also Rad-
chenko et al. 2007).
Myrmica caeca Jerdon, 1851: 116, w, India. Incertae sedis in Myrmicinae (see also Radchenko and
Elmes 2001b).

The original description of the single specimen of this species (Jerdon 1851: 116) fol-
lows: "Worker, length l / 5 t h of an inch [i.e. ca. 5 mm]; head somewhat oval, rather small;
no eyes; antennae short, thick, inserted near the mouth; and oblique groove on each side

664
of forehead [i.e. frons] for the laying of the antennae; jaws triangular, hooked at the tip, and
finely serrated; thorax narrow, of uniform width, granulose, with an elevation posteriorly
ending in two small spines; abdominal pedicles [i.e. petiole and postpetiole] raised, rounded,
pointed backwards, the first the highest; abdomen long, oval; head, thorax and legs reddish
brown; abdomen glossy brown".
For its time this was quite a complete description and it is obvious that "Myrmica" caeca
does not belong to the genus Myrmica. Moreover, if the original description is correct then
using the combination of the features described (e.g. lack of the eyes, presence of the
antennal scrobes, the short antennae, inserting very closely to the anterior head margin,
triangular [i.e. rather short than elongate] "hooked" and serrate mandibles etc.) it is im-
possible to place M. caeca to any known Myrmicinae genus, and in fact to any other of the
subfamilies having workers with a 2-segmented waist.

Myrmica carbonaria Smith F., 1858: 127, w, q, m, Madeira. Transferred to Monomorium by Roger
1863:31.
Myrmica carinata Smith F., 1859: 148, w, Indonesia. Transferred to Tetramorium by Bolton 1977:
86.
Myrmica cingulata Schenck, 1852: 104, w, q, m, Germany. Junior synonym oi Leptothorax nylanderi
(Foerster): Mayr 1855: 447 (now - Temnothorax).
Myrmica clypeata Mayr, 1853: 282, w, Austria. Transferred to Temnothorax by Bolton 2003: 271.
Myrmica concinna Heer, 1867: 32 (caste?), Yugoslavia (Miocene). Incertae sedis in Myrmicinae (see
. also Radchenko et al. 2007).
Myrmica consternes Walker, 1859: 374, w, m, Sri Lanka. Junior synonym of Monomorium minutum
(Jerdon): Donisthorpe 1932b: 576.
Myrmica contigua Smith F., 1858:125, q, Sri Lanka. Junior synonym oiMonomorium pharaonis (Lin-
naeus): Mayr 1886a: 359.
Myrmica corticalis Schenck, 1852: 100, w, q, m, Germany. Transferred to Temnothorax by Bolton
2003:271.
Myrmica debilis Foerster, 1850a: 52, m, Germany. Junior synonym of Stenamma westwoodi West-
wood: Mayr 1863: 454; revived from synonymy (as Stenamma debile): DuBois 1993: 314.
Myrmica diffusa Jerdon, 1851: 13, w, q, m, India. Transferred to Crematogaster by Roger 1863: 37.
Myrmica diligens Smith F., 1865: 73, w, New Guinea. Transferred to Tetramorium by Bolton 1977: 87.
Myrmica domestica Shuckard, 1838: 627, w, q, m, Great Britain. Junior synonym oi Formica pharao-
nis: Roger 1862: 295 (given as Myrmica pharaonis; now - Monomorium).
Myrmica duisburgi Mayr, 1868: 87, w, Baltic Amber. Transferred to Agroecomyrmex Wheeler by
Wheeler 1915: 57.
Myrmica flavidula Nylander, 1849: 43, w, Russia. Junior synonym oi Diplorhoptrum fugax (Latreille):
Mayr 1855: 450 (now - Solenopsis Westwood).
Myrmica fodicus Jerdon, 1851: 115, w, q, m, India. Junior synonym oi Myrmicaria carinata (Smith F.):
Emery 1892: 166.
Myrmica formosa Smith F., 1858: 128, w, Australia. Transferred to Podomyrma by Taylor and Brown
1985: 82.
Myrmica fragilis Smith F., 1858: 124, w, Singapore. Junior synonym oiMonomorium pharaonis: Mayr
1886a: 359.
Myrmica fuscipennis Smith F., 1861b: 46, q, Indonesia. Transferred to Pristomyrmex Mayr by Emery
1901:567.
Myrmica fuscula Nylander, 1846a: 935, w, q, m, Finland. Junior synonym olMyrmica caespitum: Smith
F. 1851; 118 (now - Tetramorium).
Myrmica galbula Losana, 1834: 326, w, Italy. Junior synonym oi Aphaenogaster barbarus (Linnaeus):
Roger 1863: 29 (now - Messor).

665
Myrmica glyciphila Smith F., 1858: 125, w, Sri Lanka. Transferred to Monomorium by Roger 1863: 32.
Myrmica gracilescens Smith F., 1860a: 64, w, Sulawesi. Transferred to Pheidole by Donisthorpe
1932a:-458.
Myrmica gracillima Smith F., 1861a: 34, w, Israel. Transferred to Monomorium by Mayr 1862: 753.
Myrmica hirtula Nylander, 1849: 45, w, Finland. Transferred to Harpagoxenus Forel by Emery
1922: 266.
Myrmica humilis Smith F., 1858: 123, q, India. Transferred to Cardiocondyla Emery: current transfer.

The main diagnostic features, given in the original description by F. Smith (1858,
p. 123) are: "Length 1.25 lines [i.e. ca 2.6 mm]... the first node of peduncle clavate, the
second much wider than the first and globose, the metathorax armed with two short acute
spines". The small size of this queen and distinctly widened postpetiole led us to transfer
Myrmica humilis to the genus Cardiocondyla rather than to any other genera distributed
in southern India (the species was described from Bombay).

Myrmica impura Foerster, 1850a: 48, w, Germany. Junior synonym of Tetramorium caespitum: Mayr
1855: 426; revived from synonymy (as Tetramorium impurum): Kutter 1977: 159.
Myrmica incerta Smith F., 1863: 21, q, Indonesia. Transferred to Pheidole by Donisthorpe 1932a: 471.
Myrmica insolens Smith F., 1861b: 47, q, Sulawesi. Transferred to Tetramorium by Emery 1901: 567.
Myrmica interrupta Schenck, 1852: 106, w, q, m, Germany. Transferred to Temnothorax by Bolton
2003:271.
Myrmica irritans Smith F., 1858: 127, w, South Africa. Transferred to Pheidole by Dixey and Longstaff
1907:331.
Myrmica jucunda Smith F., 1861: 34, w, Lebanon. Incertae sedis in Myrmicinae: current status.

The main diagnostic feature of this species, mentioned in the original description (Smith
F. 1858, p. 123) follows: "Length 1.5 line [i.e. ca2.5 mm]. Brownish-black, very smooth and
shining; ... the eyes small and placed midway at the sides of the head. Thorax narrowed
posteriorly, with the division between the meso- and metathorax strongly impressed; the
metathorax with two very minute spines; the first node of the abdomen, when viewed side-
ways, is wedge-shaped, the second globose. Abdomen subovate, widest towards the apex".
Based on the small size and "very smooth and shining" body we indisputably exclude
this species from the genus Myrmica, while the description given is insufficient to place
definitely Myrmica jucunda to any ant genus known from the Lebanon (it may be Tetra-
morium, Monomorium, Temnothorax, Pheidole etc).

Myrmica jurinei Heer, 1849: 163, q, Yugoslavia (Miocene) (see also Mayr 1867b: 59). Incertae sedis
in Myrmicinae (see also Radchenko et al. 2007).
Myrmica kirbii Sykes, 1835: 99, w, q, m, India. Transferred to Crematogaster by Smith F. 1858: 136.
Myrmica kollari Mayr, 1853: 283, w, q, m, Austria. Transferred to Tetramorium by Mayr 1855: 424.
Myrmica lacteipennis Zetterstedt, 1838: 452, m, Sweden. Junior synonym of Myrmica acervorum
(Fabricius): Nylander 1846a: 936 (now - Leptothorax Mayr).
Myrmica laevigata Smith F., 1855: 130, w, Great Britain. Junior synonym of Pheidole megacephala
(Fabricius): Roger 1863: 30.
Myrmica laevissima Smith F., 1860b: 108, w, Indonesia. Junior synonym of Solenopsis geminata
(Fabricius): Donisthorpe 1932a: 463.
Myrmica laeviuscula Foerster, 1850a: 54, q, Germany. Junior synonym of Formicoxenus nitidulus:
Mayr 1855: 418.
Myrmica leonina Losana, 1834: 332, w, Italy. Junior synonym of Myrmica rubida (Latreille): Roger
1859: 252 (now - Manica Jurine).

666
Myrmica lippula Nylander, 1849: 41, w, Russia. Junior synonym of Stenamma westwoodi: Forel 1889:
258 (see also Radchenko 2007a).
Myrmica longiceps Smith F., 1858:128, w, Australia. Transferred to Aphaenogaster by Mayr 1876: 98.
Myrmica longipes Smith F., 1857a: 70, w, Singapore. Transferred to Pheidole by Bolton 1995: 324.
Myrmica macrocephala Heer, 1849: 158, q, Switzerland (Miocene). Incertae sedis in Myrmicinae (see
also Radchenko et al. 2007; Dlussky and Rasnitsyn 2007).
Myrmica maligna Smith F., 1865: 72, q, Indonesia. Transferred to Podomyrma by Donisthorpe
1932a: 473.
Myrmica mellea Smith F., 1859: 148, w, Indonesia. Junior synonym of Solenopsis geminata:
Donisthorpe 1932a: 455.
Myrmica minkii Foerster, 1850a: 63, w, Germany. Junior synonym of Asemorhoptrum lippulum (Ny-
lander): Mayr .1863: 395 (now - Stenamma) (see also Radchenko 2007a).
Myrmica modesta Foerster, 1850a: 49, w, Germany. Junior synonym of Myrmica caespitum: Curtis
1854:215 (now-Tetramorium).
Myrmica modesta Smith F., 1860b: 108, w, Indonesia. Junior synonym of Tetramorium bicarinatum
(Fabricius): Bolton, 1977: 94.
Myrmica moesta Smith F., 1863: 21, w, Indonesia. Transferred to Vollenhovia Mayr by Donisthorpe
1932a: 471.
Myrmica molassica Heer, 1849: 162, q, Switzerland (Miocene). Incertae sedis in Myrmicinae (see also
Radchenko et al. 2007).
Myrmica montana Labram and Imhoff, 1838, w, q, m, Switzerland. Junior synonym oiMyrmica rubida:
Mayr 1855: 400 (now -Manica).
Myrmica muscorum Nylander, 1846b: 1054, w, q, m, Finland. Transferred to Leptothorax by Mayr
1855:439.
Myrmica mutica Nylander, 1849: 39, w, q, m, Russia. Transferred to Messor by Ruzsky 1903: 315.
Myrmica nebulosa Novak 0., 1878: 91, m, Germany (Oligocene). Incertae sedis in Myrmicinae (see also
Radchenko et al. 2007).
Myrmica nitidula Nylander, 1846b: 1058, w, Finland. Transferred to Formicoxenus Mayr by Mayr
1855:418.
Myrmica nylanderi Foerster, 1850a: 53, Germany. Transferred to Temnothorax by Bolton 2003: 271.
Myrmica oblonga Smith F., 1860b: 107, w, Indonesia. Transferred to Vollenhovia by Emery.1897: 560.
Myrmica obscurata Motschoulsky, 1863: 16, w, Sri Lanka. Incertae sedis in Myrmicinae: current
status.

Motschoulsky (1863, p. 16) described Myrmica obscurata from the Mts. Nuta-Ellia near
Colombo (Sri Lanka) and noted that this species is similar to M. pilinodis: "Forma et sculp-
tura ut in M. pilinodi [Sic\]", but differ from the latter mainly by the colour and by the
somewhat bigger size (1 3 / 5 1. [i.e. ca. 3.4 mm])". We can not place Myrmica obscurata to
any genus, but are sure that it does not belong to Myrmica (see also Notes to M. pilinodis,
below).

Myrmica obsoleta Heer, 1849: 160, q, Switzerland (Miocene). Incertae sedis in Myrmicinae (see also
Radchenko et al. 2007).
Myrmica ominosa Gerstäcker, 1859:263, w, Mozambique. Transferred to Monomorium by Roger 1863:31.
Myrmica opaca Smith F., 1861b: 47, w, Sulawesi. Transferred to Rombonella Wheeler by Bolton
1976: 294.
Myrmica pallida Nylander, 1849: 42, w, Italy. Transferred to Aphaenogaster byRoger 1863: 29.
Myrmica pallidula Nylander, 1849: 42, w, Italy. Transferred to Pheidole by Smith F. 1858: 173.
Myrmica parallela Smith F., 1859: 147, w, Indonesia. Transferred to Tetramorium by Donisthorpe
1932a: 455.
Myrmica parvula Schenck, 1852: 103, w, Germany. Transferred to Temnothorax by Bolton 2003: 271.

667
Myrmica pedestris Smith F., 1861b: 46, q, Sulawesi. Transferred to Vollenhovia by Mayr 1867a: 94.
Myrmica pellucida Smith F., 1857a: 71, w, Singapore. Junior synonym of Tapinoma melanocephalum
(Fabricius): Mayr 1886a: 359.
Myrmica pertinax Smith F., 1861b: 46, w, Sulawesi. Transferred to Vollenhovia by Donisthorpe
1932a: 468.
Myrmica pilinodis Motschoulsky, 1863: 16, w, Sri Lanka (misspelled as pallidinodis or pallinodis:
Dalla Torre 1893: 113; Emery 1922: 356). Incertae sedis in Myrmicinae: current status.

This species was described based on workers from the Mts. Nuta-Ellia near
Colombo (Sri Lanka). Several features of Myrmica pilinodis , particularly its small size ( 1 1 / 5
1. [i.e. ca 2.5 mm]) and strongly transversed 2-7 t h funicular joints led us to exclude this
species from the genus Myrmica. However, based on the original description it is impossi-
ble to place it to any Myrmicinae genera.

Myrmica polita Smith F., 1860b: 108, w, Indonesia. Junior synonym of Pheidologeton diversus
(Jerdon): Donisthorpe 1932a: 463.
Myrmica poneroides Smith F., 1860b: 197, q, Indonesia. Transferred to Vollenhovia by Mayr
1886a: 361.
Myrmica punctata Smith F., 1860b: 10, w, Indonesia. Transferred to Pristomyrmex by Mayr
1886a: 362.
Myrmica punica Smith F., 1861a: 34, w. Israel. Transferred to Tetramorium by Mayr 1870: 975.
Myrmica pusilla Heer, 1849: 164, q, Yugoslavia (Miocene). Transferred to Crematogaster by Han-
dlirsch 1907: 873.
Myrmica quadrispinosa Smith F., 1865: 72, w, Indonesia. Transferred to Pheidole by Mayr 1879: 677.
Myrmica recedens Nylander, 1856: 94, w, France, Italy, Spain. Transferred to Temnothorax by Mayr
1861: 68 (see also Bolton 2003: 271).
Myrmica rediana Gené, 1841: 399, w, Italy. Junior synonym of Crematogaster scutellaris: Mayr
1855: 469.
Myrmica rubriceps Nylander, 1849: 44, w, Sicily. Junior synonym of Crematogaster scutellaris: Mayr
1855:469.
Myrmica rufa Jerdon, 1851: 114, w, India. Transferred to Crematogaster by Roger 1863: 37.
Myrmica ruficeps Smith F., 1861b: 46, q, Sulawesi. Transferred to Pheidole by Mayr 1867a: 96.
Myrmica rufitarsis Foerster, 1850b: 485, w, Algeria. Junior synonym of Aphaenogaster barbarus: Roger
1863:29 (now-Messor).
Myrmica rufotestacea Foerster, 1850b: 489, w, Algeria. Transferred to Messor by Emery 1908b: 439.
Myrmica rugiceps Heer, 1849: 160, Switzerland (Miocene). Transferred to Pheidolegeton Mayr by Ass-
mann 1870: 40.
Myrmica rugifrons Smith F., 1858: 124, q, India. Transferred to Monomorium: current transfer
(provisionai).

F. Smith (1858, p. 123) originally placed Myrmica rugifrons to the "Subdivision 7 (genus
Monomorium [Sicì] Mayr)". Based on the some important features, given in the original de-
scription, such as palp formula 2,2, body length 3 lines [i.e. 6.3 mm - queen described],
and "the hinder margin of vertex emarginated... metathorax concavely truncate at the apex,
a short spine on each side at the verge of the truncation" (loc. cit., p. 124), we provision-
ally place "Myrmica rugifrons" to the genus Monomorium.

Myrmica scabrosa Smith F., 1859: 147, w, Indonesia. Transferred to Tetramorium by Donisthorpe
1932a: 455.
Myrmica scalpturata Nylander, 1856: 83, w, Algeria. Junior synonym of Alta arenaria: Roger 1862: 292
(now - Messor).

668
Myrmica semipolita Nylander, 1856: 86. Italy. Transferred to Aphaenogaster by Roger 1863: 29.
Myrmica simillima Smith F., 1851: 118, w, Great Britain. Transferred to Tetramorium by Mayr
1861:61.
Myrmica simpliciuscula Nylander, 1856: 92, w, France. Junior synonym of Leptothorax interruptus
(Schenck): Roger 1859: 258 (now - Temnothorax)
Myrmica sordidula Nylander, 1849: 44, Sicily. Transferred to Crematogaster by Mayr 1855: 472.
Myrmica striatula Nylander, 1849: 40, Russia. Junior synonym of Myrmecina latreillei: Mayr
1855:421.
Myrmica sublaevis Nylander, 1849: 33, w, Finland. Transferred to Harpagoxenus by Forel 1893: 167.
Myrmica subopaca Smith F., 1858: 127, w, q, Madeira. Transferred to Monomorium by Mayr
1862:753.
Myrmica suspiciosa Smith F., 1859: 148, w, Indonesia. Junior synonym of Pheidole megacephala:
Donisthorpe 1932a: 455.
Myrmica tarda Jerdon, 1851: 115, w, India. Junior synonym of Meranoplus bicolor (Guérin-Méneville) :
Emery 1892: 166.
Myrmica terminalis Shuckard, 1838: 627, w, Great Britain. Transferred to Crematogaster by Dalla
Torre 1893: 87.
Myrmica tertiaria Heer, 1849: 159, q, Yugoslavia (Miocene) (see also Mayr 1867b: 59). Incertae sedis
in Myrmicinae (see also Radchenko et al. 2007).
Myrmica testaceopilosa Lucas, 1849: 301, w, Algeria. Transferred to Aphaenogaster by Roger 1863: 29.
Myrmica thoracica Smith, 1859: 149, w, Indonesia. Transferred to Cardiocondyla by Donisthorpe
1932a: 455.
Myrmica tigreensis Guérin-Méneville, 1849: 351, m, Ethiopia. Transferred to Myrmicaria Saunders by
Smith F. 1858: 141.
Myrmica trachylissa Smith F., 1858: 126, q, Borneo. Transferred to Pristomyrmex by Mayr
1886a: 359.
Myrmica trinodis Losana, 1834: 327, w, Italy. Junior synonym of Pheidole megacephala: Roger
1863: 30.
Myrmica umbripennis Smith F., 1863: 21, q, Indonesia. Transferred to Pristomyrmex by Donisthorpe
1932a: 471.
Myrmica vastator Smith F., 1857a: 71, w, Singapore. Transferred to Monomorium by Mayr
1886a: 359.
Myrmica venusta Smith F., 1858: 126, w, Syria. Transferred to Monomorium by Roger 1863: 32.
Myrmica vexator Smith F., 1861b: 47, w, Indonesia. Junior synonym of Monomorium destructor:
Donisthorpe 1932a: 468.

669
г

CHAPTER 4. ZOOGEOGRAPHY AND EVOLUTION OE THE GENUS MYRMICA

Once world-wide exploration became commonplace for 19th century natural scientists
they soon recognized that quite different groups of plant and animal species lived in
different parts of the world, therefore they divided the world into geographical areas that
reflected these distributions. Arguably Alfred Russell Wallace (1823-1913) was the most
influential, he established six major zoogeographical regions each with various sub-regions
(Wallace 1876); these became the basis for subsequent studies and are still largely
recognized today. World-wide, species of the genus Myrmica are found only in Wallace's
Nearctic, Neotropical, Palaearctic and Oriental Regions that comprise most of the Northern
Hemisphere (Map 1 in Chapter 2). As this monograph is concerned only with the Old World
Myrmica species, only the last two regions are relevant. Based both on the abundance of
species and variety of species groups, the Palaearctic Region is the main centre of diversity
for the genus Myrmica. Even so, a considerable number of species inhabit the Oriental
Region (India, Taiwan, southern China, Vietnam, Burma, Thailand), where they live ex­
clusively in mountain forests and alpine meadows at altitudes over 1200-2000 m and some­
times as high as 4800 m a.s.l. Not long ago, Myrmica was considered to be strictly
a Northern Hemisphere genus until the rediscovery of a specimen in Emery's collection that
was collected south of the Equator near south-eastern Borneo, an island that Wallace
included in his Oriental Region (see notes on M. emeryi in Chapter 3.3 and below).
Ecologically speaking, Myrmica species can be considered as meso- or even hygrophilous
ants, restricted to temperate habitats (see Chapter 1.3). They have evolved life-styles that
enable them to survive cold seasonal environments but physiologically they can not cope
with high temperatures, individuals exposed to air temperatures > 26°C soon die.
Obviously soil moisture and soil temperatures interact so that as a rule of thumb in hotter
habitats one is more likely to find Myrmica nests living in humid habitats near water. They
are completely absent in true desert and semi-desert habitats, although a few species can be
found within the Desert Zone, where they are restricted to intrazonal, wet and often halo-
phytous sites. In the Oriental Region, which mostly has a moist subtropical and tropical
climate, the distribution of Myrmica is usually restricted by temperature to cooler mountain
habitats. In the Palaearctic Myrmica distribution is mainly limited by habitat temperature
in the north (too cool) and by a combination of temperature and humidity in hot southern
regions. Thus while a few Myrmica species have been recorded from the humid mountains
of North-West Africa (Morocco, Algeria and Tunisia) there are no records from the drier
southern Mediterranean countries (Libya, Egypt), the Arabian Peninsula and the Middle
East (i.e. Israel, Jordan, Lebanon, Syria and Iraq). Several species have been recorded from
the northern mountains of Iran, Afghanistan and Pakistan, but the southern limit of the
distribution in this region is poorly understood.
We recognise nine centres of diversity for Myrmica within the Palaearctic and
Oriental Regions, in most cases these are well established zoogeographical centres of di­
versity for many other animal and plant species. They are: 1. the Central Asian mountains

671
(abbreviated to C-Asia); 2. South and South-East Asian tropical and subtropical moun-
tains, including south-western China (SE-Asia); 3. the mountains of the Tibetan Plateau
(Tib); 4. the Middle Asian mountains (M-Asia); 5. Siberia including Mongolia (Sib); 6. the
far East of Russia, north-east of China and Japan (Far-E); 7. the Forest Zone of Europe and
West Siberia (Eur-Sib); 8. the Mediterranean zone (Med) and 9. the western part of the
Steppe Zone (W-step) (Map 144). For each centre we recorded the presence/absence of
each of the 142 "good" Myrmica species recognized here (Table 4.1) and similarly - the
number of species in each species group (Table 4.2). The C-Asian and SE-Asian faunas are
the most diverse, both in terms of number of species and species groups, also they support
more of the oldest lineages within the genus (see Chapter 2 and below).

Map 144. CentresofMyrmica diversity. 1 - Central Asian mountain; 2 - South and South-East Asian;
3 - Tibetan; 4 - Middle Asian mountain; 5 - Siberian-Mongolian; 6 - Far Eastern; 7 - Euro-West-
Siberian; 8 - Mediterranean; 9 - West Steppe.

4.1. The centres of Myrmica species diversity

(1) CENTRAL ASIAN MOUNTAIN

We define the Central-Asian centre as the mountain ranges of Hindukush, Karakorum,
and the south-western macro-slope of the Himalaya that includes the north-eastern
territories of Afghanistan, Pakistan and India together with Nepal and Bhutan (i.e. the
southern and western mountains that border the Tibetan Plateau). Interestingly, this more

672
or less coincides with the northern part of Wallace's (1876) Indian subdivision of his
Oriental Region.
About a quarter of the Old World Myrmica fauna are found in this centre, comprising
33 species from seven species groups (see Tables 4.1 and 4.2). Thirty one species (94%) are
endemic to the centre; the two exceptions are M. tenuispina that has probably penetrated
north-eastern Afghanistan from the Middle Asian Mountains, and M. kozlovi (that lives at
the highest known altitude for all ants - up to 4800 m a.s.l.) that has probably crossed the
Himalaya from Tibet (see detailed notes in Chapter 3.3). Furthermore, the majority (7) of
the ten Old World species that have such unusual combination of features that we cannot
place them to any of our species groups are found in this region. Apart from these unusual
species, only the rugosa-group appears to be endemic to this centre while the remaining six
species groups are also represented in other eastern centres (see Chapter 4.2.1).
The Central Asian centre of diversity is interesting because its species (and species
groups) share a feature that we consider represents the plesiomorphic state for Myrmica -
the female castes have an antennal scape that is only slightly curved at the base, not angu-
lar and with no trace of a lobe or carina. In addition, the majority of species, for which the
morphology of the males is known, have.males with a short antennal scape, another feature
that we consider as basal for this genus. Furthermore, many of the species and species
groups have morphological characters that are never or very rarely found in the Myrmica
species of more northerly centres of diversity, these features include transverse rugosity of
the alitrunk dorsum {pachei- and inezae-groups), punctato-striated first gastral tergite
(M. brancuccii), smooth and shiny surfaces of the alitrunk and waist (M. nitida), and mul-
tidentate mandibles with more than 10 denticles (M. hecate) that is a feature characteris-
tic of Manica species. Also a third of the 21. known species of the ritae-group are endemic
to this region, including the boltoni species complex. We believe the ritae-group is the least
evolved morphologically of modern Myrmica species groups because several of the extinct
Myrmica species are very similar to extant species from this group (see Radchenko et al.
2007),
Taken together, we suggest that these features show that the Myrmica fauna of this
centre of diversity probably has remained fairly isolated from that of the other centres for
a very long time, perhaps since the Himalayan uplift > 25 Ma. For most of this time the high
Himalayan ranges to the north and east have formed an almost insuperable barrier for most
ants, certainly Myrmica species, while further south tropical and sub-tropical conditions
limited opportunities for interchange with other faunas. Such long-term isolation has led to
a mixture of relicts and the evolution of many derived forms which are distinctive from
almost all other Myrmica species, both Palaearctic and Nearctic. We are sure that many
more locally endemic species remain to be discovered when these territories are explored
more fully.

(2) SOUTH- AND SOUTH-EAST ASIAN TROPICAL AND SUBTROPICAL CENTRE

We consider this centre to include territories of southern and south-western China, Tai-
wan, and the northern mountains of Vietnam (and probably Laos), Burma and Thailand and
perhaps the whole of SE Asia and even Indonesia (based on the rediscovery of M. emeryi
from near Borneo - see below). Again it is interesting that this centre more or less coincides
with one of Wallace's (1876) subdivisions, the Indochinese subdivision of the Oriental
Region. In this centre all Myrmica species live in mountain forests and meadows, usual-
ly at altitudes between 1800 and 2500 m a.s.l. being restricted by temperature at lower

673
altitudes (see above and Chapter 1.3). In the north of this centre, some species, such as
M. angulata, live at lower altitudes, in China it was recorded as low as 500 m although in
Vietnam it lives in the forests at altitudes above 1500 m; on the other hand, some Myrmica
species live at very high altitudes, e.g. in Yunnan several were collected at altitudes up to
4700 m a.s.l.
A total of 30 Myrmica species (including M. emeryi) have been recorded from this cen­
tre of diversity and all except M. curiosa (which has an uncertain taxonomic position) can
be assigned to one of six species groups. It is interesting that while each of these species
groups has members in at least one other of the centres of diversity, all species are endemics
to this centre (see Tables 4.1 and 4.2). The fauna is dominated by members of the ritae- and
раchei-groups (14 and 11 species respectively), which we consider to be quite old lineages
morphologically (see also Jansen et al. 2010). Four other groups are represented by one
species each: M. arisana (rubra-group) and M. mirabilis (luteola-group) are found only on
the Island of Taiwan, while M. mixta (inezae-group) and M sinoschencki (schencki-group)
were recorded from the north of this centre, in the mountains of the Sichuan province of
China.
It seems probable to us that this fauna has been isolated for at least as long as that of
the Central Asian centre. This region of southern China and South-East Asia has had
a relatively stable, warm climate for a long time (compared to other centres) so that
probably even in the late Eocene and Oligocene it had a Myrmica fauna that was restricted
to the cooler habitats on high mountain systems. Possibly the ancient fauna was less
diverse than that isolated in the more northern C-Asian centre, comprising mainly species
from the pachei- and ritae species groups. During subsequent climatic changes species were
sometimes restricted to the tops of high mountains while at others times could spread be­
tween mountains, at least different mountains in the same mountain chain, with the much
warmer lowlands always restricting exchange between populations. Thus we suggest that
this centre of diversity represents a huge refuge for some of the ancient lineages of Myr­
mica, which have evolved into many different locally endemic species. The fact that within
this centre, species that belong to the same species group are much more "distinct" mor­
phologically and easier to separate from one another compared to most species in Euro­
pean or Siberian species groups, we believe supports this argument.
We included M. emeryi in this fauna despite the fact that it was apparently taken in the
Island Pulo Laut, SE of Borneo, now part of Indonesia, which accords with the label on the
type specimen (see notes to M. emeryi in Chapter 3.3). While Wallace (1876) included Bor­
neo in his Oriental Region it was part of a distinct Indo-Maiayan sub-region which com­
prised most of Indonesia, Borneo Island and the Philippines. This Oriental Region is clearly
demarcated from Sulawesi and islands to the east by the famous Wallace line still recognized
today, but the separation between the Indochinese and Indo-Malayan subdivision is less
clear. At various times the Islands of Borneo and Sumatra have been linked to the In­
dochinese region, perhaps most recently only 15,000 years ago, so it is not unreasonable that
a few Myrmica species might persist on mountains in Borneo and elsewhere in Wallace's
Indo-Malayan region, perhaps even in the Philippines. Finally we emphasize that many of
the places where Myrmica might live within our huge SE-Asian centre of diversity are
remote and difficult to access, even more so if we include the Indo-Malayan sub-region.
A decade or so ago only seven Myrmica species were known from this centre and these
were considered "rare curiosities", but during the last 10 years colleagues have been able
to collect ants from a few locations within this vast area and made the Myrmica specimens

674
available for our study. Consequently we have added another 23 species and know enough
to predict that the SE-Asian centre probably supports the largest and most diverse regional
Myrmica fauna in the World. We can not imagine how many species will eventually be de-
scribed from this region, but we confidently expect many more remain to be discovered
when other remote mountain sites are studied.

(3) TIBETAN MOUNTAIN CENTRE

We include the Tibetan Plateau and some of its surrounding mountain systems in this
centre. At first, we were not sure whether to treat this region as a separate centre because
up to the present time, only four species belonging to the kurokii-, smythiesii- and tibetana
species groups are known from Tibet (see Tables 4.1 and 4.2), but we decided to do so be-
cause the centre appears to be so strongly isolated from the rest of the Palaearctic Region.
Three of the four Myrmica species are endemic to Tibet. Only M. kozlovi crosses the
Himalaya barrier, occurring also in the C-Asian centre, and we think it most likely that this
altitude tolerant-species (see Chapter 3.3) originated from Tibet. In terms of species groups,
the Tibetan fauna is most related to that of the adjacent C-Asian and M-Asian centres.
There has been little study of the Myrmica fauna, indeed the ant fauna generally, of this
huge area since the Myrmica species were described by Mayr in 1889 and by Ruzsky in
1915, and since that time almost no additional material has been obtained. We are sure that
many more than just four Myrmica species live in this centre of diversity and expect a lot
of forms will be described in future, otherwise the status of this proposed centre would need
be revised.

(4) MIDDLE ASIAN MOUNTAIN CENTRE

It includes the Tien-Shan and Pamiro-Alai mountain systems. This and the remaining
proposed centres of Myrmica diversity within the Palaearctic Region are less isolated from
each other than the three centres discussed above. Fourteen species from five species groups
have been recorded in M-Asia, the six non-endemic species are widespread boreal or steppe
elements (e.g. M. rubra, M. sulcinodis, M. bergi, M. deplanata, M. schencki) that probably
have penetrated from the north and west. Only the dshungarica species group is endemie
to M-Asia while just over half the species (eight) are endemics (see Tables 4.1 and 4.2).
However, unlike the C-Asian and SE-Asian centres most of the endemie species are
not highly specialized and distinct morphologically; for example, M. tobiasi belongs to the
speczioides-complex of the scabrinodis species group and resembles several species from
Kazakhstan and Europe, while M. kirghisorum and M. saposhnikovi are members of the
lobicornis-group and are similar to some European and Siberian species. Even species of the
endemie dshungarica-group are similar morphologically to the rubra-group. Only M. tenui-
spina and M. orthostyla (tibetana-group) have a more or less unique feature, the shape of
the frontal carinae and lobes. We conclude that this centre might have been isolated for
long periods in the past, developing several distinctive species, but at others times it has
been less isolated from adjacent regions to the north and west so that the influence of the
boreal and steppe fauna is well visible.

(5) SIBERIAN-MONGOLIAN CENTRE

We restrict this to the north-eastern part of the old Siberian sub-region of the Palaearc-
tic as used by Wallace and some other authors; it includes East and South Siberia (east of

675
the Enisei River and the Altai Mts.), Mongolia and the northern part of the Russian Far
East. The dominant biomes of this vast region are various kinds of taiga forests with steppe
habitats in the south. It should be remembered that these Siberian steppes are in the soil
permafrost zone and probably are somewhat similar to the so-called tundra-steppes that
existed in Pleistocene Europe; they are very different, both floristically and faunistically,
from the modern European and Kazakhstan steppes (Lavrenko 1954; Lavrenko et al. 1991).
Not surprisingly, steppe Siberian Myrmica species (e.g. M. kasczenkoi, M. pisarskii,
M. koreana, etc.) do not occur in the European steppes, and similarly no European steppe
species live in East Siberia.
Five species groups containing 19 species are found in this centre, the lobicornis species
group is clearly the dominant one containing about 60% (11) of the recorded species. At the
present time, the more northern, taiga regions of this centre are not effectively isolated from
the European and West Siberian taiga fauna. Consequently, they support several trans-
palaearctic boreal species: M. rubra and M. ruginodis from the rubra-group, and
M. lobicornis and M. sulcinodis from the lobicornis-group. The ranges of two other species
of the lobicornis-group (M. angulinodis and M. transsibirica) extends west beyond the Eni-
sei River into the eastern part of West Siberia. Otherwise, the Myrmica fauna of the Sib
centre well differs from the Euro-Sib centre (Tables 4.1 and 4.2). It contains the endemic
(arnoldii) species group and seven endemic species from the lobicornis-group plus two
other endemics, M. divergens (scabrinodis-group) and M. tschekanovskii (uncertain taxo-
nomic position).
It is possible to explain this mixture of endemic and transpalaearctic elements by the his-
tory and development of the taiga biome. Taiga forests arose in the north-east of Asia at the
end of the Miocene and beginning of the Pliocene, and by the end of the Pliocene it extended
into North Europe (Vulf 1944; Lavrenko 1946; Baranov 1959). The subsequent Pleistocene
glaciations almost completely demolished the biota of the taiga and partly destroyed that of
the mixed and deciduous forests, especially in Europe. The recolonization of modern
European taiga started after the last glaciations most probably by way of migration from
east to west, from Siberia. We suggest that the much higher richness of the boreal ant fauna
in general and the genus Myrmica in particular, in the east of the Palaearctic Region sup-
ports this idea. For example, the lobicornis- and rubra-groups dominate in boreal condi-
tions of the Palaearctic, yet only seven from a total of 26 species occur in Europe.

(6) FAR EASTERN CENTRE

In this we include the southern part of the Primorsky Krai of Russia, the north-east of
China, the Korean Peninsula and Japan. This area was an important refuge during the Pleis-
tocene epoch where ancient Tertiary flora and fauna persisted. In general, the fauna, in-
cluding the ant fauna, of our Far-E centre contains many Oriental (tropical) elements which
led Wallace and later authors to group it together with south-western China as the
Manchurian sub-region of the Palaearctic. However, none of the Myrmica species charac-
teristic of our SE-Asian centre are present in this Far-E centre, which has led us to treat it
separately (see also Radchenko 2005, 2007b).
As in the Siberian-Mongolian centre, species from the lobicornis group dominate: more
than half (8 of 14) Myrmica species recorded here belong to that group. However, the Myr-
mica fauna of this centre appears to be much more isolated than that of the Siberian cen-
tre and has much fewer similarities to that of the Western Palaearctic (Tables 4.1 and 4.2).
Seven species (half the total) are endemic and only two of the boreal transpalaearctic species

676
are found here -M. ruginodis and M. sulcinodis, but the latter has not reached Japan. More-
over, many species have quite unusual morphological features that are only found in species
groups or species complexes endemic in East Asia (e.g. M. luteola, M. kurokii, M. ademonia
and M. excelsa). It seems probable to us that much of the Myrmica fauna of this centre has
quite ancient roots, but has been strongly influenced in more recent times by the invasions
of boreal elements from the north.

(7) EURO-WEST-SIBERIAN FOREST CENTRE

This is another vast region that includes most of the European sub-region of the
Palaearctic as originally defined by Wallace, comprising Central and Northern Europe, the
Forest and Forest-Steppe Zones of East Europe and West Siberia up to and including the
Altai Mts. The predominant vegetation types of this region are various types of coniferous,
mixed and deciduous forests and various types of grassland and meadows, many of them
created or modified by man's activities. This centre of diversity supports 22 Myrmica species
belonging to six species groups. Indisputable the scabrinodis-group dominates this centre,
containing more than half (12) of the species, the other species groups containing only
1-3 species each. Despite the fact that there are few geographical barriers to isolate this
centre from the adjacent Mediterranean, West-Steppe and Siberian-Mongolian centres, one
endemic species group (the myrmicoxena-group) and seven endemic species occur here.
It is interesting to note however, that the myrmicoxena-group and four of the seven
endemic species are social parasites (Tables 4.1 and 4.2). Many of the species common
with other centres are widespread, often boreal species (e.g. M. rubra, M. ruginodis,
M. lobicornis, M. sulcinodis, M. schencki).

(8) MEDITERRANEAN CENTRE

This largely coincides with the traditional view of the Mediterranean subregion of the
Palaearctic. We include here the Iberian Peninsula, North-West Africa, southern France, the
Apennine Peninsula (south of Alps), Dalmatia, Balkans, Greece, Asia Minor, Transcauca-
sus, Iran and the Kopet-Dagh Mts. As mentioned at the start of this chapter, there are very
few Myrmica species present south of the Mediterranean Sea. The common factor that
characterises this centre is the so-called Mediterranean climate of hot dry summers, rela-
tively long mild spring and autumn periods and a quite short colder winter period, with
most precipitation occurring in the winter months. It is the most geographically fragmented
of our centres of diversity and can be subdivided to several smaller parts, e.g. Iberian, Apen-
nine, Balkans, and Asian (Asia Minor, Transcaucasus, Iran and Kopet-Dagh Mts.) that are
isolated from each other by mountains or seas and consequently have diversified floristically
and faunistically. Each of these sub-centres is geographically more connected with the
centres to the north than to each other.
It was not surprising therefore, to find that the Myrmica fauna of this centre is one of
the richest with 32 species from six species groups. It is more diverse than the SE-Asian
centre and is surpassed only by the Himalayan fauna (C-Asian centre). As in the Euro-
Siberian centre, species from the scabrinodis-group dominate (13 species, more than 40%
of the fauna). The fragmented' nature of the centre no doubt contributes to it having the
largest percentage of endemics (66% or 21 species), including one socially parasitic species
group (M. laurae). Seven of the endemic Myrmica species live in the Iberian Peninsula
and in North Africa, four in the Apennine Peninsula, one in Greece, six in Asia Minor and
Transcaucasus, and a further two in the Balkans, Crimea and Turkey.

677
 Despite the high level of endemism, the Myrmica fauna of the Mediterranean centre has
considerable similarities with that of the adjacent Euro-Siberian and Western-Steppe cen-
tres (Table 4.3). As the centre is largely isolated by climate rather than geography, it is not
surprising that many boreal, transpalaearctic and European forest species are present in the
northern mountains of the centre. Indeed many of them may have found refuge in the
various sub-centres during ice periods. We feel that there is still much work needed to fully
understand the Myrmica fauna of this centre and we confidently expect many more locally
endemic species of Myrmica to be described in future.

(9) WEST-STEPPE CENTRE

We consider this centre to include the Steppe Zone from Romania to Kazakhstan and
the Middle Asian deserts. This somewhat specialized region is characterised by the vegeta-
tion and climate which tends to be hot and dry. As Myrmica ants are not well adapted to
such conditions, it is rather surprising that the fauna is as rich as it is: 16 species from four
species groups are found here. Even though the level of endemism in the steppe Myrmica
fauna is the lowest among the nine centres of diversity, there are nevertheless four endemics,
living in intrazonal, wet and usually halophytous habitats (Table 4.1).
As in the Euro-West-Siberian and Mediterranean centres, the scabrinodis-group domi-
nates the West-Steppe centre (9 species, more than half the total). Apart from three boreal
species (M. rubra, M. ruginodis and M. lobicornis), three species belong to the schencki-
group (with even one endemic, M. inucta, from Kazakhstan). In general, this centre seems
to comprise the least specific Myrmica fauna compared to that of the eight other centres
(Table 4.3).

4.2. Comparison of the regional faunas of Myrmica

4.2.1. Diversity between the nine centres

We used Jaccard's coefficient (JC) to compare the Myrmica fauna of our nine centres of
diversity, this simply measures similarity by placing emphasis on the proportion of species
(or groups) held in common between centres. First, we compare centres using the co-
occurrence of the species groups (Table 4.3); we ignore the three mono-specific socially-
parasitic groups and the 10 species with uncertain taxonomic position (these could also be
considered as mono-specific species groups). Considering these 13 species as mono-
specific groups and including them into the analysis of groups produced exactly the same
pattern as that outlined below but reduced the JC values between groups (this is because
each of these mono-specific groups occur only in a single centre of Myrmica diversity).
It is immediately obvious from Table 4.3 that the C-Asian, Tibetan and SE-Asian centres
have low similarity to each other or to any of the other centres (for convenience we refer to
these three as the "oriental centres" although strictly some are not entirely in.the Oriental
Region). Based on species groups, the remaining six centres (M-Asian, Far-Eastern, Euro-
West Siberian, Siberian-Mongolian, Mediterranean and the Western Steppe) have average
of JC's = 0.60 (for convenience we refer to these as the "six northern centres").
Of the oriental centres, the Tibetan centre has low similarity to all others (Table 4.3) but
confidence in that result can not be high because the fauna ofthat region is under-recorded
compared to adjacent regions. The better studied C-Asian (Himalayan) centre is very

678
different from all others: the rugosa-group (nine species) is unique to this centre as
are seven of the 10 species with uncertain taxonomic position. It shares the ritae-, pachei-
and mezae-groups with the SE-Asian centre, species in these groups are not found
elsewhere/Despite having many unusual species, the values of JC between the SE-Asian

Table 4.1. Distribution of the Myrmica species in the species diversity centres. Abbreviations:
C-Asia = Central Asian mountains; SE-Asia = South- and South-East Asian tropical
and subtropical mountains; Tib = Tibetan plateau; M-Asia = Middle Asian mountains;
Sib = Siberian and Mongolian mountains and forests; Far-E = Far Eastern Palaearctic; Eur-Sib
= European and West-Siberian forest zone; Med = Mediterranean; W-step = Western Steppe.

!
Species diversity centres
Species
C-Asia SE-Asia Tib M-Asia Sib .Far-E Eur-Sib Med W-step
1 ademonia +
2 afghanica +
3 aimonissabaudiae +
4 aloba +
5 alperti +
6 anatolica +
7 angulata +
8 angulinodis + + +
9 arisana +
10 arnoldii +
11 bactriana +
12 bakurianica +
13 bergi + +
14 bibikoffi +
15 boltoni +
16 brancuccii +
17 cachmiriensis +
18 cagnianti +
19 caucasicola +
20 collingwoodi +
21 commarginata +
22 constricta +
23 curiosa +
24 deplanata + +•

25 displicentia .. +
26 divergens +
27 draco +
28 dshungarica +
29 eidmanni + +
30 emeryi +

679
 Table 4.1. Continued.

Species diversity centres

Species
C-Asia SE-Asia Tib M-Asia Sib Far-E Eur-Sib Med W-step
31 ereptrix +
32 excelsa +
33 ferganensis +
34 forcipata +
35 foreliana +
36 fortior +
37 gallienii + +
38 georgica +
39 gigantea +
40 hecate +
41 hellenica +
42 heterorhytida +
43 hirsuta +
44 hlavaci +
45 indica +
46 inezae +
47 inucta +
48 jennyae +
49 jessensis +
50 juglandeti +
51 kabylica +
52 kamtschatica + +
53 karavajevi +
54 kasczenkoi +
55 kirghisorum +
56 koreana + + +
57 kotokui +
58 kozakorum +.
59 kozlovi + +
60 kryzhanovskii +
61 kurokii +
62 laurae +
63 lemasnei +
64 lobicornis + + + +
65 lobulicornis + +
66 lonae + +
67 luteola +
68 margaritae +
69 martensi +

680
 Table 4.1. Continued.

Species diversity centres

Species
C-Asia SE-Asia Tib M-Asia Sib Far-E Eur-Sib Med W-step
70 mirabilis +
71 mixta +
72 multiplex .+
73 myrmicoxena + •

74 nitida +
75 obscura +
76 onoyamai +
77 ordinaria +
78 orthostyla +
79 pachei +
80 pararitae +
81 pelops +
82 petita +
83 phalacra +
84 pisarskii +
85 pleiorhytida +
86 poldii +
87 polyglypta +
88 pulchella +
89 ravasinii +
90 rhytida +
91 rigatoi +
92 ritae +
93 rubra + + + + +
94 ruginodis + + + + +
95 rugosa +
96 rugulosa +
97 rupestris +
98 ruzskyana +
99 sabuleti + + +
100 salina +
101 saposhnikovi +
102 scabrinodis + + + +
103 schencki + + + +
104 schoedli +
105 schulzi +
106 sculptiventris +
107 serica +
108 siciliana +

68i
 Table 4.1. Continued.

Species diversity centres

Species
C-Asia SE-Asia Tib M-Asia Sib Far-E Eur-Sib Med W-step
109 sinensis +
110 sinoschencki +
111 slovaca + +
112 smythiesii +
113 specioides + + +
114 spinosior +
115 stangeana +
116 sulcinodis + + + + +
117 symbiotica +
118 taibaensis +
119 tamarae +
120 tenuispina + +
121 tibetana +
122 titanica +
123 tobiasi +
124 transsibirica + +
125 tschekanovskii +
126 tulinae + +
127 turcica +
128 urbanii +
129 vandeli + +
130 varisculpta +
131 villosa +
132 vittata +
133 wardi +
134 weberi +
135 weii +
136 wesmaeli +
137 williamsi +
138 wittmeri +
139 xavieri +
140 yamanei +
141 yunnanensis +
142 zojae +
total number
33 30 4 14 19 14 22 32 16
of species
number of endemic 31 30 3 8 8 7 7 21 4
species (%) (94) (100) (75) (57) (42) (50) (32) (66) (25)
 Table 4.2. Distribution of the Myrmica species groups in the species diversity centres
(abbreviations of the centres see in Table 4.1).

Species groups Species groups diversity centres

(number of species) C-Asia SE-Asia Tib M-Asia Sib Far-E Eur-Sib Med W-step
1 arnoldii (1 ) 1
2 dshungarica (4) 4
3 inezae (3) 2 1
4 karavajevi (3) 1 2
5 kurokii (2) 1 1 1
6 laurae (1 ) 1
7 lobicornis (22) 3 11 8 5 7 1
lobicornis-complex
2 4 4 3 4 1
(11)
excelsa-complex (3) 1 2 1
kasczenkoi-complex
5 1
(5)
sulcinodis-complex
1 1 2 1 1
(2)
xavieri-complex (1) 1
8 luteola (2) 1 1
9 myrmicoxena (1) 1
10 pachei (14) 3 11
11 ritae (21) 7 14
12 rubra (4) 1 1 2 2 2 2 2.
13 rugosa (9) 9
14 scabrinodis (26) 2 2 12 14 9
scabrinodis-
1 2 4 1
complex (4)
bergi-complex (3) 1 1 1 2
rugulosa-complex (3 2 1
sabuleti-complex (6) 5 4 1
specioides-complex
1 2 4 5
9)
15 schencki (11) 1 2 2 2 1 5 4
16 smythiesii (5) 3 2
17 tibetana (3) 1 1 2
number of species
7 6 3 5 5 5 6 6 4
groups
number of endemic 1 1 0
1 0 1 1 1 0
species groups
number of species
with uncertain 7 1 1 1
taxonomic position

683
centre and the six northern centres are somewhat higher (compared to the C-Asian and
Tibetan centres) due to the presence of species from the rubra- and schencki-groups.
However, these two groups have been found only in the northern Chinese part of the
centre and when the distribution of the ant fauna of China is better known, the Chinese
boundary between the SE-Asian and Far-Eastern centres might be redrawn.
The values of JC between pairs of the six northern centres of diversity, that partition up
the vast northern Palaearctic land mass, more or less reflects the geographical distance
between them. The Far Eastern centre and the M-Asian centre have relatively low simi­
larity to one another (JC = 0.25) and the other four centres (JC's are < 0.50 in both cases).
The Euro-West Siberian, Siberian-Mongolian, the Mediterranean and the Western Steppe
centres all have species from the rubra-, schencki-, lobicornis- and scabrinodis-groups,
consequently similarity values are higher (values of JC are generally > 0.80, Table 4.3).
However, if one splits the lobicornis- and scabrinodis-groups into their various com­
plexes the differences between eastern and western centres (especially between the
Siberian-Mongolian and Euro-West Siberian) is magnified (see Table 4.2 for distribution of
complexes).
We repeated the analysis (Table 4.4) based on species presence listed in Table 4.1, which
further amplified the pattern seen in the analysis of species groups (above). Of the three
oriental centres, only C-Asia shares any species with other centres: M. kozlovi (Tib) and
M. tenuispina (M-Asia). Consequently, the three oriental centres have very little similarity
to each other and no similarity to any of the other centres except M-Asia. The six northern
centres are more similar to each other, although similarities are not high; on average the
centres have in common about 13% of their combined species diversity (i.e. JC = 0.13) and
this never exceeds 31% (Table 4.4). As with the analysis based on species groups, simi­
larity between these centres mainly increases with geographical proximity (compare Map
144 with the values of JC in Table 4.4).

Table 4.3. Jaccard's coefficient (JC) of the similarity of Myrmica species groups between the nine
centres of Myrmica species-diversity. For abbreviations of centre names see Table 4.1. The
diagonal cells give the number of species groups in each centre; the bottom left cells give the
number of the species groups in common; the upper right cells give the values of JC which varies
from 0 (no similarity) to 1 (identical). [Jaccard's coefficient is calculated as JC = с /'(a + b-c),
where a and b are the number of species in two centres compared and с is the number of species
in common].

Centres - C-Asia SE-Asia Tib M-Asia Sib. Far-E Eur-Sib Med W-step
C-Asia 7 0.30 0.29 0.09 0 0.09 0 0 0
SE-Asia 3 6 0 0.10 0.25 0.38 0.22 0.22 0.25
Tib 2 0 2 0 0 0.17 0 0 0
M-Asia 1 1 0 5 0.50 0.25 0.43 0.43 0.50
Sib 0 2 0 3 4 0.50 0.80 0.80 1.00
Far-E 1 3 1 2 3 5 0.43 0.43 0.50
Eur-Sib 0 2 0 3 4 3 5 1.00 0.67
Med 0 2 0 3 4 3 5 5 0.80
2

W-step 0 0 3 4 3 4 4 4

Ó84
Table 4.4. ]accard's coefficient (JC) of the similarity in Myrmica species between the nine centres
of Myrmica species diversity; the layout of the table is the same as that in Table 4.3.

Centres C-Asia SE-Asia Tib M-Asia Sib Far-E Eur-Sib Med W-step
C-Asia 33 0.00 0.03 0.02 0.00 0.00 0.00 0.00 0.00
SE-Asia 0 30 0.00 0.00 0.00 0.00 0.00 0.00 0.00
Tib 1 0 4 0.00 0.00 0.00 0.00 0.00 0.00
M-Asia 1 0 0 14 0.10 0.04 0.09 0.05 0.15
Sib 0 0 0 3 19 0.27 0.21 0.11 0.21
Far-E 0 0 0 1 7 14 0.09 0.05 0.07
Eur-Sib 0 0 0 3 7 3 22 0.26 0.31
Med 0 0 0 2 5 2 11 32 0.14
W-step 0 0 0 4 6 2 9 6 16

Based on species co-occurrences, the six northern centres can be divided into Western
and Eastern sub-regions (the latter comprising trie Far-Eastern and Siberian-Mongolian
centres), with the border between them being the boundary between the West and East
Siberia (i.e. the Enisei River) (Map 145). There is very little similarity between the faunas
of these sub-regions, they have 59 and 26 recorded Myrmica species respectively with only
eight species common to both. Two are the very widespread transpalaearctic boreal species,
M. ruginodis and M. sulcinodis; three other species are nearly as widespread having
a boreal type of distribution, occupying chiefly different kinds of taiga and mixed
forests (M. rubra, M. lobicornis and M. angulinodis); two are schencki-group species
(M. schencki and M. koreana) that are an example of geographic vicariation (with
overlapping ranges in eastern part of the West Siberia and west of the South Siberia),
inhabiting dry meadows, sandy dunes, steppe and steppe-like habitats even in the Forest
Zone; finally, only M. scabrinodis from among the species associated with European
deciduous and mixed forests occurs east of the Enisei River (Table 4.5).

Map 145. Borders of the West and East Palaearctic.

685
 Table 4.5. The Myrmica species common to pairs of centres ranged from
West to East across the Palaearctic.

M-Asian Siberian-
, Centres Mediterranean West Steppe :. Far East
mountains Mongolian
Euro-W- ruginodis ruginodis rubra ruginodis ruginodis
Siberian rubra rubra rubra
lobicornis lobicornis lobicornis
sulcinodis sulcinodis sulcinodis sulcinodis
scabrinodis scabrinodis scabrinodis
specioides specioides angulinodis angulinodis
sabuleti sabuleti
lobulicornis schencki schencki schencki
lonae gallienii
tulinae slovaca
vandeli
Mediterranean ruginodis rubra ruginodis ruginodis
rubra rubra
lobicornis lobicornis
sulcinodis sulcinodis sulcinodis
scabrinodis scabrinodis
sabuleti
specioides
West Steppe bergi ruginodis ruginodis
rubra rubra
lobicornis
schencki schencki
deplanata koreana koreana
scabrinodis
M-Asian rubra
mountains sulcinodis sulcinodis
schencki
Siberian- ruginodis
Mongolian sulcinodis
angulinodis
koreana
eidmanni
kamtschatica
transsibirica

4.2.2. Interpretation of the diversity between the centres

For more than thirty million years the Palaearctic and Orientai Zoogeographie Regions
have comprised a huge contiguous land-mass with few oceanic barriers to the spread and
exchange of species between regions. The only true physical barriers have been various
mountain systems of which we believe the Tibetan Plateau has had the greatest effect (see
Chapter 4.3, below). Otherwise the main barrier to the spread and exchange of Myrmica
species has been climatic; as indicated in Chapter 1.3, most modern Myrmica species are
adapted to relatively warm, but not hot, and moist habitats, and have evolved specialized
physiology and life history strategies that enable individuals and colonies to survive in
strongly seasonal habitats, provided summer temperatures are sufficiently warm! Over the

686
time during which Myrmica has evolved, the climate of the Earth has varied widely due to
a complex interaction between the heat from the sun, oceanic and atmospheric circulation
and mountain building events, that has caused prolonged hot, cool, wet and dry periods
throughout geological time. The position and relief of any spot on Earth created local
climatic variation within the overall trends.
Latitudinal position obviously underlies climate differences due to the amount of sun-
light reaching the Earth's surface that decreases as one moves north (or south) from the
equator and which is regulated by the predictable astronomical movements of the Earth and
Sun. This creates the familiar climatic zones, i.e. the tropical, subtropical, temperate and
polar zones with their different lengths of season and annual mean temperatures. Within
a zone mean temperatures can vary according to short-term cycles (e.g. sunspot cycles) and
longer term factors (e.g. changes in global circulation and composition of the atmosphere, and
size of the Earth's ice caps) , In geological time continental drift might shift a particular spot
on the Earth's crust between climatic zones. With respect to the current zonal distribution
of Myrmica ants at sea level one can generalise that north of the arctic circle summers are too
short for colonies to complete their life cycle, and south of latitude 35°N summer tempera-
tures are too hot for them to forage and compete with ants of other genera. While the cooler
tops of high mountain systems enable Myrmica species to live south of latitude 35°N, high
mountains form barriers in the north. '
Apart from zonal climate the exact position east or west (longitude) within
a continent determines climate on a provincial scale. Provincial climatic variation is greatly
influenced by physiographic factors such as distance from an ocean, the size and position
of inland seas (e.g. Caspian Sea) and the position, orientation and height of mountains.
Physiographic factors that seem to have existed for ever in terms of human history change
quite quickly on a geological time scale (e.g. the extent and height of the Tibetan Plateau).
At the present, the main provincial difference that affects Myrmica distribution at sea level
in the Palaearctic is the degree of continentality, i.e. the trend for increasingly hot and dry
summers and cold winters towards the centre of a continent. This is greatly skewed with
eastern regions having a more continental climate than western ones.
Comparison of the distributions of present-day taxa suggests to us that there are five
major provinces for Myrmica species diversity centred on the Tibetan Plateau, and it seems
highly probable that the formation of the Tibetan Plateau had a huge impact on the subse-
quent spread and speciation of the Old World Myrmica fauna. While there is general agree-
ment that the Tibetan Plateau formed when the Indian sub-continent collided with Asia,
there has been considerable debate regarding the timing and resulting pattern of uplift of the
Earth's crust (see Wang et al. 2.008). Various models have suggested that the process began
> 50 Ma with Plateau growth starting ih the late Oligocene (30 Ma) and proceeding slowly,
alternatively some models suggest a more abrupt uplift about 7 Ma. Wang et al. (2008) be-
lieve geological data from Tibet indicates at least a two-phased process; they suggest that
by 40 Ma a geographically smaller but higher "Proto-Tibetan" Plateau had already formed,
generally exceeding altitudes of 5000 m a.s.l. and by 20 Ma the Plateau had grown to the
east with the up-thrust of the Kunlun Mountain chain (in present day China). During the
last 20 million years they suggest that the great Himalayan ridge formed in the southwest
and the Qilian Mountains of China in the northwest, while the original plateau has reduced
in altitude to less than 5000 m a.s.l.
We hypothesise that by the Late Eocene a fairly diverse Myrmica fauna had already
formed, of which ants that morphologically resembled ritae-group species lived at least in

687
regions where, amber was generated (see Radchenko et al. 2007 and below for further
discussion of this). As the Tibetan Plateau grew, it formed a major barrier for exchange of
fauna between what is now the Palaearctic and Oriental Regions, the barrier being both
physical and climatic, thus by 20 Ma the faunas were already quite different. During the last
20 million years the Himalaya has further isolated the fauna of the C-Asian centre from
the SE-Asian one. At the same time, Tibet started developing its own specialist high altitude
tolerant fauna as it reduced in altitude, derived perhaps from species invading from the east,
the increasingly high Kunlun and Qilian ranges tending to isolate its fauna more
recently. For most of the last 20 million years the habitats available to Myrmica in the
C-Asian and SE-Asian centres have been mountain zones within a generally sub-tropical
climate. Species have been able to find refugia by migrating up or down mountains during
periods of climate change, but at the same time the subtropical conditions at the base óf
mountains has restricted them to their mountain "islands". Thus there are a considerable
number of "old" lineages persisting in these regions as well as a relatively large number of
unusual endemic species.
Very probably the 15-20 degree difference in latitude between the mountainous
territories of the Oriental Region and the areas to the north of the Tibetan Plateau had
already led to the establishment of differing suites of Myrmica species in these regions
20 Ma. However, the fauna isolated on the north and northeast of the Tibetan Plateau
would have always been "more connected" geographically with the main Palaearctic
centres, especially in warmer periods. Isolation would have occurred during various periods
of desertification and in cold periods when Myrmica were forced into different southern
refugia.
Take for example, the Middle Asian Mountain centre that has about 60% endemic
species: it is much more isolated from the adjacent mountain systems (Tibet, Himalaya)
than from the territories laying in the north and west. Most of the species are not able to
cross the extremely high mountain ranges of the north-east Afghanistan (Hindukush and
Karakorum) while the Takla Makan desert separates the eastern Tien-Shan from the Kun-
lun range of the Tibetan Plateau. M-Asia has most species in common with Western steppe
(JC = 0.15), because steppe species can fairly easily reach Tien-Shan from the north-west.
On the other hand,-many boreal species cannot cross dry and hot semi-desert habitats of
Dzhungaria and north-western Kazakhstan and only two boreal species (M. rubra and
M. sulcinodis) are found in the Middle Asian Mountains.
The nature of the connections between the other centres is illustrated by the East Si-
berian-Mongolian centre which shares species with its western neighbour (the Euro-West
Siberian centre) and with its eastern neighbour (the Far Eastern centre), but these last two
have only three very widespread species in common (M. ruginodis, M. sulcinodis and
M. angulinodis). The Western Steppe centre has the lowest proportion of endemic species
(25%), which is not surprising, because it is defined mainly by ecological factors rather than
by geography. The result we found the most surprising when making these analyses was
the richness and originality of the Myrmica fauna of the Mediterranean centre. The num-
ber of species supported is of the same order of magnitude as found in the long-established
C-Asian and SE-Asian centres with 65% being endemics, the highest proportion among the
six northern centres.
Finally, our work during the last decade on the Myrmica from the Himalaya, China and
South-East Asia has enabled us to make some important and unanticipated generalisa-
tions about the genus (we have described 43 new species from this area). Previously, all

688
authors including ourselves, have considered the genus Myrmica to be characteristic of the
Palaearctic temperate zone, being one of the keystone ant genera of the Forest Zone of
Eurasia, i.e. coniferous (boreal), mixed and deciduous forests and temperate grasslands.
However, from among the 142 Old World Myrmica species, at most only 47, about one
third, can be regarded as Eurasian Forest Zone dwellers. Almost a half the species (63) live
in the Oriental Region - in the Himalaya and South and South-East Asia, and possibly fur-
ther south (see notes on M. emeryi above). A further 32 species (23%, probably an under-
estimate of the true total) are endemics of the warmer and drier Mediterranean Zone,
Middle Asian mountains and Steppe Zone. Thus, while from an ecological and bioceno-
logical point of view it may be correct to consider Myrmica a typical genus of the Eurasian
Forest Zone, from a faunistic and zoogeographical point of view it is a genus of southern
mountains. Many of the Oriental species are basa! morphologically for Myrmica (see below),
so perhaps we can turn on its head the old idea that many Himalayan and Oriental Myrmica
species (or Oriental Myrmica fauna as a whole) are "exotic", "peculiar" or "unusual" when
compared to the well-studied European species, it is probable that the boreal fauna is
derived and "exotic".

4.3. Origin and evolution of the genus Myrmica

4.3.1. Putative ancestor and estimated time of Myrmica origin

The genus Myrmica belongs to the tribe Myrmicini of the subfamily Myrmicinae (see
Chapter 4.2.1). For many years, the oldest palaeontological evidence for Myrmicinae
genera came from Middle Eocene deposits in the USA (Wilson 1985; Dlussky and Rasnit-
syn 2002), but recently Dlussky et al. (2004) pushed the age much further back when they
described the genus Afromyrma from the South'African shales that are dated as Turonian
(ca. 90 Ma). This agrees with the confirmed age of the existence of the modern subfamilies
(Formicinae, Ponerinae, Dolichoderinae, Myrmeciinae and Aneuretinae) of between 80 and
90 Ma found in the ambers and shales of different continents, dating from the Late Creta-
ceous Turonian-Campanian Ages (Grimaldi et al. 1997; Dlussky 1999; Grimaldi and Agosti
2000; Grimaldi and Engel 2004; Engel and Grimaldi 2005; Dlussky and Rasnitsyn 2007).
Even at that time some representatives of these subfamilies were quite highly specialized so
one might suppose that the modern subfamilies originated even earlier, perhaps in the Ceno-
manian Age (97.5-91 Ma), although only Armaniidae and Sphecomyrminae are known
from the Albian and Cenomanian Ages (113-91 Ma).
Based mainly on .molecular analyses ("molecular clock") but also including taking
account of some morphological and palaeontological evidence, Moreau et al. (2006) put
the estimated time of origin of the extant ant subfamilies in the Jurassic epoch (> 150 Ma)
while Brady et al. (2006) suggest the Early Cretaceous age (ca. 115-135 Ma). Based on
palaeontological data we consider that the Early Cretaceous time of origin of extant ants (i.e.
excluding Armaniidae and Sphecomyrminae) is more probable than a Jurassic one. The
same authors estimate different times for when the Myrmicinae lineage was established,
varying from ca. 100-115 Ma (Moreau et al. 2006) to ca. 80-90 Ma (Brady et al. 2006)
which brackets the time ca. 100 Ma that was supposed by Ward and Brady (2003) and
Dlussky and Rasnitsyn (2007). Therefore we suppose the Myrmicinae originated at least 90
Ma and a time of 100 Ma seems quite reasonable.

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 Emery (1921) placed the tribe Myrmicini at the base of his "genealogical tree" of the
Myrmicinae, based only on plesiomorphic characters. We agree the tribe Myrmicini is most
probably ancestral for the subfamily Myrmicinae because its members have a plesiomor-
phic state of many features and it seems to be the most primitive morphologically. For
example, the female castes have 11-12-segmented antennae and the males have 12-13-
segmented antennae; individuals usually have well developed pectinate spurs on the
middle and hind tibiae, and 6-segmented maxillary and 4-segmented labial palpes (at least
in some genera). Additionally, the species generally have well developed eyes and a simple
unmodified shape of the head, mandibles, alitrunk and waist (Bolton 2003; Radchenko et
al. 2007). However, the morphological evidence is less clear as to which genus might
represent the ancestral form of the tribe Myrmicini.
In many genera of Myrmicini the wing venation (both males and gynes) has a ple-
siomorphic state (see also Chapter 2.1). Brown and Nutting (1950) believed that a com-
pletely closed set of cells {l+2r, 3r, rm and mcu) and a quadrangular cell rm was the
primary, initial type of wing venation in the subfamily Myrmicinae; such wing-types are
common in the subfamily Ponerinae and some other primitive subfamilies. As some Pogo-
nomyrmex males had quadrangular cell rm and they believed it to be triangular in most
extant Myrmicinae genera, Brown and Nutting inferred that the genus Pogonomyrmex
represented the archetypal form. However, Barry Bolton (pers, comm.) has found quad-
rangular or even pentagonal rm cells in some Myrmica, Messor and even Aphaenogaster
(besides Pogonomyrmex) species, and has recorded many examples of closed cell 3r in more
than 30 Myrmicinae genera. Recently, Dlussky and Radchenko (2009) discovered males of
two extinct Myrmicini genera in ca. 40 million years old Late Eocene European ambers.
They considered that one of the genera, Protomyrmica Dlussky and Radchenko, represents
the most primitive wing venation in the subfamily Myrmicinae.
Taking account of many different characters of both male and female castes of Myrmicini
genera, one finds that all possess a mixture of plesiomorphic and apomorphic features. Let
us consider the following three examples. First, Manica species have plesiomorphic charac-
ters, including a full complement of mesosomal sutures with some large worker specimens
having a rudimentary scutellum, and males have a filiform antennal funiculus without
a developed antennal club, but on the other hand they have specialized mandibles with
a greater number of teeth, and the propodeum of workers and queens lacks spines. Secondly,
we have suggested (Radchenko and Elmes 2001b; Radchenko et al. 2007) that workers of the
ancestral Myrmicini were probably most similar to modern Myrmica or perhaps Manica
species, because the other genera possess more derived features (see also Bolton 2003), but
the forewing of Myrmica and Manica have cells 1 +2r and rm partly separated by a reduced
vein 2+3RS (apomorphic state). Thirdly, while the wing venation of Pogonomyrmex and
Hylomyrma is more primitive (see above) they have a reduced number of the maxillary and
labial palpes (apomorphy). Even the two fossil genera Protomyrmica and Plesiomyrmex
Dlussky and Radchenko are not strong contenders to be ancestral: although they have very
primitive wing venation and antennal structure (in the first genus the antennae are filiform
and in the second they are "sphecoid", i.e. the length of the funicular segments gradually de-
creasing from the second one towards the apex, and the apical segment is the shortest); they
have also apomorphies not seen in any other extinct or extant Myrmicinae genera; see
Dlussky and Radchenko 2009). Thus while Protomyrmica and Plesiomyrmex were probably
dérivâtes of the ancient, basal stem of the tribe Myrmicini and persisted long enough to
co-exist with probably younger Myrmicini genera, such as Myrmica, it is unlikely that they
or any other known Myrmicini genus was the direct ancestor of the tribe.

690
 If this is the case, then the time of origin of the ancestor of the tribe Myrmicini would
seem to be considerably older than the Late Eocene, but how old is the tribe Myrmicini?
Jansen et al. (2010) based on the molecular-phylogenetic analysis deduced the probable
time of origin of the tribe as 45-55 Ma (Middle Eocene-Paleocene). Taking account that the
tribe Myrmicini most probably is basal for all modern Myrmicinae genera (see above), we
generally agree with this date, but suppose that an ancestor already existed in the Early
Eocene or even Late Paleocene (i.e. earlier than 50 Ma). There is even less Paleontologi-
cal evidence for the time of origin of the genus Myrmica. Compared to some modern
genera, specimens of fossil Myrmica are quite rare and are found exclusively in the Late
Eocene ambers (Radchenko et al. 2007) ; by this time they were already rather diverse so we
suppose that the genus Myrmica differentiated sometime in Middle- or even Early Eocene
epoch (ca. 45-50 Ma).
Furthermore, biogeographical data provides some clues to the diversification of
Myrmicini and origin of its modern genera (e.g. Myrmica, Manica, Pogonomyrmex and
Hylomyrma). Myrmica species are distributed mainly in the Holarctic, but also in the
northern' parts of the Orientai and Neotropical Regions. Manica has an exclusively
Holarctic distribution: M. rubida (Latreille) lives in the mountains of Europe and Cauca-
sus, M. yessensis Azuma is restricted to the volcanic mountains of Japan, and four
Nearctic species (including one social parasite) inhabit mountains of USA and Canada.
Moreover, compared to Myrmica, the genus Manica has a somewhat primitive morphology
(see above) and amore "primitive" biology (Cammaerts and Cammaerts 1984,1985,1987),
and this combined with its disrupted distribution suggests that it might be a quite old and
relict genus. Pogonomyrmex species are distributed in the semi-arid and arid regions of the
southern Nearctic and northern Neotropical Regions, while Hylomyrma is a Neotropical
genus. This suggests that Pogonomyrmex and Hylomyrma almost certainly originated in
the New World as probably also did Manica, whereas the genus Myrmica probably had an
Old World origin (see below and Jansen et al. 2010).
If one accepts this scenario, then the common ancestor of the modem Myrmicini
genera should have already existed before the complete separation of North America and
Eurasia. The northern Atlantic started opening by the end of the Cretaceous epoch (ca. 65
Ma), but land connection between America and Europe persisted via the so-called Atlantic
Bridges. The most important for the interchange of flora and fauna was the Thulean Bridge
that connected Europe to Greenland through the British Isles, persisting until the Early
Eocene (ca. 50 Ma) (McKenna 1983; Tiffney 1985; see also Samartin et al. 2001). On this
basis one would predict that the Myrmicini ancestor had already evolved by the time the
Thulean Bridge was broken (50 Ma), which corresponds with putative time of origin based
on molecular and palaeontological evidence (see above). Further consideration of the ori-
gin and evolution of the Myrmicini genera is outside the main task of this monograph. We
now consider possible ways by which the genus Myrmica evolved and adapted to produce
the 142 extant Old World species.

4.3.2. Evolution within the genus Myrmica and origin of the species groups

4.3.2.1. Ancestral Myrmica forms and place of their origin

Less than 10 pieces of Late Eocene European ambers contain fossilized Myrmica, from
among more than 17,000 known examples containing ants (see Dlussky 1997; Dlussky and
Rasnitsyn 2007, 2009). Fossils from the more recent Oligocene (38-25 Ma) and Miocene

691
(25-7 Ma) deposits of Eurasia that are associated with temperate conditions, contain quite
a rich ant fauna, including several Aphaenogaster species that are ecologically similar to
Myrmica (Heer 1849, 1867; Mayr 1867b; Emery 1891b; Théobald 1937; Dlussky 1981;
Zhang 1989; Dlussky and Rasnitsyn 2007). However, to date no fossil Myrmica species
have been found in these deposits. This scarcity of fossil Myrmica suggests that compared
with many modern genera, Myrmica ants were generally rare in the Miocene epoch, at least
populations were probably ecologically specialized with quite limited local distributions,
until sometime in the Pliocene epoch (7-2.5 Ma). Unfortunately, there are no data on the
Pliocene ant fauna of Eurasia to indicate how adaptive radiation proceeded to the extent
that Myrmica is now one of the dominant ant genera of the temperate regions of the
Holarctic, whether measured in terms of the number of species, colonies or individuals.
The morphology of the fossil Myrmica (Radchenko et al. 2007) suggests that the female
castes of the ancestral species had a long and smoothly curved antennal scape, with no
thickening, carinae or lobes. There are no equivalent fossil males, but a short scape is the
plesiomorphic evolutionary condition in males for all ants (Dlussky 1983; Dlussky and
Fedoseeva 1988). The males of the extinct fossil Myrmicini genera Protomyrmica and
Plesiomyrmex have a short scape (Dlussky and Radchenko 2009) and it is believed that the
males of all other Myrmicini genera have a short scape (Barry Bolton, pers. comm.). Thus
we are almost certain that males of the ancestral Myrmica species had a short scape and that
the two morphological features that are most important in defining some modern Myrmica
species groups (a sharply angled antennal scape of the female castes and a longer antennal
scape among males) were evolved during the adaptive radiation of Myrmica.
Based on these and other morphological features we have suggested that the ritae
species group can be considered as fairly basal in Myrmica evolution (Radchenko 1994b;
Radchenko and Elmes 2001b; Radchenko et al. 2007). This view is supported by the ob-
servation that four of the five known fossil Myrmica species from the Late Eocene European
ambers are very similar to the modern ritae-group species and based solely on morphologi-
cal criteria would be placed to this group (Radchenko et al. 2007). We also suggested that
ancestral Myrmica species were primarily arboreal (loc. cit.); this style of life is very
unusual for the vast majority of modern Myrmica, but recent field studies shown that some
ritae-group species might live and forage entirely on trees (Radchenko et al. 2007). We sug-
gested that the genus originated and evolved in the subtropical Eurasian forests during the
Eocene epoch, being adapted to exploit the cooler mountain tops, and as these conditions
have persisted more or less continuously in the northern part of the Oriental Region several
ancient lineages have survived there.
The recent molecular-phylogenetic study of Myrmica by a group of scientists from
Helsinki, Finland (Jansen et al. 2009, 2010), suggested that all the modern species of Myr-
mica studied had a common ancestor about 36 Ma (see Fig. 332). This ancestor was pre-
sumably morphologically similar to the ritae-group-like specimens found in European
amber. Jansen et al.'s results indicate that about 35 Ma the ancestral form gave rise to two
distinct lineages. One of the lines was ancestral to most of the American species studied;
the scabrinodis-group species diverged from this lineage about 25 Ma, which much later
(about 10 Ma) also gave rise to the lobicornis- and schencki-group species. The second of
the lines that derived from the common ancestor 35 Ma contains most of the Oriental
Region species (included to the phylogeny) and which on morphological criteria we believe
to represent "old" lineages. A line that gave rise to modern rugosa-group species diverged
from this lineage about 32 Ma; interestingly this lineage appears to have also given rise to

692
 a North American species (M. wheeleri) that shared a common ancestor with modern
rugosa-group species about 27 Ma. According to this phylogeny the modern rubra- and
ritae-group species shared a common ancestor with the rugosa-group about 32 Ma and
diverged from each other about 30 Ma. The unusual species M. luteola appears to have
shared a common ancestor with modern ritae-group species about 27 Ma (Fig. 332). From
the little that is known about the ecology of species from the ritae-, rubra- and rugosa-
groups, they appear to forage more readily in small trees and shrubs compared to the other
major lineage (that gave rise to scabrinodis-group species etc.) that we believe have many
"more derived" morphological features that we suggest may be associated more with
a "ground foraging" life-style.
Almost certainly the adaptive radiation of Myrmica species was connected with the
cooling and aridization of the climate that started in the Late Oligocene and continued
through the Miocene and Pliocene epochs, the period of time during which the Tibetan
Plateau formed (see Wang et al. 2008 and below). By the Pleistocene (the last 2-3 million
years) the modern species groups and many species had formed and the alternating glacial
ages played an important role in subsequent speciation within groups. Throughout this long
period of time the climate of subtropical India, South and South-East Asia was less
impacted and temperate adapted species could find refugia on high mountains that became
more and more isolated from the rest of Eurasia as the Tibetan plateau increased in size. Not
only ritae-group Myrmica species but also many other species of plant and animal found
in the Late Eocene European ambers still persist in this region (Larsson 1978). Thus we
expect that many of the other unusual Myrmica species of this region might reflect the
morphology of other basal forms of Myrmica that co-existed with the ritae-group at the end
of the Oligocene epoch. For example, M. mirabilis that it is rather similar to Manica by the
general shape of its body, even though is clearly a Myrmica by most features.
It seems logical to postulate that the ancestor of the genus Myrmica first arose in the Old
World rather than in the New World, where all known species (with the exception of
M. wheeleri Weber) have more derived morphological characters with none being similar to
the ritae-like species found in the Late Eocene amber. Furthermore, the morphological
diversity among American Myrmica species is less than that among those of the Old World
and the abundance of species, though surely underestimated (see Chapter 1.1), appears to
be much less (currently < 35 vs. 142 species). The "younger age" of much of the American
fauna is supported by the results of Jansen et al. (2010 - see Fig. 332).
If the genus Myrmica had an Old World origin then one might ask: "Where was the cen-
tre of origin?" Based on modern fauna one might suspect that it was South-East Asia be-
cause the Central Asian Himalayan and South-East Asian centres of diversity contain many
species that represent "older", more basal species groups (see above). However, current
distribution may not be relevant to this question which may never be answered satisfacto-
rily. The limited palaeontological evidence from amber shows that ritae-group species and
other "ancient" forms of Myrmica were certainly present in Europe during the Late Eocene
(ca. 40 Ma) having probably originated about 10 million years earlier (see above). How-
ever, during the entire Cretaceous epoch until the end of the Middle Oligocene (i.e. from ca.
135 to 30 Ma) Europe was more or less continuously separated from Asia by the Turgai
Strait that covered the modern West-Siberian Plain, connecting the Polar Sea and Tethys
Sea, and separated the ancient continents of Gondwana and Laurasia during the Mesozoic
(Kuznetsov 1962; Zonenstein and Gorodnitsky 1977a, b; Briggs .1987; Zonenstein 1988).
The Turgai Straitwas a shallow sèà that posed a huge natural barrier to the exchange of

693
Figure 332. Chronogram of a Myrmica phylogeny redrawn from that published by Jansen et al. 2010
(Fig. 2 ibid.); the scale gives age in Ma and the major geological epochs. The lineages that terminate
in Nearctic species are truncated by a triangle. Figure redrawn by Günther Jansen, March 2010.

694
species in many plant and animal groups, especially ants. Thus if the genus Myrmica first
evolved in the European part of Laurasia we would not expect them to reach Asia until the
Turgai Straight began to disappear and land connection between Europe and Asia was re-
established. On this basis we think the centre of origin was very unlikely to have been the
South or South-East Asia despite this being the modern centre of diversity of the ritae-
group (Map 146). This might be one of numerous examples where modern taxonomic
diversity does not correspond with the place of origin of a systematic group.

Map 146. Distribution of theritae-groupspecies.

The closure of the Tethys Sea is supposed to have started when the Indian subcontinent
collided with Asia at the end of Eocene and by the Late Oligocene (ca. 30 Ma) the Turgai
Strait had dried up, causing the climate of Siberia to become more continental (loc. cit). The
Indian-Asian collision also created the Tibetan plateau, which by 30 Ma was already very
high but much less extensive in area than at present (Wang et al. 2008). Even so, it is
believed that this led to considerable changes in the regional (even global) atmospheric
circulation that provoked major changes and cooling in the climate of the Northern Hemi-
sphere. By the end of the Oligocene, these global changes had stimulated the formation of
extensive cool- and drought-resistant deciduous forests in the north while the rich sub-
tropical evergreen forest persisted in Southern and partly Central Europe. Both types of
forest became connected to the similar biomes in the east. As a result, many floristic and fau-
nistic elements were able to interchange, migrations probably occurring in both east and
west directions. We hypothesise it was during this somewhat limited period (35-30 Ma)
that some lineages of Myrmica began adapting to the cooler temperate forests while others
spread to the subtropical evergreen forest of South and South-East Asia and possibly parts
of East Asia, where they became increasingly isolated by a combination of climate change
and the continuing growth of the Tibetan Plateau. The major dichotomy in the phylogeny
at about this time, suggested-by Jansen et al. (2010) (see Fig. 332), could well represent the

695
adaptation to the cooler forests. If Wang et al. (2008) are correct, then the formation of the
Himalaya in the west and the Qilian Mountains in the east during the late Miocene epoch
(ca. 15 Ma) would have further isolated the fauna in the warmer forests to the south and
southeast from the cooler-adapted fauna in the other parts of the Palaearctic. It would also
have effectively increased the isolation between the regions that we recognise as the
C-Asian (southwest Himalaya) and the SE-Asia diversity centres. The loss of the evergreen
broad-leaves forests and its replacement by of other types of vegetation continued through-
out the Palaearctic Region during the Neogene (roughly the last 23 million years). It caused
the extinction of most "old" forms of Myrmica besides many other groups of plants and
animals that were associated with that type of forest with the exception of those species that
had already found refuge in the Oriental Region.
If the genus Myrmica evolved in Europe then when did it reach America? It is highly un-
likely that species from any of the ancient lineages (e.g. ritae-like species) migrated into
North America before the Thulean land bridge was closed. The break-up of the Laurasian
supercontinent, which resulted in the separation of North America and Greenland from
Eurasia is believed to have started by 55 Ma, before Myrmica evolved (ca. 50 Ma, see above).
On the other hand, the eastern Beringian land bridge between Siberia and North America
was open more or less continuously during the Palaeogene (ca. 65-23 Ma) and intermit-
tently since. It is probable that after the closure of the Turgai Strait, the more derived tem-
perate-climate adapted forms of Myrmica spread across the northern Palaearctic and
reached North America by this route, perhaps as early as 23 Ma with several subsequent
waves of invasion.
Thus we hypothesise that the ancestors of the North American Myrmica will be shown
to be a subset of the Palaearctic species derived during the early Neogene from the original,
late Eocene forms. This proposed chronology is supported to some extent by the recent
phylogenetical studies of Jansen et al. (2010). The apparently more limited morphological
diversity of the North American Myrmica can be explained by this hypothesis, though
a much better understanding of the distribution and diversity of the North American species
is needed before this idea can be explored fully.

4.3.2.2. Origin of the species groups and their probable relationships

As there are no post-Eocene fossils on which to base a scenario for more recent Myrmica
evolution, discussion of this aspect is mainly based on morphological and zoogeographical
considerations combined with observations on ecology where known. As indicated above,
we have published a suggestion that two significant morphological divergences occurred
during Myrmica evolution in the Neogene (Radchenko 1994b; Radchenko and Elmes
2001b). One was the formation of a sharply angled antennal scape in the female castes,
often with variously developed lobes or carinae on the outer surface of the bend. The female
castes of all known Himalayan, South- and South-East Asian Myrmica species, belonging
to several very distinct different species groups, have a smoothly curved scape without any
lobes or carinae. An angled scape is associated with species, which mainly occur in Europe,
Middle Asia and Siberia. We suggest that an angled scape in workers has a functional adap-
tation to foraging on the soil surface and more cryptically, under leaves and moss, whereas
long and feebly curved scape may have been more advantageous for arboreal life-style or
foraging on shrubs and vegetation in different kinds of tall-grass habitats. An acutely angled
bend appears to enable the ants to retract the antennae more closely against the sides of
their head and we are sure that when the necessary morphological and histological studies

696
are made, the width of the frons and shape and size of the antennal lobes and other
thickenings will be shown to have a functional relationship with the musculature, which
controls the movement of the antennae. The second major development was the evolution
of a longer antennal scape by males. The functional explanation for a longer male scape is
harder to explain, we suggested that it might be related to swarming behaviour and the dis-
tance and height males fly to find queens. Mating behaviour remains a poorly studied area
of Myrmica biology (see Chapter 1.3).
When considering the possible trends in the evolution in the genus Myrmica, we pos-
tulate that homoplasies iii female morphology have been rare: for example the evolution of
the different basic shape of scape in female castes started independently in different ances-
tral forms of Myrmica and subsequent convergences have not occurred; also morphologi-
cal reverses are impossible (e.g. if the scape evolved to be angled with a certain type lobe or
carina, it cannot evolve back to the smoothly curved state). Similarly, we assume reversals
in the length of scape of males have not occurred (e.g. longer scape cannot evolve back to
the shorter one), although this may be less certain. We also assume that the angled scape
in the female castes was an evolutionary more valuable step than the elongation of the male
scape, and consequently give this character more weight.
Among the Old World Myrmica we recognize 17 species groups and 10 species with
unique autapomorphies (see Chapter 2.2). As far as it goes, the molecular phylogeny pub-
lished by Jansen et al. (2010) provides strong support for our concept of species groups, but
suggests that our idea of morphological species complexes within groups have no relation
to phylogeny. Five species groups are social parasites or have features related to social
parasitism {karavaievi-, laurae-, myrmicoxena-, luteola- and arnoldii-groups) while the
rugosa- and scabrinodis-groups include some social parasites. Only four species groups
have the derived character of female castes with an angled scape (scabrinodis-, schencki-,
lobicornis, and the laurae-group of social parasites), while the remaining eight have female
castes with gently curved antennal scape (dshungarica-, inezae-, kurokii-, pachei-,
ritae-, rubra-, rugosa-, smythiesii- and tibetana-groups, plus the other four groups of social
parasite).
Species groups having females with a curved antennal scape. As has been well dis-
cussed above we consider- the modem riiae-group to be morphologically similar to the
oldest known Myrmica species living at the end of the Eocene but this does not necessarily
imply that they are any more related genetically to these ancestral forms than any other
modern species (see Fig. 332).
The inezae-group includes three species, distributed in the Himalaya and south-western
China (Map 147); males are not known for any species. No specimens were included in
Jansen et al.'s (2010) molecular phylogeny. The females share several features with the
ritae-group, but well differ from the latter by the strongly prominent and not notched ante-
rior clypeal margin, by the rounded apically propodeal lobes, etc. We suppose this group will
be shown to be a relict of an old fauna closely related to the ritae- and rugosa-group line-
ages because differences in their morphologies do not suggest any drastic adaptive mor-
phological reorganization. The rugosa-group (nine species from the Himalaya- Map 148)
also seems morphologically primitive although superficially more like "modern" Myrmica
than say the riiae-group because the female castes of the species tend to have rather
"generalized" feature of Myrmica, e. g. smoothly curved scape, wide frons and not-extended
frontal lobes, frontal carinae that merge with the rugae that extend to the occipital margin.
Also the males have a short scape. Generally, rugosa-group species seem more adapted to

697
 Map 147. Distribution of the inezae-group species.

high altitude habitats than n'tee-group species, so we suppose that this group was directly
derived from a ritae-like ancestor and was isolated in C-Asia gradually adapting to the high
altitudes of the Himalaya (see above), where it underwent speciation. The female castes
of the pachei-group also have rather "generalized" features, but males have a long scape.
It comprises 14 species (and we suspect many more, none of which have been included in
a molecular phylogeny), distributed in the Himalaya and south western and southern China
(Map 149). The main diagnostic feature of this group is a transversale rugose (at least in
part) alitrunk dorsum. From the little that is known about its ecology, we suppose that like

Map 148. Distribution of the rugosa-group species.

698
 Map 149. Distribution of the pachei-group species.

the rugosa-group it derived from a ritae-like ancestor that adapted to somewhat cooler tem-
perate, open conditions that are found at slightly lower altitudes on more northern moun-
tains in the Oriental Region, where local isolation has led to a variety of local forms.
The tibetana-group is interesting, comprising three species distributed in Tibet, north-
eastern Afghanistan and Middle Asian Mts. (Map 150). The female castes have several de-
rived features, especially a wide frons with feebly curved frontal carinae combined with
widely extended frontal lobes. Such shape of the frontal lobes is very rare among Old World
Myrmica, resembling the American incompleta-group and being somewhat similar to the
arnoldii- and myrmicoxena-groups. Males of the two Middle Asian species have short

Map 150. Distribution of the tibetan a -group species.

699
antennal scape, but the situation with the males of M. tibetana is less clear (see Notes to
this species in Chapters 2.2 and 3.3). It seems probable that its ancestral form may have
been derived relatively early-on from the rugosa -group rather than directly from the ritae-
group, because the modem species live at high altitudes, like many п/gosa-group species.
Possibly the ancestral species of the tibetana-group lived on the north of the early Tibetan
Plateau becoming isolated in the Middle Asian mountains when the Himalaya formed
with eventually M. tibetana penetrating the C-Asian centre of diversity from the north (as
discussed above).
The remaining four species groups (dshungarica-, smythiesii-, kurokii- and rubra-
groups) are superficially very similar to each other and differ mainly by the shape of the
frontal carinae and the clypeus (see Chapter 2.2). The males of all these groups have a long
antennal scape and the female castes have a fairly unspecialized morphology. It seems
equally possible that they are paraphyletic, independently evolving from basal, ritae-like or
rugosa-like ancestral species, or that they all derived from a common ancestor that had
evolved a long scape in the males. In either scenario we think that these lineages arose quite
early in the evolution of modern Myrmica (this seems to be the case for the rubra-group -
Fig. 332). It seems probable that the ancestors of the smythiesii-group (five species found
in the Himalaya and Tibet, Map 151) and the kurokii-group (two species distributed from
Himalaya and Tibet through northern China to Japan, Map 152) were isolated on the
southern oriental side of the early Tibetan Plateau and gradually adapted to very high alti­
tude conditions; M. kurokii only relatively recently having "escaped" the isolation of Tibet
and spread across China to Japan. Conversely the ancestors of the dshungarica-group (four
species restricted to the Middle Asian Mts. and NE Afghanistan, Map 153) and the rubra-
group were isolated on the northern side of the early Tibetan Plateau. The dshungarica-
group persisted as a relict fauna in the mountains of Middle Asia whereas the rubra-group
species appear to have retained more of the "arboreal" foraging behaviours of the ancient
ritae-like forms and adapted and migrated with the Forest Zone of Eurasia (Map 154).

Map 151. Distribution of the smythiesii-group species.

700
 Map 152. Distribution of the kurokii-group species.

Currently, there are only four rubra -group species, but they are among the most well-
known and obvious components of the ant fauna of their areas. M. kotokui lives in the
extreme east of the Palaearctic, and M. arisana is a Taiwanese endemic. We thought that
M. arisana might quite possibly have been derived from M. kotokui accidentally introduced
by man to Taiwan sometime in the last 5,000 years, but the molecular phylogehy suggests
that it is quite basal to the modern rubra-group species (Fig. 332). Finally the rubra -group
contains two of the most common and widespread boreal transpalaearctic species (M. rubra
and M. ruginodis). During the Pleistocene these species have at times been very widespread,

Map 153. Distribution of the dshungarica-group species.

701
 Map 154. Distribution of theтаЬга-groupspecies.

as at present, and restricted to various southern refugia during ice periods that might have
produced several "glacial lineages" among the present day populations.
Species groups having females with a bent antennal scape: the phylogeny of Jansen et
al. (2010) (see Fig. 332) suggests that the ancestor of these groups emerged quite early in
the evolution of Myrmica (about 35 Ma). We believe that the force that drove selection for
some of the characteristic morphology of these groups was the need to forage on the soil sur­
face and more cryptically under leaves and moss as the climate cooled and broad-leaved
forests receded and grassy regions became more common following the huge climatic and
ecological fluctuations following the initial Tibetan up-thrust at the end of the Eocene
epoch. Based on morphological criteria we believe that the four species groups below are
all more recently derived from the ancient fauna of the Palaearctic compared to the species
groups having curved antenna. The phylogeny illustrated in Fig. 332 lends considerable
support for this view. In general, these more temperate, open, grassy habitats have changed
more rapidly over the last 15 million years or so than the habitats used by most of the species
with curved antenna and this has led to the variety and abundance of Myrmica species with
bent antenna throughout the Holarctic Region.
Currently the scabrinodis-group contains the most species from among the Myrmica
species groups (26, including three socially parasitic species). All males have a short scape
(SI1 < 0.60, usually < 0.40) and that of the females is strongly curved at their base, often
appearing bent to more or less form a right angle. The shape is quite distinct from the more
gradual, open curve of the antenna seen for example, in the rubra -group. Most species have
a horizontal lobe on the outer surface of the scape bend, though in different species it varies
greatly in shape and size and in some species it is so poorly developed that it appears only
as a carina or ridge, while in a few species it is completely absent. This variability combined
with high diversity among other morphological features within the group led us to divide it
into species complexes (see Chapter 2.2).
The scabrinodis-group has a distinctly West Palaearctic distribution (Europe, North­
west Africa, Caucasus, Asia Minor, Iran, West Siberia, Kazakhstan and Middle Asia) and

702
only two species were found in Siberia east of the Enisei River (Map 155 and above). The
centre of species diversity is the Euro-Caucasian region, where more than 50% of the Myr-
mica species (25 from 48) belong to the scabrinodis-group (five being social parasites - see
below). Most species could be considered as mesophilic dwellers of the Deciduous and
Mixed Forests Zones, where they inhabit many different types of open woodlands, bogs,
marshes and meadows. In many habitats they dominate the ant fauna, both in terms of nest
density and biomass. Generally the members of the specioides-complex are more ther-
mophilic, adapted to the warmer and drier conditions of the sub-Mediterranean regions,
where they live in open and sparse forest and grasslands.
On morphological criteria we had supposed that the ancestor of the scabrinodis-group
evolved somewhere on the territory of modern Europe from an ancestor that lived about 40
Ma (an argument to support the ancient origin of this group may be the plesiomorphic short
antennal scape of the males) ; however the molecular phytogeny (Fig. 332) suggests a more
recent date - at the start of the Oligocene. Initially, the putative more-cryptic, ground-sur-
face foraging ancestor was probably rare, experiencing considerable competition from other
thermophilic ant species that were well-adapted to that niche. However, the group seems
to have been well placed to take advantage of the cooler, more open conditions created as
the temperate deciduous forests started replacing the old broad-leaved forests during the
Oligocene and eventually spread over the vast connected areas across Europe and Asia as
the Turgai Straight dried up. It is not known how far the scabrinodis-group ancestral species
penetrated into Siberia during this time, possibly the present distribution represents the
maximum range the group has ever achieved (Map 155). It seems probable that the speci-
ation in this group that probably started in the Miocene became more active during the
Pliocene as further cooling of the climate caused dramatic changes in the patterns of vege-
tation. It seems probably that the Ice periods of the Pleistocene intensified the process of
local adaptation and we suspect that currently over much of its range, species in the scabri
nodis-group are undergoing active speciation triggered by the changes in habitat distribution

Map 155. Distribution of the scabrinodis-group species.

703
since the end of the last Ice Age (ca. 16 thousand years ago). Currently, Europe appears to
be an active centre of species diversification with the establishment of many closely related
geographical races, subspecies and species, many of which are morphologically cryptic with
adaptation of behaviour and physiology being more important (see Chapters 1.3 and 3.3).
We wait with interest to see what pattern emerges when more species are added to the mo-
lecular phylogeny that produced Fig. 332.
The species of the schencki-group are all also characterised by having males with a short
antennal scape (SI, < 0.50) and by this criteria we suspected that it also has a quite ancient
pedigree (see below). Twenty years ago only four species were included in this group (Rad-
chenko 1994b), but more recently seven additional species have been described (Seifert
2003; Radchenko, Elmes and Alicata 2006; Radchenko et al. 2008, 2009a, c; Elmes et. al.
2009). The antennal scape of the female castes of the schencki-group species is always
strongly angled at the base (more or less right angled) and always has a vertically-
projecting plate, lobe, tooth or ridge. This lobe or ridge can vary considerably in size and
in the exact details of its structure but it always forms a shield-like plate along the short
(vertical) surface of scape (see e.g. Figs 60, E; 242, D).
The schencki-group is distributed from the Atlantic Ocean to Japan (Map 156), but only
a few species have large ranges (e.g. M. schencki, M. deplanata and M. koreana), all the oth-
ers have very restricted distributions, often known only from the type localities (see Chap-
ters 2.2 and 3.3). This kind of fragmented distribution usually characterizes a relict of
a more ancient widely distributed and probably more diverse fauna. We would class all the
schencki-group species as thermophiloús, most live in the more southern, warmer parts of
the Palaearctic (Mediterranean Region, Steppe Zone, central China and central Japan) and
even M. schencki that is distributed in more northerly habitats is considered to be one of
the most thermophiloús species among northern Myrmica species, specializing on relatively
hot dry habitats.
One habit of M. schencki is that colonies make a woven tunnel, funnel or chimney at
the entrance to their nest, and a guard worker often sits in the entrance of the funnel.

Map 156. Distribution of the schencki-group species.

704
The species is well known for this behaviour and in parts of Europe it is referred to as the
Chimney Ant. From our personal observation it seems that at least some colonies of all the
other species of this group make similar chimneys, though these may vary in size and de­
sign. We suspect that this is quite an old behaviour evolved to protect the nest from other
ants in the open grassy habitats preferred by these species. The schencki-group is the only
Old World Myrmica species group that not only shares significant-morphological features
with Nearctic species, but has the propensity to make woven chimneys at the nest entrance
(personal observation). In fact, based on current knowledge many common and widespread
North American forms share the main diagnostic features with the Palaearctic schencki-
group species and could formally be placed to this group. However the results published by
Jansen et al. (2009, 2010) suggest that a close relationship between any of the modern Old
World species groups and modern American species is unlikely, though it remains to be
fully tested whether there is a close relationship between the schencki-group and any of the
American "schencki-likе" species when more species from both continents are added to
the phylogeny.
We suspect that the schencki-group ancestor first adapted to ground-living and
foraging in leaf litter in drier and warmer conditions than was the case for the scabrinodis-
group ancestor and if so, the time of origin of the schencki-group should be somewhat more
recent. The most probable time is the Oligocene-Miocene boundary, when dry, light and
sparse Mediterranean-like forests, as well as steppe-like landscapes formed. Although the
bent scape of the female caste with its ornamentation may have evolved for the same func­
tional and biomechanical purpose as that of scabrinodis-group species we do not think that
the schencki-group ancestor evolved directly from an early scabrinodis-group species, we
think it more probable that they shared a common ancestor (this is view has been supported
by the phylogeny - Fig. 332). We suggest that the group probably originated in Southern Eu­
rope or even somewhere in the Asia Minor-Caucasian region and became well established
during the Miocene. Through subsequent dry periods and extensions of steppe grasslands
in the Northern Hemisphere they probably spread right across the Palaearctic reaching the
Japanese Islands when they were connected with the Asian mainland (probably the Early
Pliocene).
The lobicornis-group is second only to the scabrinodis-group in terms of number of
species (22). By the main diagnostic features of a sharply angled scape with a vertical lobe
or dent (in the majority of species) the queens and workers are very similar to those of the
schencki-group. However, the scape lobe does not form a shield-like plate along the short
(vertical) surface of scape (see Chapter 2.2). Several species have the lateral portions of
their clypeus raised into a sharp ridge in front of the antennal insertions so that the anten­
nal sockets are distinctly separated from the clypeal surface. This feature is also found in the
tribe Tetramoriini but is unique to this group of the genus Myrmica. Also an important dif­
ference is that males of this group have the more derived, relatively long scape (SIX > 0.60).
The lobicornis-group has a transpalaearctic distribution (Map 157) with many species
having wide ranges. While one would hardly classify this group as psychrophilic, they are
generally much more cold-tolerant than scabrinodis-group species. The vast majority of
species of the lobicornis-group are typical boreal dwellers, inhabiting the Taiga Zone and
the cold Siberian steppes that overlay the permafrost soil, but on the other hand, several
species live in the much warmer Mediterranean Region, i.e. the mountains of the Iberian
Peninsula and Asia Minor. The highest regional diversity of lobicornis-group species
occurs in Siberia, 11 of the 22 species are found here, eight live in the Far East (three being

705
 Map 157. Distribution of the lobicornis-group species.

endemic to that region) and a further two species live in the mountains of Middle Asia.
Thus 16 of the 22 lobicornis-group species live within the vast territories of the eastern
Palaearctic.
We suppose that lobicornis-group shared a common ancestor with the schencki-group,
rather than it derived from one of the early schencki-group species (the phytogeny illus­
trated in Fig. 332 supports the suggestion despite the schencki-group being represented by
only three species). Moreover, descent from a schencki-likе ancestor would imply a rever­
sion of the shield-like structure on the scape in the schencki-group female castes to the
more ancestral state and not all species of the lobicornis-group have a developed lobe or dent
at the scape base. Probably, the ecological range of the ancestor of the lobicornis-group
species overlapped the range of the ancestor of the schencki-group, but being more
tolerant to cooler conditions was better adapted to spread in the open cooler forests. We had
supposed that the main time of diversification within the lobicornis-group probably took
place somewhere in Siberia during the Late Miocene and especially in the Pliocene, when
the climate got cooler and deciduous forest and warm steppe were substituted by taiga and
cool steppe, but the phylogeny illustrated in Fig. 332 suggests that diversification occurred
throughout the Miocene epoch.
The social parasites. Here we consider groups having combinations of morphology nor­
mally associated with social parasites (Kutter 1973). We do not consider the problems and
different theories of origin of the social parasitism in ants in general and Myrmica in par­
ticular (see Savolainen and Vepsäläinen 2003 and references therein; see also Chapter
1.3.6), rather the various groups are described in relation to the other groups outlined above.
The laurae-group contains a single workerless social parasite, M. laurae, which is found
in Central Italy (Map 158) where it lives in the nests of M. scabrinodis and M. spinosior.
Based on the combination of features of its queens, it could be placed in the scabrinodis
species group, but its males have a long scape (SI1 > 0.75) and 12-segmented antennae
and so differ from any other male in that group. Probably, M. laurae derived directly
from a scabrinodis-group species and the apomorphic features of its males (reduction of

706
 Map 158. Distribution of the laurae-group species.

the antennal segments and elongation of the scape) may have evolved as an adaptation to
mating behaviour (see notes to M. laurae in Chapter 3.3).
The arnoldii-group also has only one species, M. arnoldii. It has been found in a rela-
tively limited area of South Siberia (from Tuva to eastern Transbaikalia) and northern Mon-
golia (Map 159). While it possesses many features of the so-called "socially:parasitic
syndrome" (see Radchenko and Elmes 2003a), it lives independently and is probably not
strictly a social parasite, though possibly, like M. vandeli, it might spread by temporary so-
cial parasitism. M. arnoldii seems like an alien element in the Siberian fauna because it is

Map 159. Distribution of the arnoldii-group species. '

707
not morphologically similar to any other species distributed in the same or adjacent regions.
Although the queens strongly resemble those of the Central European social parasite,
M. myrmicoxena, a close relationship between the two species seems quite improbable not
only from the zoogeographical point of view, but also because males of M. arnoldii have
12-segmented antennae, while in the latter species they are 13-segmented (in both species
the scape is short). Based on morphology, M. arnoldii could have had many evolutionary
routes, e.g. directly from M. ritae-like species, or from the more derived species groups. The
recent molecular-phylogenetic analysis by Jansen et al. (2010) placed M. arnoldii in the in-
completa species group of the North-American Myrmica and indicates that it derived from
a common ancestor of M. incompleta Provancher and M. alaskensis Wheeler about 15-20
Ma. At first sight, this seems strange, but M. arnoldii does shares several morphological
features with the species of the incompleta-group (e.g. shape of scape and frontal lobes
in the female castes, and short scape of males). The common ancestor could have been
a Palaearctic species and the ancestors of the M. incompleta and M. alaskensis lineages
migrated independently to America via the Beringian bridge leaving M. arnoldii as a relict
species, it is more plausible that M. arnoldii (or its ancestor) represents a later reverse
invasion from North America via Beringia.
The luteola-group species have been suggested as being social parasites based on some
morphological features and somewhat dubious ecological data (see Chapter 3.3) so they
might strictly not be social parasites, at best being temporary social parasites. The group
contains two morphologically remarkable species, M. luteola and M. mirabilis, distributed
in the south-eastern Palaearctic and on Taiwan respectively (Map 160). Sexual castes of the
latter species are unknown, but queens of M. luteola possess a unique feature for Myrmica:
they have no propodeal spines, only blunt tubercles; its males have a short scape. Both
species are large and robust (M. mirabilis is one of the largest Myrmica in the World) and
superficially are similar to Manica species (see Elmes and Radchenko 1998; Radchenko
and Elmes 2003a). The molecular phylogeny (Fig. 332) suggests M. luteola is an old species
derived directly from a basal stock of the genus, perhaps temporary social parasitism has
enabled this archaic form to persist in less favourable climates.

Map 160. Distribution of the lulteola-group species.

708
 The karavajevi- and myrmicoxena-groups include four workerless socially-parasitic
species, distributed in Europe and North-West Africa (Maps 161 and 162) (Radchenko and
Elmes 2003a). The species of the karavajevi-group lives in nests of scabrinodis-group
species whereas M. myrmicoxena was found in nests of M. lobulicornis (the lobicornis-
group). Queens of these social parasites have curved scape without ridge, lobe or carina,
males of the karavajevi-group have 12-segmented antennae and long scape, but antennae
in M. myrmicoxena males are 13-segmented and with short scape. Additionally, both sexes
have a reduced spur on the mid- and hind tibiae, and rather derived wing venation. Species
in both these groups apparently somewhat contradict Emery's rule that postulates that
socially-parasitic species are closely related to their hosts (see Savolainen and Vepsäläinen
2003). None of these species could have derived from their modern hosts because the scape
of the females in the karavajevi-group are "more primitive" than in their hosts, and
M. myrmicoxena males have a short scape while their host-species, M. lobulicornis, has the
derived long scape. Thus the morphology suggests that if the species in these groups share
a common ancestor with their modern hosts they underwent speciation a long time ago and
survived by "shifting hosts" as the host lineage diversified. The molecular phylogeny pro-
duced by Jansen et al. (2010) strongly supports the morphological data: M. karavajevi is
placed in the scabrinodis-group, but is quite distantly related to most of the modern species,
differentiating from their common ancestor about 15-20 Ma.
As a generalisation from Jansen et al. (loc. cit.) it seems that many of the species with
the suite of morphological features associated with social parasitism are not recent additions
to the Myrmica fauna. However, they showed that some species are quite recently related
to their hosts (a strict form of Emery's rule) while others (e.g. M. karavajevi) have a com-
mon ancestor with their hosts at a species group level (see for further discussion of this
Jansen et al. 2010). It would seem to us that opportunities for a parasitic life style occur
when a group of species is undergoing rapid change due to local selection pressurés, such
as we hypothesise is occurring in the scabrinodis-group at this time. This results in various
forms that might be taxonomically close to their host species. If any of these forms is to

Map Ì61. Distribution of the karavajevi-group species.

709
 Map 162. Distribution of the myrmicoxena-group species.

survive as parasites in geological time they either must adapt and make host shifts as their
hosts become scarce and die-out (probably due to general ecological reasons rather than the
effect of parasitism) or revert to a free living life-style. There are probably great costs asso­
ciated with either choice so most parasites are short-lived in geological terms. Thus at any
time one might expect to find some successful less host-specific parasites with quite an­
cient lineages and other more recently derived forms that are closer to their current host
species. The problem in studying this process further using the genus Myrmica is that the
ecology of most of the "old" lineages (e.g. the ritae-, inezae- and расhei-groups) has not
been studied sufficiently intensively to find social parasites, if they exist. For example, the
chance of finding a host nest containing M. hirsuta is < 1 in 100 (Elmes 1994), in many
Myrmica species groups probably less than 100 nests of all species together have been sam­
pled. It seems probable that the older groups do have social parasites because a single queen
of M. ereptrix has been found in a nest of a rugosa-group species. On the other hand, some
populations of the rubra-group species, M. rubra and M. ruginodis that we believe has more
recently expanded across the northern Palaearctic, have evolved specialized microgyne
morphs that in the case of M. rubra behave as pseudo-social parasites (see Chapter 1.3.6,
also Elmes 1976; Savolainen and Vepsäläinen 2003).

710
CHAPTER 5. KEYS FOR THE IDENTIFICATION OF THE MYRMICA SPECIES
OF THE OLD WORLD

The Myrmica fauna is very different between the major Zoogeographie regions (see
Chapter 4) and some regions contain almost entirely locally endemic species. Therefore we
have compiled seven separate regional Keys for the identification of the Myrmica species of
the Old World. In our opinion, this approach is more convenient for users and makes the
identification of species easier and more reliable. A few widespread species (e.g. M. rubra,
M. ruginodis, M. sulcinodis) occur in several of the regions and consequently appear in
several keys. The keys are based mainly on worker characteristics that also occur in the
queen caste, important male characters are included in the Keys.
Users should appreciate that a correct identification is most probable when they have
several specimens from the same colony, preferably of all three castes. In some cases it is
impossible to discriminate between several different species based on female castes without
corresponding males and vice-versa, similarly there can be less confidence of an identifica-
tion based on single specimens than identifications made on samples from the same colony.
If users wish to identify a Myrmica specimen we suggest that they start with the key for the
region where the specimen was taken; if unable to identify it we suggest that they then try
the Keys for adjacent regions. If you are still unable to identify the specimen (especially
when all castes are available) then, given the high rate of endemism in Myrmica (Chapter
4) there is a high probability that you have a new species.
There are eight Keys comprising ones to the extant species of the seven regions (Map
163) and one for fossil Myrmica species, these are numbered as follows:
5.1. West Europe, from the Atlantic Ocean to the border of the former USSR in the east,
including the Iberian Peninsula, Italy, Balkans and the north-west of Africa in the south.
5.2. East Europe, from the border of the former USSR in the west and including West
Siberia and the Altai Mts., the Caucasus, Asia Minor, Iran, the Kopetdagh Mts., the plains
of Kazakhstan and Middle Asia (excluding the Middle-Asian mountains, i.e. Tien-Shan,
Alai and Pamir).
5.3. East Siberia, from the Enisei River and Altai Mts. in the west and Japan and the Pa-
cific Ocean in the east, including the Russian Far East, Mongolia, northern China and the
Korean Peninsula.
5.4. Middle-Asian mountains comprising the Tien-Shan and Pamiro-Alai.
5.5. Tibetan Plateau.
5.6. Himalaya, including Pakistan, North India, Nepal, Bhutan and Afghanistan.
5.7. South and South-East Asia, including Taiwan, southern and partly central China,
Burma, Thailand, and Vietnam.
5.8. Fossil Myrmica species, found in the European Eocene ambers.
For many years morphometries and indices calculated from them have proved very use-
ful in the taxonomy oîMyrmica (see Chapter 1.5.1) and we make use of various indices in
the Keys. Generally in the Keys we have used indices calculated from simple measurements

711
that can be made quickly and accurately by most people with access to a microscope with
a magnification of at least x 50. These are head-width (HW) and head-length (HL), scape
length (SL), alitrunk length (AL), frons width (FW) and frontal lobe width (FLW), petiole
length (PL) and height (PH), postpetiole width (PPW) and length (ESL) and width
between the tips of the propodeal spines (ESD); from these we calculate the following
indices: CI = HL/HW; FI = FW/HW; FLI = FLW/FW; SIj = SL/HL; SI 2 = SL/HW;
PIj = PL/PH; PPI 4 = PPW/HW; ESLI = ESL/HW; ESDI = ESD/ESL (for full details of
these measurements and indices and others used elsewhere in this monograph and our other
publications see Chapter 1.5.3 and Fig. 333; all figures for extant species see in Chapter
3.3.2, and for fossil species in Chapter 3.4).

Figure 333. Explanation of measuring (A-G-workers; H, I-males). A-head, frontal view; В-scape,
lateral view; C, H - alitrunk, lateral view; D . I - alitrunk, dorsal view; E - waist, lateral view; F - waist,
dorsal view; G - hind tibia. For abbreviations of measurements see Chapter 1.5.3.

712
Map 163. Regions, for which different Keys are compiled. 1 - West Europe and north west of Africa;
2 - East Europe, West Siberia and Altai Mts., plains of Kazakhstan and Middle Asia, Caucasus, Asia
Minor, Iran, and Kopetdagh Mts.; 3 - East Siberia, Russian Far East, Mongolia, northern China, Korean
Peninsula, and Japan; 4 - Middle-Asian mountains; 5 - Tibet; 6 - Himalaya; 7 - South and South-
East Asia.

5.1. A key for the identification of the Myrmica species of West Europe
and North Africa (workers*)

* There are several socially-parasitic Myrmica species found in West Europe and North
Africa that all use other species of Myrmica as their hosts. The queens and males of all the
socially-parasitic species well differ visually from the workers, queens and males of their
hosts. Those species that can produce workers are included in this key. However, most are
workerless, therefore we provide a separate identification key to the queens and males of
the socially parasitic Myrmica species (excluding the facultative social parasite M. vandeli) :
if you suspect a queen or male is a social parasite please use this Key (also see Radchenko
and Elmes 2003a).

1. Frontal carinae curved outwards to merge with the rugae that surround antennal
sockets (e.g. Fig. 213, A). Scape very smoothly curved at the base, never angled and
without any trace of a lobe or carina (e.g. Fig. 213, B) 2
-. Frontal carinae merging with the rugae that extend to the occipital margin, they do
not curve outwards and do not merge with rugae that surround antennal sockets
(e.g. Fig. 239, A). Scape strongly curved, often angled at the base, usually with (but
sometimes without) a lobe, ridge or carina (Figs 13, B, C; 91, B; 99, B, C; 152, B, C;
222, B, C; 230, B-D; 239, B, C; 264, B, C; 273, B, C) 3
2(1). Petiolar node with rounded dorsum, without a flattened dorsal plate, node of
petiole and postpetiole smooth, at most very finely striated; propodeal spines

713
 short, mean ESLI 0.26 (Fig. 213, С, D). Males: scape and tibiae with numerous,
rather long standing hairs (Fig. 215, B, E). - Europe, Caucasus, West Siberia,
northern Kazakhstan, Tien-Shan, Pamir, East Siberia, to the east until Transbaikal
region M. rubra (Linnaeus)
-. Petiolar node flattened, with a dorsal plate, node of petiole and postpetiole with quite
coarse, short sinuous longitudinal rugae; propodeal spines longer, mean ESLI 0.41
(Fig. 216, C, D). Males: scape and tibiae with sparser and short standing hairs
(Fig. 218, B, E). - Transpalaearctic species, distributed from Atlantic to Pacific
Oceans, including Japan, Caucasus, absent in Tien-Shan . . . M. ruginodis Nylander
3(1). Scape with a vertical lobe or dent and sharply angled at the base (e.g. Figs 152, B-D;
204, B-E; 242, B-D; 315, B, C) 4
-, Scape never with a vertical lobe or dent, varying from gradually curved to sharply
angled at the base, often with horizontal lobe, ridge or carina of various shapes
(e.g. Figs 13, B, C; 91, B; 99, B, C; 222, B, C; 230, B-D; 239, B, C; 264, B, C; 273,
B, C) : 13
4(3). Frontal carinae very strongly curved, frons extremely narrow (the narrowest among
all known Myrmica species), FI < 0.19, FLI > 1.95 (Fig. 204, A). Scape at the base
with an extremely massive, vertical, plate-like lobe (Fig. 204, B-E).-Albania, former
Yugoslavia, Greece, Asia Minor, Transcaucasus M. ravasinii Finzi
-. Frontal carinae less curved, frons wider, FI > 0.20, FLI < 1.90 (e.g. Figs 152, A; 242,
A; 315, A). Scape at the base with vertical lobe ranging from moderately sized plate­
like structure to a small dent (e.g. Figs 152, B-D; 242, B-D; 315, B, C) 5
5(4). Foot of scape with extended, shield-like plate (e.g. Figs 242 D; 254, E). Males: scape
short, SIj < 0.50 (e.g. Figs 62, B; 244, B, C) 6
-. Foot of scape without extended shield-like plate (Fig. 152 D). Males: scape long,
SIj > 0.60 (e.g. Figs 154, B; 317, B) 10
6(5). Frontal carinae strongly curved, frons narrower, species' means FI < 0.27 (e.g. Fig.
242, A) 7
-. Frontal carinae less curved, frons wider, species' means FI > 0.28 (e.g. Fig.
60, A) 9
7(6). Surface of foot of scape strongly excavated (seen in profile, Fig. 191, B, D; for
the shape of scape lobe see also Fig. 191, C, E). Alitrunk with fairly coarse, more
or less straight, longitudinal rugae (Fig. 191, F, G). - Greece (Peloponnesus
Peninsula) M. pelops Seifert
-. Surface of foot of scape not excavated (seen in profile, Fig. 242, B; for the shape of
scape lobe see also Figs 175, B-D; 242, C, D). Alitrunk with finer sinuous longitudi­
nal rugae (Figs 175, E, F; 242, E, F) 8
8(7). Frons narrow, mean FI < 0.24, very rarely FI of individuals > 0.25. Males:
scape more strongly curved at the base forming less obtuse angle (mean 121°,
range 101-130°) (Fig. 244, B, C). - Central and North Europe, Caucasus,
West Siberia, northern Kc. ikhstan, Tien-Shan, East Siberia, to Enisei river in
the east M. schencki Viereck
-. Frons distinctly wider, mean FI 0.27, rarely FI of individuals < 0.25. Males: scape less
strongly curved at the base forming more obtuse angle (mean 133°, range 126-144°)
(Fig. 177, B, C). - Italy south of Alps M. obscura Finzi
9(6). Frontal lobes less extended, FLI 1.23-1.36, mean 1.29. Propodeal spines long, ESLI
0.32-0.42, mean 0.38 (Fig. 254, F, G). Distinctive, unusual shape of, scape lobe

74
 see Fig. 254; B-E. Males: scape longer and distinctly curved at the base, subequal
to the length of 1-3 funicular segments together, mean SI2 0.45 (Fig. 256, B, C).
- Sicily M. siciliana Radchenko, Elmes et Alicata
-. Frontal lobes more extended, FLI 1.36-1.52, mean 1.41. Propodeal spines shorter,
ESLI 0.18-0.30, mean 0.24 (Fig. 60, F, G). For the shape of scape lobe see
Fig. 60, B-E. Males: scape very short and not curved at the base, shorter
than 1 st and 2 nd funicular segments together, mean SI2 0.34 (Fig. 62, B).
- Central Europe, Balkans, south of East Europe, Transcaucasus, Iran, Turk-
menistan, south of West Siberia, northern Kazakhstan, Tien-Shan, to Altai Mts. in
the east M. deplanata Emery
10(5). Anterior clypeal margin in female castes and males narrowly rounded, slightly promi-
nent and not-notched medially (Figs 320, A; 321, A; 322, A). Males: scape very long,
SIj > 0.90, SI2 > 0.95. - Spain M. xavieri Radchenko, Elmes et Savolainen
-. Anterior clypeal margin in female castes and males widely rounded, not promi-
nent and notched medially (e.g. Fig. 152, A). Males: scape shorter, SIj < 0.85,
SI2 < 0.90 11
11(10). Petiole with well developed peduncle, its anterior surface not steep, distinctly con-
cave, meeting the dorsal one to form an obtuse, rounded angle, petiolar node with-
out a distinct dorsal plate, its dorsum convex or at most slightly flattened (seen in
profile); propodeal spines longer, mean ESLI 0.35 (Fig. 315, D). Scape at the base
with small, often dentiform lobe (Fig. 315, B, C). Frons wider, mean FI 0.37.
- Iberian Peninsula, French Pyrenees M. wesmaeli Bondroit
-. Petiole with short peduncle, its anterior surface steep, only slightly concave,
meeting the dorsal one at an acute or right angle, petiolar node with a distinct, de-
clined posteriorly, flattened dorsal plate (seen in profile); propodeal spines shorter,
species' means ESLI < 0.30 (e.g. Fig. 152, E). Scape at the base usually with
a distinct lobe (its size may vary, but it is never dentiform) (e.g. Fig. 152, B, C).
Frons narrower, species' means FI < 0.35 12
12(11). Frons relatively narrow, mean FI 0.28. - Boreal species: Europe, Caucasus,
Asia Minor, Siberia, northern Kazakhstan, to the east until Transbaikal
region M. lobicornis Nylander
-. Frons relatively wide, mean FI 0.34. - Iberian Peninsula, Canary Islands, southern
France, Italy, Switzerland, Austria M. lobulicornis Nylander
13(3). Alitrunk and waist with very coarse, straight (not sinuous) longitudinal rugae; peti-
ole with a very short peduncle, a steep anterior surface that meets the dorsal one
at an almost right angle, dorsal plate well developed and flattened (Fig. 273, D, E).
Scape strongly but gradually curved at the base, at most slightly angled, sometimes
with a weak longitudinal ridge on the foot of scape (Fig. 273, B, C). Large and ro-
bust species, usually bicoloured, with reddish alitrunk and blackish head dorsum
and gaster. Males: scape long, SI, > 0.80. - Boreal species, distributed from Ireland
to Kamchatka, absent in Japan M. sulcinodis Nylander
-. Alitrunk, and especially waist, with less coarse, usually sinuous longitudinal ru-
' gosity; petiole having a different shape, scape of various shapes (e.g. Figs 13, B, C;
91; B; 99, B, C; 222, B, C; 230, B-D; 239, B, C; 264, B, C). Males: scape shorter,
SI, < 0.80, usually < 0.70 14
14(13). Clypeus posteriorly with large fovea (Fig. 276, A). - Italy (social parasite)
M. symbiotica (Menozzi)
-. Clypeus without fovea (e.g. Fig. 239, A) 15

715
15(14). Scape gradually curved at the base or at most slightly angled, without an horizon-
tal lobe, at most with a short horizontal ridge or very weak carina (seen under mag-
nification > 5 0 x ; e.g. Figs 13, B, C; 91, B; 222, B, C) 16
-. Scape strongly angled at the base, with an horizontal lobe, carina Orwell developed
ridge (seen under magnification > 5 0 x ; e.g. Figs 99, B, C; Figs 230, B-D; 239, B,
C; 259, B, C; 264, B, C) 20
16(15). Scape at the base bent in an "ideal" curve, with no trace of an angle, carina or ridge
(Fig. 91, B). - Europe, Caucasus, south of West Siberia . . . . M. gallienii Bondroit
-. Scape at the base either slightly angled or with a weak horizontal ridge (e.g. Figs
13, B, C; 222, B, C) 17
17(16). Scape at the base slightly angled, with no trace of a ridge (Fig. 222, B, C).
- Central, North and East Europe (absent in British Isles), Balkans "and Greece
mountains, Caucasus M. rugulosa Nylander
-. Scape at the base with short ridge or weak carina (Figs 13, B, C; 47, B; 55,
B, C) 18
18(17). Small species, mean HW 0.79, mean AL 1.28. Scape at the base slightly angled and
with a very weak ridge (Fig. 55, B, C). Tibial spur well developed and pectinate.
Pronotal dorsum reticulated. Males: middle and hind tibiae with short hairs (Fig. 57,
E). - Ukraine, Poland, southern Finland, Germany, Austria, Switzerland, Czech Re-
public, Romania, Italy, Serbia, Bulgaria and Greece M. constricta Karawajew
-. Larger species, species' means HW 0.97... 1.05, means AL 1.56... 1.64. Scape at the
base only curved, not angled, with a more developed ridge or weak carina (Figs 13,
B, C; 47, B). Tibial spur very often partly reduced, short, simple or only weakly
pectinate (Fig. 13, F). Males: middle and hind tibiae with very long hairs (e.g. Fig.
15, E). . 19
19(18). Only frons with longitudinal rugae, remainder parts of head dorsum with reticula-
tion (Fig. 13, A). Scape at the base with a weak carina (Fig. 13, B, C). Petiolar
node without a distinct dorsal plate (Fig. 13, D). Generally less hairy species.
- Iberian Peninsula, Balearic Isl., French Pyrenees M. aloba Forel
- . Head dorsum mostly with longitudinal rugae, reticulation, if present, only on
occiput (Fig. 47, A). Scape at the base with fine ridge only (Fig. 47, B). Petiolar
node flattened to form a declined posteriorly dorsal plate (Fig. 47, C). Generally
more hairy species. - Morocco M. cagnianti Espadaler
20(15). Frons very narrow, FI < 0.30, mean 0.27. Lobe at the base of scape quite large, but
does not extend to form a longitudinal carina on the dorsal surface of scape (Fig.
259, C). - Central Europe, Balkans, central and southern parts of East Europe,
southern part of West Siberia, northern Kazakhstan M. slovaca Sadil
-. Frons wider, FI > 0.30, species' means > 0.33. Base of the scape with ornamenta-
tion ranging from a ridge or a distinct carina to a large, well developed lobe, but if
the lobe is large, it extends to form a longitudinal carina on the dorsal surface of
scape (e.g. Figs 99, B, C; 230, B-D; 239, B, C; 264, B, C) 21
21 (20). Entire head dorsum with coarse reticulation, only lower part offrons with sinuous
rugae (Fig. 39, A). Males: scape relatively long, longer than 4 basal funicular seg-
ments together, SIj > 0.60 (Fig. 41, B). Head margins with numerous long hairs
(Fig. 41, A). - Switzerland, Germany (social parasite) M. bibikoffi Kutter
-. If present at all, the reticulation on head dorsum is much less extensively
developed (e.g. Figs 230, A; 239, A) 22

716
22(21). Postpetiole very wide, PPI4 > 0.55; whole body with very abundant, long hairs;
frons wide, FI 0.40-0.42 (Fig. 103, A, D, E). Males: scape relatively long, as
long as 4 basal funicular segments together, SIj > 0.50 (Fig. 105, B). Head mar-
gins and waist with numerous long hairs (Fig. 105, A, C, D). - England, Denmark,
Sweden, southern Finland, Netherlands, Germany, Austria, Hungary, southern
Poland, Czech and Slovak Republics, Serbia, European part of Russia (social
parasite) M. hirsuta Elmes
-. Postpetiole narrower, PPI4 < 0.45; body with less abundant and shorter
hairs (e.g. Figs 230, E, F; 239, D, E). Males: scape of various length (e.g. Figs 101,
B;304, B) 23
23(22). Frons wide, mean FI 0.40; frontal lobes moderately extended, mean FLI 1.20 (for
Balkans populations). Scape at the base distinctly angled with a narrow horizon-
tal lobe, carina or even distinct ridge (size of this structure is quite variable) (Fig.
99, B, C). - Caucasus, Asia Minor, Ukraine (Crimea), Bulgaria, Greece, Italy, Croa-
tia, Slovenia and Serbia M. hellenica Finzi
-, Frons generally narrower, species' means FI < 0.38; frontal lobes distinctly more ex-
tended, means FLI > 1.30. Scape at the base with a much more pronounced, larger
horizontal lobe or carina (e.g. Figs 230, B-D; 239, B, C) 24
24(23). Hairy species, petiole with more than 10 (usually with 12-20) long, thin
and often curved hairs. Entire alitrunk dorsum with longitudinal, slightly
sinuous rugae, without reticulation; postpetiolar dorsum with partly re-
duced sculpture (Fig. 302, D, E). Anterior clypeal margin shallowly but dis-
tinctly notched medially (Fig. 302, A). Spurs on middle and hind tibiae at
least partly rduced and usually not pectinate (Fig. 302, F). Queens: petiolar
node with coarse, regular longitudinally-concentric rugosity, without reti-
culation (Fig. 303, D, E). Males: scape relatively long, as long as 4 basal
funicular segments together, SIj > 0.50. Head margins with numerous long hairs
(Fig. 304, A). -West Europe, Ukraine (Transcarpathian Prov.) (facultative social
parasite) M. vandeli Bondroit
-. Generally less hairy species, petiole with less than 10 (usually not more than 8)
long, straight, thick hairs. Alitrunk dorsum with strongly sinuous longitudinal rugae
and often also with reticulation; postpetiolar dorsum with coarser sculpture.
Anterior clypeal margin not-notched medially (e.g. Figs 230, A, E, F; 239, A, D,
E). Spurs on middle and hind tibiae as a rule well developed and pectinate. Queens:
petiolar node with various sculpture, but never with coarse, regular longitudinally-
concentric rugosity (e.g. Figs 231, D, E; 240, C, D). Males: scape shorter; if the
same length (i.e. SIj > 0.50) then head margins with very short and sparse hairs
(e.g. Fig. 232, A) : 25 or 30

• At this point if you have no males or wish to determine species primarily on

worker characters go to couplet 25
- Otherwise, to determine species primarily on male characters go to alternative
key couplet 30

25(24). Scape at the base with very large, massive lobe, which is clearly raised over the
dorsal plane of scape (seen in profile, Fig. 158, B, C). Frons narrow, mean FI 0.32.
Males: scape relatively long, as long as 4-4.5 basal funicular segments together,

717
 SIj > 0.50 (Fig. 160, В). - Central and North Europe, Balkans (from north­
western Italy to Bulgaria), East Europe, Asia Minor, Caucasus, West Siberia and
northern Kazakhstan M. lonae Finzi
-. Scape at the base with smaller and not massive lobe, which is never raised over the
dorsal plane of the scape (seen in profile, e.g. Figs 230, B-D; 239, В, С) Males:
scape of various lengths 26
26(25). Frons relatively narrow, species' means FI < 0.35. Scape at the base with rela­
tively large lobe that is more extended posteriorly than anteriorly (e.g. Fig. 230,
B-D) (either M. sabuleti or M. tulinae which can only be separated with certainty
on male characters) 27
-. Frons relatively wide, species' means FI > 0.36. Scape at the base with smaller lobe
that is more extended anteriorly than posteriorly, or sometimes with only a narrow
carina (e.g. Figs 239, В, С; 264, В, С) 28
27(26). Workers: spur of hind and middle tibiae well developed, pectinate. Males: scape
relatively long, as long as 4-4.5 basal funicular segments together, SIj > 0.50
(Fig. 232, B). - Europe (to southern Sweden, Finland and Norway in the north),
Caucasus M. sabuletiMeinert
-. Workers: spur of hind and middle tibiae often reduced to some extent. Males:
scape short, shorter than 1-3 basal funicular segments together, SIj < 0.40 (Fig.
297, B). -Turkey, Georgia, France, Italy, Netherlands, Poland
M. tulinae Elmes, Radchenko et Aktac
28(26). Propodeal spines relatively short, mean ESLI 0.36; petiolar node without a dis­
tinct horizontal dorsal plate, its posterior surface declines gradually to postpetiole
(seen in profile, Fig. 264, D). Frons relatively wide, mean FI 0.38; frontal lobes
less extended, mean FLI 1.32. Males: scape short, SI1 < 0.45. Middle and hind
tibiae and basitarsi with short hairs, they are shorter than maximum width of tibia
(Fig. 266, E). - Europe (to southern England and Denmark in the north), Cauca­
sus, Asia Minor, Iran, Turkmenistan, south of West Siberia, northern Kazakhstan,
to Altai Mts. in the east M. specioides Bondroit
-. Propodeal spines longer, species' means ESLI > 0.40; petiole usually with a dis­
tinct horizontal or slightly declined posteriorly dorsal plate, its posterior surface
falls abruptly to postpetiole, often appearing as a distinct step (seen in profile, e.g.
Fig. 239, D). Frons somewhat narrower, species' means FI 0.36 ... 0.37; frontal
lobes variously extended. Males: scape of various length. Middle and hind tibiae
and basitarsi with longer hairs, they are longer than maximum width of tibia (Figs
241, E; 269, E) 29
29(28). Workers: frontal lobes more extended, mean FLI 1.42. Males: scape short, as long
as 3-3.5 basal funicular segments together, SIj < 0.45 (Fig. 241, B). - Europe,
Caucasus, Asia Minor, northern Kazakhstan, West Siberia; the easternmost
confirmed locality - vicinity of Krasnoyarsk (ca. 93°E) and 100 km N of Kansk
(ca. 96°E) M. scabrinodis Nylander
-. Workers: frontal lobes less extended, mean FLI 1.33. Males: scape relatively long,
as long as 4-4.5 basal funicular segments together, SIj > 0.50 (Fig. 269, B).
- South-west of Mediterranean Region (Iberian Peninsula, southern France,
Italy) M. spinosior Santschi

718
• Altenative key based primarily on male characters

30(24). Males: scape relatively long, as long as 4-4.5 basal funicular segments together,
SI, > 0.50 (e.g. Fig. 232, B) (M. lonae, M. sabuleti, M. spinosior, these species can
only be separated with certainty on worker characters) 31
-. Males: scape short, as long as 3-3.5 basal funicular segments together, SI, < 0.45
(e.g. Fig. 241, B) . . 33
31(30). Workers: scape at the base with very large, massive lobe, which is clearly
raised'over the dorsal plane of scape (seen in profile, Fig. 158, B, C). Frons
narrow, mean FI 0.32. - Central and North Europe, Balkans (from north-western
Italy to Bulgaria), East Europe, Asia Minor, Caucasus, West Siberia and northern
Kazakhstan : , . . . . . M. lonae Finzi
-. Workers: scape at the base with smaller and not massive lobe, which is never
raised over the dorsal plane of scape (seen in profile, e.g. Figs 230, B-D; 267,
B, C) 32
32(31). Workers: frons narrow, mean FI 0.33. - Europe (to southern Sweden, Finland and
Norway in the north), Caucasus M. sabuleti Meinert
-. Workers: frons wider, mean FI 0.37. - South-west of Mediterranean Region
(Iberian Peninsula, southern France, Italy) M. spinosior Santschi
33(30). Males: middle and hind tibiae and tarsi with short hairs, they are shorter than
maximum width of tibia (Fig. 266, E). Workers: frons relatively wide, mean
FI 0.38; frontal lobes less extended, mean FLI 1.32. Scape at the base usually with
• narrow lobe or even carina (Fig. 264, B, C). Propodeal spines shorter, mean ESLI
" 0.36; petiolar node without a distinct horizontal dorsal plate, its posterior surface
declines gradually to postpetiole (seen in profile, Fig. 264, D). - Europe (to
southern England and Denmark in the north), Caucasus, Asia Minor, Iran, Turk-
menistan, south of West Siberia and northern Kazakhstan, to the Altai Mts. in the
east , M. specioides Bondroit
-. Males: middle and hind tibiae and tarsi with very long erect, often curved
hairs, they are distinctly longer than maximum width of tibia (e.g. Fig. 241, E).
Workers: frons narrower, species' means FI 0.34 ... 0.36; frontal lobes more
extended, species' means FLI > 1.40. Scape at the base usually with more
developed, sometimes large lobe (Figs 239, B, C, 295, B, C). Propodeal spines
longer, means ESLI > 0.40; petiole usually with a distinct horizontal or slightly de-
clined posteriorly dorsal plate, its posterior surface falls abruptly to postpetiole,
often appearing as a distinct step (seen in profile, e.g. Fig. 239, D) (M. tulinae and
M. scabrinodis, these species can only be separated with certainty on worker
characters) 34
34(33). Workers: frons relatively narrow, mean FI 0.34. Scape at the base with relati-
vely large lobe (Fig. 295, B, C). - Turkey, Georgia, France, Italy, Netherlands,
Poland M. tulinae Elmes, Radchenko et Aktaç
-. Workers: frons wider, mean FI 0.36. Scape at the base usually with much smaller
lobe, or even with carina (Fig. 239, B, C). - Europe, Caucasus, Asia Minor, northern
Kazakhstan, West Siberia; the easternmost confirmed locality - vicinity of Krasno-
yarsk (ca. 93°E) and 100 km N of Kansk (ca. 96°E) M. scabrinodis Nylander

71-9
A key for the identification of the socially-parasitic Myrmica species
of West Europe and North Africa (queens)
1. Scape at the base strongly angled or only curved, but always with a horizontal lobe
or at least well developed carina (Figs 40, B, C; 104, B; 148, B, C) 2
-. Scape at the base only curved, always without lobe or carina (e.g. Fig. 126, B) . . . . 4
2(1). Scape at the base gradually curved, with narrow horizontal carina (Fig. 148, B, C).
Eyes with conspicuous hairs, length of the longest hairs 0.035 mm (Fig. 148, A).
- Italy M. laurae (Emery)
-. Scape at the base strongly angled, with more developed horizontal lobe (Figs 40, B,
G; 104, B). Eyes usually without hairs, or if present, they are inconspicuous and very
short, length of the longest hairs < 0.01 mm (Figs 40, A; 104, A) 3
3(2). Large species (HW 1.22-1.24 mm). Frons quite narrow (FI 0.34-0.35). Only frons
with longitudinal sinuous rugae, remainder part of head dorsum with coarse reticu-
lation (Fig. 40, A). - Switzerland, Germany M. bibikoffi Kutter
-. Smaller species (HW 0.80-1.10 mm). Frons wider (FI 0.39-0.46). Head dorsum
mainly with longitudinal, slightly sinuous rugae, only its rear third with reticula-
tion (Fig. 104, A). - England, Denmark, Sweden, southern Finland, Nether-
lands, Germany, Austria, Hungary, southern Poland, Czech and Slovak Republics,
Serbia M. hirsuta Elmes
4(1). Scape relatively short, SI] < 0.76, SI2 < 0.70. Petiolar node (seen in profile) sub-
triangular (Fig. 171, C). - Switzerland M. myrmicoxena Forel
-. Scape longer, SIj > 0.80, SI2 > 0.85. Petiolar node (seen in profile) with broadly
rounded dorsum (e.g. Fig. 126, C) 5
5(4). The basal half of the first gastral tergite with conspicuous, fairly long, suberect
hairs; postpetiolar dorsum with distinct reticulation (Fig. 150, C, D). - France,
Spain M. lemasnei Bernard
-, First gastral tergite with only sparse, short, decumbent pubescence; postpetiolar
dorsum without reticulation (e.g. Fig. 126, C, D) 6
6(5). Surface of head, alitrunk and waist finely but densely punctated, appears dull; peti-
olar and postpetiolar nodes with relatively short, straight or slightly downward-
curved hairs (Fig. 121, A, C, D). -Algeria M. kabylica (Cagniant)
-. Surface of head, alitrunk and waist smooth or at most superficially punctated, ap-
pears shiny; petiolar and postpetiolar nodes with fairly long, downward-curved hairs
(Fig. 126, A, C, D). - East, Central and North Europe M. karavajevi (Arnoldi)

A key for the identification of the socially-parasitic Myrmica species

of West Europe and North Africa (males)
1. Antennae 12-segmented 2
-. Antennae 13-segmented 5
2(1). Eyes with conspicuous hairs, length of the longest hairs 0.04 mm (Fig. 149, A).
Propodeum with pointed teeth (Fig. 149, C, D). - Italy M. laurae (Emery)
-. Eyes usually with no hairs, or if present, they are inconspicuous and very short, length
of the longest hairs < 0.01 mm (e.g. Fig. 127, A). Propodeum with short blunt denti-
cles (e.g.'Fig. 127, C, D) 3

720
3 (2). Surface of alitrunk (especially of scutum and scutellum) finely but densely punctated,
appearing dull (Fig. 122, C, D). - Algeria M. kabylica (Cagniant)
-. Surface of alitrunk not punctated, smooth and appearing shiny (Figs 127, C, D; 151,
C, D) 4
4(3). First gastral tergite with only sparse short decumbent pubescence (Fig. 127, C, D).
- East, Central and North Europe . . M. karavajevi (Arnoldi)
-. Basal half of first gastral tergite with conspicuous, fairly long suberect hairs (Fig. 151,
C, D). - France, Spain M. lemasnei Bernard
5(1). Head margins with short hairs; antennal scape short (SIj = SI 2 = 0.35) (Fig. 171, H).
Switzerland . . . M. myrmicoxena Forel
-. Head margins with long hairs; antennal scape longer (SIj > 0.40, SI2 > 0.45) (Figs
41, A, B; 105, A, B) '....- 6
6(5). Lateral and posterior parts of head dorsum with distinct reticulation (Fig. 41, A)
- Switzerland, Germany M. bibikoffi Kutter
-. Lateral and posterior parts of head dorsum without reticulation, at most with longi-
tudinal striation or sometimes only punctated (Fig. 105, A). - England, Denmark,
Sweden, southern Finland, Netherlands, Germany, Austria, Hungary, southern
Poland, Czech and Slovak Republics, Serbia M. hirsuta Elmes

5.2. A key for the identification of the Myrmica species of East Europe,
West Siberia, northern Kazakhstan, Caucasus, Asia Minor, Turkmenistan
and Iran (workers*)
* Does not include M. karavajevi Arnoldi which is a worker-less social parasite living
in nests of several Myrmica species. Its queens have well developed ventral process on the
petiole (Fig. 126, C), males have 12-segmented antennae, both queens and males are small:
in queens HW < 0.90, AL < 1.40 mm, in males HW < 0.75, AL < 1.30 mm. On forewing
vein 2+3RS completely reduced and cells 1 +2r and rm are fused (Fig! 4, G in Chapter 2.1)
(see also the Key for the socially parasitic species of West Europe).
1. Lateral portion of clypeus raised into a sharp ridge in front of antennal insertions, so
that antennal sockets are distinctly separated from clypeal surface (similar to that of
Tetramorium) (Figs 279, A; 290 A) 2
-. Lateral portion of clypeus not raised into a sharp ridge in front of antennal insertions,
so that antennal sockets lay on the same level with clypeal surface (e.g. Figs 60, A;
213, A; 230, A; 273, A) 3
2(1). Head dorsum with coarse rugosity, frons between frontal carinae level with the eyes
with < 10 rugae (Fig. 279, A). Metanotal groove very shallow to almost absent (Fig.
279, C). - Asia Minor M. tamarae Elmes, Radchenko et Aktac
-. Head dorsum with finer rugosity, frons between frontal carinae level with the eyes
with > 13 rugae (Fig. 290, A). Metanotal groove distinct (Fig. 290, D). - South Siberia
(including Altai Mts.), southern part of Russian Far East, north-eastern China,
Korean Peninsula, Japan (Hokkaido) M. transsibirica Radchenko
3(1). Frontal carinae curve outwards to merge with the rugae that surround antennal
sockets (e.g. Fig. 213, A). Scape very smoothly curved at the base, not angled and
without any trace of lobe or carina (e.g. Fig. 213, B) 4

721
 -. Frontal carinae merging with the rugae that extend to the occipital margin, they do
not curve outwards and do not merge with rugae that surround antennal sockets
(e.g. Figs 60, A; 230, A; 273, A). Scape strongly curved, often angled at the base, with
or without lobe, ridge or carina (e.g. Figs 60, B-E; 91, B; 99, B, C; 152, B-D; 204,
B-E; 222, B, C; 230, B-D; 239, B, C; 242, B-D; 264, B, C; 273, B, C) 5
4(3). Petiolar node with rounded dorsum, without a flattened dorsal plate, node of peti-
ole and postpetiole smooth, at most very finely striated; propodeal spines short,
mean ESLI 0.26 (Fig. 213, C, D). Males: scape and tibiae with numerous, rather
long standing hairs (Fig. 215, B, E) - Europe, Caucasus, West Siberia, northern
Kazakhstan, Tien-Shan, Pamir, East Siberia, to the east until Transbaikal
region M. rubra (Linnaeus)
-. Petiolar node flattened, with a dorsal plate, node of petiole and postpetiole with quite
coarse, short sinuous, longitudinal rugae, propodeal spines longer, mean ESLI 0.41.
(Fig. 216, C, D). Males: scape and tibiae with sparser and short standing hairs (Fig.
218, B, E). - Transpalaearctic species, distributed from Atlantic to Pacific Oceans,
including Japan, Caucasus, absent in Tien-Shan M. ruginodis Nylander
5(3). Scape with a vertical lobe or dent and sharply angled at the base (e.g. Figs 60, B-E;
152, B-D; 204, B-E; 242, B-D) 6
-. Scape never with vertical lobe or dent, varying from gradually curved to sharply
angled at the base, often with horizontal lobe, ridge or carina of various shapes (e.g.
Figs 91, B; 99, B, C; 222, B, C; 230, B-D; 239, B, C; 264, B, C; 273, B, C) 14
6(5). Frontal carinae very strongly curved, frons extremely narrow (the narrowest among
all known Myrmica species), FI < 0.19, FLI > 1.95 (Fig. 204, A). Scape at the base
with an extremely massive, vertical plate-like lobe (Fig. 204, B-E). - Albania, former
Yugoslavia, Greece, Asia Minor, Transcaucasus M. ravasinii Finzi
-. Frontal carinae less curved, frons wider, FI > 0.20, FLI < 1.90 (e.g. Figs 60, A; 152,
A; 242, A). Scape at the base with vertical lobe ranging from moderately sized plate-
like structure to a small dent (e.g. Figs 60, B-E; 152, B-D; 242, B-D) 7
7(6). Head and alitrunk with strongly reduced sculpture, only with fine striation, surface
between striae smooth and appears shiny, only petiole and postpetiole finely punc-
tated (Fig. 112, A, F, G). Whole body appears shiny black. - Kazakhstan
M. inucta Radchenko et Elmes
-. Head and alitrunk with well developed, often quite coarse rugosity, surface
between rugae does not appear smooth and shiny (e.g. Figs 60, A, F, G; 242, A, E,
F). Body colour various, but never entirely black 8
8(7). Foot of scape with an extended shield-like plate (e.g. Figs 60, E; 242, D). Males:
scape short, SIj < 0.50 9
-. Foot of scape without an extended shield-like plate (e.g. Fig. 152, D). Males: scape
long, SI, > 0.60 12
9(8). Frontal carinae strongly curved, frons narrower, species' means FI < 0.26 (e.g. Fig.
242, A) 10
-. Frontal carinae less curved, frons wider, species' means FI > 0.29 (Figs 60, A;
134, A) 11
10(9). Scape at the base with well developed, large lobe (Fig. 242, B-D). Frontal carinae
strongly curved, frons narrower, mean FI < 0.24. Males: scape, hind and middle
tibiae with short hairs (Fig. 244, B, C, F). - Central and North Europe, Caucasus,
West Siberia, northern Kazakhstan, Tien-Shan, East Siberia, to Eniser river in
the east M. schencki Viereck

722
 -..Scape at the base with small, dentiform lobe, (Fig. 49, B-D). Frontal carinae
less curved, frons wider, FI 0.25-0.29, mean 0.26. Males: scape, hind and mid­
dle tibiae with very long hairs (Fig. 51, B, E). - Transcaucasus
M. caucasicola Arnoldi
11(9). Frontal lobes less extended, FLI 1.36-1.52, mean 1.41 (Fig. 60, A). Propodeal
spines short, ESLI 0.18-0.30, mean 0.24 (Fig. 60, F, G). Body colour usually dark
reddish-brown. Males: scape very short, shorter than 1st and 2nd funicular segments
together, mean SI2 0.34 (Fig. 62, B). - Central Europe, Balkans, south of East
Europe, Transcaucasus, Iran, Turkmenistan, south of West Siberia, northern
Kazakhstan, Tien-Shan, to Altai Mts. in the east M. deplanata Emery
-. Frontal lobes more extended, FLI 1.50-1.67, mean 1.60 (Fig. 134, A). Propodeal
spines longer,' ESLI 0.27-0.34, mean 0.31 (Fig. 134, F, G). Alitrunk yellowish-
brown, head reddish-brown, gaster dark brown. Males: scape longer than
I st and 2 nd funicular segments together, mean SI2 0.40 (Fig. 136, B, C). - West
Siberia from Irtysh river in the west, Tarbagatai and Saur Mts., South and East
Siberia, Mongolia, southern part of Russian Far East, northern China, Korean
Peninsula M. koreana Elmes, Radchenko et Kim
12(8). Frons relatively wide, FI > 0.35. Scape at the base with small vertical dentiform
lobe or even small dentiform ridge (Fig. 17, В, С). Petiole with well developed pe­
duncle, its anterior surface not steep, distinctly concave, meeting the dorsal one to
form an obtuse, rounded angle, petiolar node without a distinct dorsal plate, with
a convex or at most slightly flattened dorsum (seen in profile, Fig. 17, D).
- Asia Minor M. anatolica Elmes, Radchenko et Aktac
-. Frons narrower, FI < 0.34. Scape at the base with a vertical (sometimes inclined
anteriorly), usually large lobe, which sometimes can be smaller but never denti­
form (Figs 152, B-D; 325, В, С). Petiole of various shape 13
13(12). Petiole with a short peduncle, its anterior surface steep, only slightly concave,
meeting the dorsal one at an acute or right angle, petiolar node with a distinct, de­
clined posteriorly, flattened dorsal plate (seen in profile, Fig. 152, E). Lobe at the
base of the scape projecting almost vertically (Fig. 152, B-D). - Boreal species:
Europe, Caucasus, Asia Minor, Siberia, northern Kazakhstan, to the east until
Transbaikal region M. lobicornis Nylander
-, Petiole with a well developed peduncle, its anterior surface not steep, distinctly
concave, meeting the dorsal one to form an obtuse, rounded angle, petiolar node
without a distinct dorsal plate and with a convex dorsum (seen in profile, Fig. 325,
D). Lobe at the base of the scape inclined anteriorly (Fig. 325, B, C). - Altai Mts.
and Southern Ural M. zojae Radchenko
14(5). Alitrunk and waist with very coarse, straight (not sinuous) longitudinal rugae; peti­
ole with a very short peduncle, a steep anterior surface that meets the dorsal one
at an almost right angle, dorsal plate well developed and flattened (Fig. 273, D, E).
Scape strongly but gradually curved at the base, at most slightly angled, sometimes
with a weak longitudinal ridge on the foot of scape (Fig. 273, B, C). Large and ro­
bust species, usually bicoloured, with reddish alitrunk and blackish head dorsum
and gaster. - Boreal species, distributed from Ireland to Kamchatka, absent in
Japan M. sulcinodis Nylander
-. Alitrunk, and especially waist, with less coarse, usually sinuous longitudinal ru­
gosity; petiole having a different shape, scape of various shapes (e.g. Figs 55, B-D;
91, B-D; 99, B-D; 158, B-D; 222, B-D; 230, B-E; 239, B-D; 264, B-D) 15

723
 15(14). Scape gradually curved at the base or at most slightly angled, without horizontal
lobe, at most with very weak short horizontal ridge (seen under magnification
> 50x; e.g. Figs 55, B, C; 91, B; 222, B, C) 16
-. Scape strongly angled at the base, with horizontal lobe, carina or well developed
ridge (seen under magnification > 50X; e.g. Figs 158, B, C; 230, B-D; 239, B, C;
264, B, C) 20
16(15). Frons extremely wide, FI > 0.50. Males: scape long, SIt > 0.75. - Asia
Minor M.jennyae Elmes, Radchenko et Aktac
-. Frons narrower, FI < 0.50. Males: scape shorter, SIj < 0.65 17
17(16). Scape at the base bent in an "ideal" curve, with no trace of an angle, carina or
ridge (e.g. Fig. 91, B). Head dorsum with longitudinal rugosity, reticulation, if
present, developed on its lateral parts above the eyes and/or near occipital margin
(e.g. Fig. 91, A). Pronotal dorsum longitudinally rugose (e.g. Fig. 91, D) . . . . 18
-. Scape at the base either slightly angled or with a weak horizontal ridge (e.g. Fig.
222, B, C). Posterior third of head dorsum, as well as pronotal dorsum, reticulate
(e.g. Fig. 222, A, E) 19
18(17). Propodeal spines short, mean ESLI 0.28, petiole low, mean PIl 1.24 (Fig. 36, C,
D). Generally bigger, mean HW 1.12, mean AL 1.80, more robust and darker
coloured species. - Delta of Danube riv., southern parts of Ukraine and European
Russia, southeastern Transcaucasus, northeastern Iran, Kazakhstan, plains and
mountains of Middle Asia M. bergi Ruzsky
-. Propodeal spines longer, mean ESLI 0.36, petiole higher, mean PIj 1.14 (Fig. 91,
C, D). Generally smaller, mean HW 1.04, mean AL 1.62, more slender and
lighter coloured species. - Europe, Caucasus, south of West Siberia
M.gallieniiBondroit
19(17). Scape at the base with no trace of a ridge (Fig. 222, B, C). - Central, North
and East Europe (absent in British Isles), Balkans and Greece mountains,
Caucasus M. rugulosa Nylander
-. Scape at the base with weak ridge (Fig. 55, B, C). - Ukraine, Poland, southern
Finland, Germany, Austria, Switzerland, Czech Republic, Romania, Italy, Serbia,
Bulgaria and Greece M. constricta Karawajew
20(15). Frons very narrow, FI < 0.30, mean 0.27. Lobe at the base of scape quite large, but
not extended to form a longitudinal carina on the dorsal surface of scape (Fig. 259,
C). - Central Europe, Balkans, central and southern parts of East Europe,
southern part of West Siberia, northern Kazakhstan M. slovaca Sadil
-. Frons wider, FI > 0.30, species' means > 0.33. Base of the scape with ornamenta­
tion ranging from a ridge or a distinct carina to a large, well developed lobe, but if
the lobe is large, it extends to form a longitudinal carina on the dorsal surface of
scape (e.g. Figs 158, B, C; 230, B-D; 239, B, C; 264, B, C) 21
21(20). Postpetiole very wide, PPI4 > 0.55; whole body with very abundant, long hairs;
frons wide, FI 0.40-0.42 (Fig. 103, A, D, E). Males: scape relatively long, as long
as 4 basal funicular segments together, SII > 0.50 (Fig. 105, В). Head margins
and waist with numerous long hairs (Fig. 105, A, C, D). - England, Denmark,
Sweden, southern Finland, Netherlands, Germany, Austria, Hungary, southern
Poland, Czech and Slovak Republics, Serbia, European part of Russia (social
parasite) M. hirsuta Elmes
-. Postpetiole narrower, PPI4 < 0.45; body with less abundant and shorter hairs

724
 (e.g. Figs 230, E, F; 239, D, E). Males: scape of various length (e.g. Figs 101, В;
304, В) 22
22(21). Hairy species, petiole with more than 10 (usually with 12-20) long, thin and often
curved hairs. Entire alitrunk dorsum with longitudinal, slightly sinuous rugae,
without reticulation; postpetiolar dorsum with partly reduced sculpture (Fig. 302,
D, E). Anterior clypeal margin shallowly but distinctly notched medially (Fig. 302,
A). Spurs on middle and hind tibiae at least partly reduced and usually not pecti­
nate (Fig. 302, F). Queens: petiolar node with coarse, regular longitudinally-
concentric rugosity, without reticulation (Fig. 303, D, E). Males: scape relatively
long, as long as 4 basal funicular segments together, SIX > 0.50 (Fig. 304, B). Head
margins with numerous long hairs (Fig. 304, A). - West Europe, Ukraine (Trans-
carpathian Prov.) (facultative social parasite) M. vandeli Bondroit
-. Generally less hairy species, petiole with less than 10 (usually not more than 8)
long, straight, thick hairs. Alitrunk dorsum with strongly sinuous longitudinal
rugae and often also with reticulation; postpetiolar dorsum with coarser sculpture
(e.g. Figs 230, E, F; 239, D, E). Anterior clypeal margin not-notched medially (e.g.
Figs 230, A; 239, A). Spurs on middle and hind tibiae as a rule well developed and
pectinate. Queens: petiolar node with various sculpture, but never with coarse,
regular longitudinally-concentric rugosity (e.g. Figs 231, D, E; 240, C, D). Males:
scape shorter; if the same length (i.e. SI, > 0.50) then head margins with very
short and sparse hairs (e.g. Fig. 232, A) 23 or 34

• At this point if you have no males or wish to determine species primarily on

worker characters go to couplet 23
- Otherwise, to determine species primarily on male characters go to alternative
key couplet 34

23(22). Scape at the base with very large, massive lobe, which is clearly raised over the
dorsal plane of scape (seen in profile) (Fig. 158, В, С). Frons narrow, mean
FI 0.32. Males: scape relatively long, as long as 4-4.5 basal funicular segments to­
gether, SI, > 0.50 (Fig. 160, B). - Central and North Europe, Balkans (from north­
western Italy to Bulgaria), East Europe, Asia Minor, Caucasus, West Siberia and
northern Kazakhstan M. lonae Finzi
-. Scape at the base with smaller and not massive lobe, which is never raised over the
dorsal plane of the scape (seen in profile, e.g. Figs 230, B-D; 239, В, С). Males:
scape of various lengths 24
24(23). More'hairy species, head margins with very abundant, relatively long but subde-
cumbent hairs (Fig. 33, A); scape with abundant suberect hairs (Fig. 33, В, С);
hairs on the body quite coarse and whitish, coarser and more abundant than in
related species. Frons relatively wide, mean FI 0.39, frontal lobes moderately
extended, mean FLI 1.34. Males: scape somewhat longer than 1-3 basal funicular
segments together, mean SIX 0.44 (Fig. 35, B). Tibiae and tarsi on both anterior and
posterior surfaces with'long suberect to erect hairs; the longest hairs on tibiae dis­
tinctly longer than their maximal width, those on the basitarsus more than twice
longer than its maximal width (Fig. 35, E). - Georgia . . . M. bakurianica Arnoldi

725
 -. Less hairy species, head margins with less abundant, usually suberect hairs (e.g.
Fig. 230, A); scape with much less abundant suberect hairs (e.g. Fig. 230, B-D);
hairs on the body not coarse. Frontal carinae and frontal lobes of various shape.
Males: scape of various length, character of pilosity on tibiae and tarsi different
(e.g. Figs 241, E; 266, E) 25
25(24). Frons relatively narrow, species' means FI < 0.35. Scape at the base with rela-
tively large lobe that is more extended posteriorly than anteriorly (e.g. Fig. 230,
B-D) 26
-. Frons relatively wide, species' means FI > 0.36. Scape at the base with smaller
lobe that is more extended anteriorly than posteriorly, sometimes with narrow
carina only (e.g. Fig. 239, B, C) 29
26(25). Frons somewhat wider, mean FI 0.35, frontal lobes less extended, mean FLI 1.50
(Fig. 140, A). Males: scape somewhat shorter than 1-3 basal funicular seg-
ments together, mean Sll 0.42. Standing hairs on tibiae and tarsi shorter, longest
hairs on tibiae not longer than the tibial width, same on tarsi less than twice
longer than tarsal width (Fig. 142, E). - Steppes from southern Ukraine till
Kazakhstan M. kozakorum sp. nov.
-. Frons narrower, species' means FI 0.33...034, frontal lobes more extended, means
FLI 1.53-1.57 (e.g. Fig. 230, A). Males: scape short (SIj < 0.40) or long (SIa >
0.50). Standing hairs on tibiae and tarsi very long, longest hairs on tibiae distinctly
longer than the tibial width, same on tarsi more than twice longer than tarsal width
(Figs 232, E; 297, E) (males of M. georgica are unknown) 27
27(26). Workers: propodeal spines shorter, mean ESLI 0.38. - Georgia
M. georgica Seifert
-. Workers: propodeal spines longer, mean ESLI 0.42 : 28
28(27). Workers: spur of hind and middle tibiae well developed, pectinate. Males: scape
relatively long, as long as 4-4.5 basal funicular segments together, SIj > 0.50
(Fig. 232, B). - Europe (to southern Sweden, Finland and Norway in the north),
Caucasus M. sabuleti Meinert
-. Workers: spur of hind and middle tibiae often reduced in different extent. Males:
scape short; shorter than 1-3 basal funicular segments together, SIj < 0.40 (Fig.
297, B). -Turkey, Georgia, France, Italy, Netherlands, Poland
M. tulinae Elmes, Radchenko et Aktac
29(25) .Frons wide, mean FI 0.43; frontal lobes weakly extended, mean FLI 1.15 (for Turk-
ish, Crimean and Caucasian populations) (Fig. 99, A). Scape at the base distinctly
angled with a narrow horizontal lobe, carina or even distinct ridge (size of this struc-
ture is quite variable) (Fig. 99, B, C). - Caucasus, Asia Minor, Ukraine (Crimea),
Bulgaria, Greece, Italy, Croatia, Slovenia and Serbia M. hellenica Finzi
-. Frons generally narrower, species' means FI < 0.40; frontal lobes distinctly more
extended, species' means FLI > 1.30. Scape at the base with a much more pro-
nounced, larger horizontal lobe or carina (e.g. Figs 239, B, C; 264, B, C) . . . . 30
30(29). Frons wider, mean FI 0.38 frontal lobes less extended, mean FLI 1.32 (Fig. 264,
A), propodeal spines relatively short, mean ESLI 0.36; petiolar node without
a distinct horizontal dorsal plate, its posterior surface declines gradually to
postpetiole (Fig. 264, D). Males: middle and hind tibiae and tarsi with short
hairs, they are shorter than maximum width of tibia (Fig. 266, E). - Europe (to

726
 southern England and Denmark in the north), Caucasus, Asia Minor,
Iran, Turkenistan,.south of West Siberia, northern Kazakhstan, to Aitai Mts. in
the east M. specioides Bondroit
-. Frons narrower, species' mean FI 0.36, shape of frontal lobes, petiole and
propodeum various (Figs 233, A, D; 239, A, D; 270, A, D; 298, A, D) 31
31(30). Frontal lobes less extended, mean FLI 1.31. Scape at the base with narrow hori-
zontal ridge or at most very small carina (Fig. 270, B, C)..Propodeal spines short
(mean ESLI 0.31), not widened at the base, thin, often needle-like; metanotal
groove weak or completely absent; petiolar node without dorsal plate, usually only
rounded, its posterior surface gradually declines to postpetiole; sides of alitrunk
with relatively coarse, regular, almost straight longitudinal rugae (Fig. 270, D, E).
Body colour rather dark, brownish-red. Males: scape very short, mean SIt < 0.33,
tibiae and tarsi with short subdecumbent hairs (Fig. 272, B, E). - Steppes from
the lower reaches of the Dnepr River to the eastern Kazakhstan, to the north -
until Samara Province of Russia M. stangeana Ruzsky
-. Frontal lobes more extended, species' means FLI > 1.40. Scape at the base with
• more developed, but never massive, horizontal carinaor lobe (Figs 233, B, C; 239,
B, C; 298, B, C). Propodeal spines longer (species1 means ESLI > 0.35), usually
widened at the base (more thorn-like) never needle-like; metanotal groove well
developed, often deep; petiolar node of various shape; sides of alitrunk with less
coarse sinuous longitudinal rugae (Figs 233, D; 239, D; 298, D). Body colour
lighter, usually ochreous- or yellowish-red. Males: scape somewhat longer, species'
means SIj > 0.36, often > 0.40). Tibiae and tarsi with various hairs (Figs 235, F;
241, E; 300, E) 32
32(31). Propodeal spines longer, mean ESLI > 1.40; petiolar node with a distinct hori-
zontal or slightly declined posteriorly dorsal plate (Fig. 239, D). Males: middle and
hind tibiae and tarsi with very long erect, often curved hairs, they are distinctly
longer than maximum width of tibia (Fig. 241, E). - Europe, Caucasus, Asia Minor,
northern Kazakhstan, West Siberia; the easternmost confirmed localities -
vicinity of Krasnoyarsk (ca. 93°E) and 100 km N of Kansk (ca. 96°E)
M. scabrinodis Nylander
-. Propodeal spines shorter, species' means ESLI < 1.36; petiolar node dorsum with
or without dorsal plate (Figs. 233, D; 298, D). Males: middle and hind tibiae and
tarsi with much shorter hairs, they are not longer than maximum width of tibia
(Figs 235, F; 300, E) 33
33(32). Frontal lobes less extended, FLI 1.40. Petiolar node with a distinct horizontal
dorsal plate (Fig. 233, D). - south of West Siberia and northern Kazakhstan
. . . M. salina Ruzsky
'-. Frontal lobes more extended, mean FLI 1.47. Petiolar node without a distinct
horizontal dorsal plate, its posterior surface declines gradually to postpetiole
(Fig. 298, D). - Asia Minor M. turcica Santschi

727
• Alternative key based primarily on male characters
(males of M. georgica are unknown)

34(22). Males: scape relatively long, as long as 4-4.5 basal funicular segments together,
SI, > 0.50 (e.g. Fig. 232, B) (M. sabuleti and M. lonae; these species can only be
separated with certainty on worker characters) 35
-. Males: scape short, as long as 3-3.5 basal funicular segments together or less,
SI, < 0.45 (e.g. Figs 241, B; 266, B) 36
35(34). Workers: scape at the base with very large, massive lobe, which is clearly raised
over the dorsal plane of scape (seen in profile) (Fig. 158, B, C). - Central and
North Europe, Balkans (from north-western Italy to Bulgaria), East Europe, Asia
Minor, Caucasus, West Siberia and northern Kazakhstan M. lonae Finzi
-. Workers: scape at the base with smaller lobe, which is never raised over the dor-
sal plane of the scape (seen in profile) (Fig. 230, B-D). - Europe (to southern Swe-
den, Finland and Norway in the north), Caucasus M. sabuleti Meinert
36(34). Males: middle and hind tibiae and tarsi on both anterior and posterior surfaces
with very long erect, often curved hairs, the longest hairs on tibiae distinctly longer
than maximum width of tibia, those on the basitarsus more than twice as long as
its maximal width (Figs 35, E; 241, E; 297, E) (M. bakurianica, M. tulinae and
M. scabrinodis; these species can only be separated with certainty on worker
characters) 37
-. Males: middle and hind tibiae and tarsi with much shorter hairs, the longest hairs
on tibiae not longer than the tibial width, same on basitarsus less than twice as long
as its width (e.g. Figs 266, E; 300, E) 39
37(36). Workers: more hairy species, head margins with very abundant, relatively long but
subdecumbent hairs (Fig. 33, A); scape with abundant suberect hairs (Fig. 33, B,
C); hairs on the body quite coarse and whitish, coarser and more abundant than
in related species. Frons relatively wide, mean FI 0.39, frontal lobes moderately
extended, mean FLI 1.34 - Georgia M. bakurianica Arnoldi
-. Workers: less hairy species, head margins with less abundant, usually suberect
hairs (e.g. Fig. 239, A); scape with much less abundant suberect hairs (e.g. Fig.
239, B, C); hairs on the body not coarse. Frons narrower, species' means FLI
0.34 ... 36, frontal lobes more extended, species' means FLI > 1.40 38
38(37). Workers: frons narrower, mean FI 0.34, frontal lobes more extended, mean FLI
1.53. Scape at the base with relatively large lobe that is more extended posteriorly
than anteriorly (Fig. 295, B, C). Spur of hind and middle tibiae often reduced in
different extent. - Turkey, Georgia, France, Italy, Netherlands, Poland
M. tulinae Elmes, Radchenko et Aktac
-. Workers: frons wider, mean FI 0.36, frontal lobes less extended, mean FLI 1.42.
Scape at the base with smaller lobe that is more extended anteriorly than posteri-
orly, (Fig. 239, B, C). Spur of hind and middle tibiae well developed and pecti-
nate. - Europe, Caucasus, Asia Minor, northern Kazakhstan, West Siberia; the
easternmost confirmed localities - vicinity of Krasnoyarsk (ca. 93°E) and 100 km
N of Kansk (ca. 96°E) M. scabrinodis Nylander
39(36). Males: scape very short, mean SI, 0.36. - Steppes from the lower reaches of
the Dnepr River to the eastern Kazakhstan, to the north - until Samara Province
of Russia M. stangeana Ruzsky
-. Males: scape longer, species' means SI, > 0.40 40

728
40(39). Males: petiole rather massive, shorter and higher, PIa 1.33 (Fig. 235, D); post-
petiole relatively wider, PPI4 0.61. - South of West Siberia and northern Kaza-
khstan M. salina Ruzsky
-. Males: petiole somewhat elongate, not massive, species' means PIj > 1.40 (e.g.
• Fig. 266, C); postpetiole relatively narrower, species' means PPI4 < 0.60: (M. spe-
cioides, M. turcica, M. hellenica and M. kozakorum; it is not possible to separate
the males of these species, however the geographical location can indicate which
is most probable):
- Europe (to southern England and Denmark in the north), Caucasus, Asia Minor,
Iran, Turkmenistan, south of West Siberia, northern Kazakhstan, to Altai Mts. in
the east M. specioides Bondroit
- Asia Minor M. turcica Santschi
- Caucasus, Asia Minor, Ukraine (Crimea), Bulgaria, Greece, Italy, Croatia,
Slovenia and Serbia M. hellenica Finzi
- Steppes from southern Ukraine to Kazakhstan M. kozakorum sp. nov.

5.3. A key for the identification of the Myrmica species of East Siberia,
Russian Far East, Mongolia, Korean Peninsula, northern China, and Japan
(workers)
1. Lateral portion of clypeus raised into sharp ridge in.front of antennal insertions, so
that antennal sockets are distinctly separated from clypeal surface (similar to that of
Tetramorium) (e.g. Fig. 80, A) 2
-. Lateral portion of clypeus not raised into sharp ridge in front of antennal insertions,
so that antennal sockets lay on the same level with clypeal surface (e.g. Figs 134, A;
' 161, A; 213, A) 3
2(1). Scape at the bend having the same width as at its mid-length (seen in profile, Fig. 80,
B); the foot of scape having a longitudinal groove and lateral ridges (seen anteriorly,
Fig. 80, C). Males: petiole and postpetiole with fine superficial microsculpture,
appearing dull. - Southern part of Russian Far East, north-eastern China, Korean
Peninsula, Japan M. excelsa Kupyanskaya
-. Scape at the bend is distinctly narrower than at its mid-length (seen in profile, Fig.
290, B); the foot of scape without longitudinal groove and lateral ridges (seen ante-
riorly, Fig. 290, C). Males: petiole and postpetiole smooth, appearing shiny. - South
Siberia (including Altai Mts.), southern part of Russian Far East, north-eastern
China, Korean Peninsula, Japan (Hokkaido) M. transsibirica Radchenko
3(1). Frontal carinae curved outwards to merge with the rugae that surround the antennal
sockets (e.g. Figs 161, A; 213, A). Scape very smoothly curved at the base, not an-
gled and without any trace of lobe or carina (e.g. Figs 161, B; 213, B) 4
-. Frontal carinae merging with the rugae that extend to the occipital margin, they do
not curve outwards and do not merge with rugae that surround the antennal sockets
(e.g. Fig. 134, A). Scape strongly curved at the base, with or without a lobe, ridge or
carina (e.g. Figs 5, B; 22, B; 66, B; 123, B; 128, B; 134, B-E; 195, B; 239, B, C; 273,
B, C; 325, B, C) 7
4(3). First gastral tergite finely but distinctly longitudinally striated at the base. Much more
hairy species; petiolar node with ca. 10 long and numerous shorter standing hairs
(Fig. 161, C, D). Frons with numerous, fine longitudinal rugulae, numbering > 20

729
 between the frontal carinae level with the eyes (Fig. 161, A). Spurs on tibiae of
middle and hind legs reduced, simple (not pectinate) (Fig. 161, E, F), Scape quite
strongly curved at the base (Fig. 161, B). Queens no bigger than the workers, their
propodeum without spines, at most with bunt tubercles (Fig. 162, C, D). Males: scape
short, Sl 2 < 0.50. - Southern part of Russian Far East, north-eastern China,' Korean
Peninsula, Japan M. luteola Kupyanskaya
-. First gastral tergite completely smooth. Less hairy species, petiolar node with 6-8
long standing hairs (e.g. Fig. 213, C, D). Frons with fewer relatively coarse, longitu-
dinal rugae, numbering < 15 between the frontal carinae level with the eyes (e.g. Fig.
213, A). Spurs on tibiae of the middle and hind legs well developed and pectinate.
Scape gently curved at the base (e.g. Fig. 213, B). Queens distinctly larger than the
workers, their propodeum with spines. Males: scape long, SI2 > 0.70 5
5 (4). Petiolar node with rounded dorsum, completely without a dorsal plate, node of petiole
and postpetiole smooth, at most very finely striated; propodeal spines short, mean ESLI
0.26 (Fig. 213, C, D). - Europe, Caucasus, West Siberia, northern Kazakhstan, Tien-
Shan, Pamir, East Siberia, to the east till Transbaikal region . . . M. rubra (Linnaeus)
-. Petiolar node with a dorsal plate developed to various extents, node of petiole and
postpetiole with coarser sculpture, rugulose or rugose; propodeal spines longer,
species' means ESLI 0.41 (Figs 137, C, D; 216, C, D) 6
6(5). Petiolar node with a distinct, sharp flattened dorsal plate and having quite coarse
short sinuous longitudinal rugae on lateral parts (Fig. 216, C). Males: clypeus usually
smooth, at most with very fine superficial punctures (Fig. 218, A). -Transpalaearctic
species, distributed from Atlantic to Pacific Oceans, including Japan, Caucasus, ab-
sent in Tien-Shan M. ruginodis Nylander
-. Petiolar, node dorsum slightly convex, with somewhat convex, not sharp dorsal plate,
laterally with finer short longitudinal rugulae (Fig. 137, C). Males: clypeus with fine
longitudinal striation (Fig. 139, A). - Southern part of Russian Far East, north-
eastern China, Korean Peninsula, Japan M. kotokuiForei
7(3). Scape sharply angled at the base, with horizontal lobe (Fig. 239, B, C).
- Europe, Caucasus, Asia Minor, northern Kazakhstan, West Siberia; the eastern-
most confirmed localities - vicinity of Krasnoyarsk (ca. 93°E) and 100 km N of Kansk
(ca. 96°E) M. scabrinodis Nylander
-. Scape of various shapes, but never with horizontal lobe (e.g. Figs 5, B; 22, B; 66, B;
. 123, B; 128, B; 134, B-E; 195, B; 273, B, C; 325, B, C) 8
8(7). Scape sharply angled at the base, with a vertical lobe (that can be inclined anteriorly)
or with dent (e.g. Figs 134, B-E; 195, B; 325, B, C) 9
-. Scape gradually curved or angled at the base, but never with a vertical or inclined
lobe or dent (Figs 5, B; 22, B; 66, B; 123, B; 128, B; 273, B, C) 17
9(8). Propodeal spines thick and blunt at their apex, directed backwards and upwards, and
strongly curved inward; petiole with very short peduncle, its anterior surface steep
and meets the dorsal one at an acute angle, so that petiolar node appears sharply an-
gled (seen in profile) (Fig. 84, C, D). - East and South Siberia, to the west till Saiany
Mts., northern Mongolia; in Russian Far East is absent . . . M. forcipata Karawajew
-, Propodeal spines thin and sharp at their apex, not curved inward; petiole of
various shapes (seen in profile) but if it is sharply angled, the frons is narrower
(FI < 0.32 7JS > 0.35 in M. forcipata) (e.g. Figs 195, C, D; 242, E) 10

730
 10(9). Lobe at the base of scape forms shield-like plate along the surface of the foot of the
scape (e.g. Figs 134, E; 178, E; 242, D). Males: scape short, Sl1 < 0.50 11
-. Lobe at the base of scape does not form shield-like plate along surface of the foot
of the scape (e.g. Fig. 152, D). Males: scape long, SIj > 0.60 13
11(10). Frontal carinae feebly curved, frons wider, FI > 0.33 (mean 0.35), and impor­
tantly the frontal lobes are not extended, FLI < 1.25 (Fig. 178, A). Scape with
a small vertical dent (Fig. 178, B-E). - Japan
M. onoyamai Radchenko et Elmes
-. Frontal carinae strongly curved, frons narrower, FI < 0.33 (species' means
< 0.30), and the frontal lobes are distinctly extended, FLI > 1.50 (e.g. Fig. 134, A).
Scape with larger, more distinct, not dentiform, vertical lobe (Figs 134, B-E; 242,
B-D) .. . , 12
12(11). Frontal carinae strongly curved, frons quite narrow, FI 0.20-0.27, mean 0.23.
Scape lobe quite large and metanotal groove deeper (Fig. 242, B-E). - Europe,
Caucasus, West Siberia, northern Kazakhstan, Tien-Shan, East Siberia, to the east
till Enisei river M. schencki Viereck
-. Frontal carinae less curved, frons wider, FI 0.27-0.32, mean 0.29. Scape lobe
smaller and metanotal groove shallower (Fig. 134, B-F). - West Siberia, to the
west till Irtysh river, Tarbagatai and Saur Mts., South and East Siberia, Mongolia,
southern part of Russian Far East, northern China, Korean Peninsula
M. koreana Elmes, Radchenko et Kim
13(10). Petiole with a short peduncle, its anterior surface (seen in profile) is steep,
' only slightly concave, meeting the dorsal one to form an acute or right angle; dor­
sal plate distinct, declined posteriorly (Fig. 152, E). Frons narrower and frontal
lobes more extended, mean FI 0.28, mean FLI 1.43. - Boreal species: Europe,
Caucasus, Asia Minor, Siberia, northern Kazakhstan, to the east till Transbaikal
region M. lobicornis Nylander
-. Petiole of other shapes (e.g. Figs 116, С; 195, С; 325, D). Frons wider and frontal
lobes less extended, species' means FI > 0.32, means FLI < 1.40 14
14(13). Anterior surface of petiole (seen in profile) weakly concave and meeting the dor­
sal one to form a slightly rounded obtuse angle, node dorsum slightly convex, quite
long and gradually sloping posteriorly, forming regular low arch together with
the posterior surface; propodeal spines not widened at the base, slightly curved
down and directed mainly backward, subhorizontal (Fig. 195, С). - South Siberia,
Mongolia . . . . M. pisarskii Radchenko
-. Anterior surface of petiole (seen in profile) distinctly concave, node dorsum with
convex plate or widely rounded, posterior surface steep; propodeal spines widened
at the base, straight, directed backward and upward, not subhorizontal (e.g. Figs
' 116, C; 325, D) . 15
15(14). Scape at the base with a large lobe that is inclined anteriorly (Fig. 325, B, C).
- Altai Mts. and Southern Ural ...' .. . M. zojae Radchenko
-. Scape at the base with small, almost vertical, lobe or dent (Figs 75, B, C;
116, B) ' 16
16(15). Frontal carinae strongly curved, frons narrower, and frontal lobes more extended:
FI 0.31-0.34, mean 0.32, FLI 1.29-1.42, mean 1.35. - Korean Peninsula,
Japan M. jessensis Forel

731
 -. Frontal carinae less curved, frons wider, and frontal lobes less extended:
FI 0.32-0.39, mean 0.36, FLI 1.18-1.40, mean 1.29. - South Siberia (from
Buryatia in the west), Mongolia, Russian Far East, northern China, Korean
Peninsula M. eidmanni Menozzi
17(8). Mesonotum and propodeum strongly constricted laterally (seen from above), their
dorsal surface narrow and delineated laterally by sharp ridges, which merge with
the outer bases of the propodeal spines (Fig. 54, D). - South and East Siberia,
Mongolia M. commarginata Ruzsky
-. Mesonotum and propodeum not constricted laterally, their dorsal surface not
delineated by ridges (i.e. of "normal" shape) (e.g. Figs. 28, D; 273, D) 18
18(17). Petiole with a distinct lobe-like subtriangular ventral process, its peduncle very
short, anterior surface almost straight and steep, node narrowly rounded so that
in profile, the petiole appears to be subtriangular (Fig. 28, C). Scape short, mean
SIj 0.70. Spur on the tibiae of hind and middle legs fairly reduced, short, often
not pectinate (Fig. 28, E, F). Males: antennae with 12 segments, scape short
SIj < 0.40. - South Siberia from Tuva to Transbaikalia, Mongolia
M. arnoldii Dlussky
-. Petiole of various shapes, but never with lobe-like ventral process (e.g. Figs 5, D;
66, C; 128, C; 145, C; 273, D). Scape longer, species' means SI, > 0.75. Spur on
hind and middle tibiae well developed and pectinate. Males: antennae with 13 seg-
ments, antennal scape of various length 19
19(18). Scape gradually curved at the base or at most very slightly angled, usually without
a ridge on the inner margin of its foot (e.g. Figs 5, B; 66, B; 128, B); if in some spec-
imens such a ridge is present (Fig. 273, C) then the alitrunk and waist have very
coarse, straight (not sinuous) longitudinal rugae, the petiole has a relatively short
peduncle and its anterior surface is almost straight, meeting the dorsal one at
a right angle, also the dorsal plate is developed, flattened, not inclined posteriorly
(see M. sulcinodis, below) 20
-. Scape distinctly angled at the base, always with ridge on the inner margin of
its foot (e.g. Figs 22, B; 123, B); if in some specimens the scape is not distinctly
angled then the petiole has a very short peduncle and its anterior surface meets
the dorsal one at an acute angle, so that petiolar node appears sharply angled
(Fig. 22, C) 25
20(19). Alitrunk with very coarse, straight (not sinuous) longitudinal rugae; propodeal
spines long, often longer than the length of the propodeal dorsum, species' means
ESLI > 0.40; petiole with a short peduncle, seen in profile its anterior surface is
almost straight, meeting the dorsal one almost at a right or somewhat obtuse angle,
dorsal plate well developed, flattened, not inclined posteriorly (Figs 5, D, E; 273,
D, E). Scape strongly curved at the base, occasionally slightly angled (Figs 5, B;
273, B) 21
-. Alitrunk with less coarse sinuous longitudinal rugosity (except for M. kasczenkoi);
propodeal spines shorter, species1 means ESLI < 0.35; petiole of various shapes,
but never with a well developed, flattened dorsal plate (e.g. Figs 66, C, D; 128, C,
D). Scape gradually, often smoothly, curved at the base, never angled (e.g. Figs 66,
B; 128, B) 22
21(20). Sides of petiolar node with coarse rugae that are very similar to those on the
alitrunk; anterior surface of petiole very steep, meeting the dorsal one almost at
a right angle; propodeal spines somewhat shorter than the length of the propodeal

732
 dorsum, mean ESLI 0.40 (Fig. 273, D). - Boreal species, distributed from Ireland
to Kamchatka, absent in Japan M. sulcinodis Nylander
-. Sides of petiolar node with coarse punctures and short rugae, which are much less
coarse than those on the alitrunk; anterior surface of petiole less steep, meeting the
dorsal one at a somewhat obtuse angle; propodeal spines very long, equal or longer
than the length of the propodeal dorsum, mean ESLI 0.44 (Fig. 5, D). - Southern
part of Russian Far East, Korean Peninsula M. ademonia Bolton
22(20). Sides of alitrunk with quite coarse, almost straight longitudinal rugae, metanotal
groove very weak or absent; petiole and postpetiole with longitudinal rugosity only,
not punctated; anterior surface of petiole weakly concave and meets with dorsal
one at a slightly rounded obtuse angle, node dorsum slightly convex, quite long and
gradually sloping posteriorly, forming with posterior surface regular low arch (Fig.
128, C, D). -Altai Mts., South Siberia, southern and southeastern Transbaikalia,
Mongolia M. kasczenkoi Ruzsky
-. Alitrunk with less coarse sinuous longitudinal rugosity, metanotal groove distinct,
often deep (except for M. kurokii); petiole and postpetiole with fine, short rugu-
lae or striae, surface between them punctated; anterior surface of petiole dis­
tinctly concave, node dorsum rounded, posterior surface steep (e.g. Fig 66,
C, D) 23
23(22). Bigger species: HW > 1.15, AL > 1.90. Surface of head dorsum between rugae
densely punctated, appearing dull (Fig. 145, A). Propodeal spines longer, mean
ESLI 0.36 (Fig. 145, C). - Southern part of Russian Far East, northern China,
Korean Peninsula, Japan M. kurokii Forel
-. Smaller species: HW < 1.10, AL < 1.80. Surface of head dorsum between rugae
at most very finely superficially micropunctated, appearing shiny (e.g. Fig 66, A).
Propodeal spines shorter, species' means ESLI < 0.30 (e.g. Fig 66, C) 24
24(23). Anterior clypeal margin simply convex, not-notched medially. Males: scape
short, SIj < 0.40. - South and East Siberia (to the west - until Altai Mts.),
Mongolia M. divergens Karawajew
-. Anterior clypeal margin shallowly notched medially. Males: scape long, SIX > 0.70.
- Kamchatka, Magadan Prov, of Russia M. displicentia Bolton
25(19). Petiole with very short peduncle, mean PIj 1.03, its anterior surface steep, very
weakly concave, meeting the dorsal one at a sharp acute angle, dorsal plate flat,
well developed, strongly inclined backward; propodeal spines usually curved in­
ward (Fig. 22, C, D). - Siberia, to the west till Kuznetsky Alatau, Russian Far East,
Mongolia, north-eastern China, Korean Peninsula M. angulinodis Ruzsky
-. Petiole longer, with well developed peduncle, species' means РI1 1.20 ... 1.21, its
anterior surface not steep, distinctly concave, meeting the dorsal one to form
a slightly rounded acute or obtuse angle, dorsal plate not developed, node dorsum
rounded or somewhat convex and steeply sloping backward; propodeal spines
straight, never curved inward (Figs 123, C, D; 293, C, D) 26
26(25). Anterior surface of petiole meeting the dorsal one to form slightly rounded
. obtuse angle, node dorsum widely rounded (Fig. 123, C). Males: antennae
with 13 segments. - South and East Siberia, Mongolia, Russian Far East, North
Korea M. kamtschatica Kupyanskaya
-. Anterior surface of petiole meeting the dorsal one to form a rounded acute angle,
node dorsum somewhat convex and steeply sloping backwards (Fig. 293, C).
Males: antennae with 12 segments. - East Siberia. . . M. tschekanovskii Radchenko

733
5.4. A key for the identification of the Myrmica species of the Middle Asian
Mountains (workers)

1. Scape sharply angled at the base and with a horizontal or vertical lobe, or at least
a small vertical dent (e.g. Figs 60, B-E; 242, B-D; 287, B, C) 2
-. Scape gradually curved at the base, or at most with a slightly marked very
obtuse angle, and never with any form of lobe or dent (e.g. Figs 36, B; 213, B;
273, B) 6
2(1). Scape with a horizontal lobe (Fig. 287, B, C). - Middle Asian Mts
M. tobiasi Radchenko et Elmes
-. Scape with vertical lobe or at least small dent (e.g. Fig. 236, B, C; 242, B-D) 3
3 (2). Lobe at the base of scape forms shield-like plate along the surface of the foot of scape
(Figs 60, E; 242, D). Petiole in profile with a distinctly concave anterior face, its node
with a flattened or somewhat convex dorsal plate (Figs 60, F; 242, E). Males: scape
short, SI, < 0.50 4
-. Lobe at the base of scape does not form shield-like plate along the foot of scape (as
in Fig. 152, D). Petiole in profile with an almost straight, steep anterior face, its node
with a posteriorly-inclined dorsal plate, appearing subtriangular (Figs 131, D; 236,
D). Males: scape long, SI1 > 0.70 5
4(3). Frontal carinae strongly curved, frons narrower, mean FI 0.23. Scape at the base with
large shield-like lobe (Fig. 242, B-D). Alitrunk with deeper metanotal groove,
propodeal spines longer, mean ESLI 0.33 (Fig. 242, E). - Europe, Caucasus,
West Siberia, northern Kazakhstan, Tien-Shan, East Siberia, to the east till Enisei
river M. schencki Viereck
-. Frontal carinae less curved, frons wider, mean FI 0.31. Scape at the base with small
lobe or even dent (Fig. 60, B-E). Alitrunk with shallow metanotal groove, propodeal
spines shorter, mean ESLI 0.24 (Fig. 60, F). - Central Europe, south of East Europe,
Transcaucasus, Iran, Turkmenistan, south of West Siberia, northern Kazakhstan,
Tien-Shan, to the east - till Altai Mts M. deplanata Emery
5(3). Scape at the base with small but distinct lobe (Fig. 131, B, C). Frontal lobes some-
what more extended, mean FLI 1.37. - Middle Asian Mts
M. kirghisorum Arnoldi
-. Scape at the base with small dent or even ridge (Fig. 236, B, C). Frontal lobes less
extended, mean FLI 1.25. - Middle Asian Mts M. saposhnikovi Ruzsky
6(1). Frontal carinae strongly curved at their anterior third, frontal lobes strongly extended,
wide and subsquare, FLI > 1.25 (e.g. Fig. 281, A) 7
-. Frontal carinae feebly curved along whole length, frontal lobes not extended, rela-
tively narrow, FLI < 1.18 (e.g. Figs 36, A; 213, A) 8
7(6). Propodeal spines not broadened at the base, straight, needle-like (Fig. 183, C). Males:
scape extremely short, SI, < 0.35. - Middle Asian Mts. . . . . . M. orthostyla Arnoldi
-. Propodeal spines somewhat broadened at the base, slightly curved (Fig. 281, C).
Males: scape somewhat longer, SI, > 0.36. - Middle Asian Mts., north-eastern
Afghanistan M. tenuispina Ruzsky
8(6). Frontal carinae curved outwards to merge with the rugae that surround antennal
sockets (Fig. 213, A). -Europe, Caucasus, West Siberia, northern Kazakhstan, Tien-
Shan, Pamir, East Siberia, to the east until Transbaikal region
M. rubra Linnaeus
-. Frontal carinae merging with the rugae that extend to the occipital margin or at least

734
 to the level of upper eyes margin, they do not curve outwards and do not merge with
rugae that surround antennal sockets (e.g. Fig. 36, A) 9
9(8). Alitrunk and waist with very coarse, almost straight (not sinuous) longitudinal
rugae; petiole with very short peduncle, its anterior face steep, meets with dorsal
surface at an almost right angle, dorsal plate well developed, flattened (Fig. 273, D,
E). Scape strongly but gradually curved at the base, at most slightly angled, some-
times with weak longitudinal ridge on the foot of scape (Fig. 273, B, C). Large and
robust species, usually bicoloured, with reddish alitrunk and blackish head dorsum
and gaster. - Boreal species, distributed from Ireland to Kamchatka, absent
in Japan M. sulcinodis Nylander
-. Alitrunk, and especially waist, with much finer sinuous longitudinal rugae; petiole
with distinct peduncle and concave anterior face (e.g. Fig. 36, C, D). Scape at the
base gradually curved (e.g. Fig. 36, B) 10
10(9). Scape curved at the base to form quite a strong bend (Fig. 36, B). More
robust, darker coloured and generally bigger species (HW 0.95-1.30, AL 1.50-
2.10 mm). Males: scape short, SIj < 0.60. - Steppe Zone from the delta of
Danube River to Altai Mts., Transcaucasus, north-eastern Iran, plains and moun-
tains of Middle Asia M. bergi Ruzsky
-. Scape very smoothly curved at the base (e:g. Fig. 72, B). More slender, lighter
coloured and generally smaller species (HW 0.75-1.05, AL 1.25-1.65 mm).
Males: scape long, SI1 > 0.80 11
11(10). Antennal sockets not surrounded by concentric rugulae, at most with a very
fine interrupted striation (Fig. 83, A). - Middle Asian Mts
M. ferganensis Karawajew
-. Antennal sockets surrounded by distinct fine, concentric rugulae (e.g. Fig.
72, A .12
12(11). Petiole higher (mean PIj 1.32), with a shorter peduncle, its node (seen in profile)
with.a slightly convex dorsal plate that appears horizontal, or slightly inclined back-
wards (Fig. 72, C). Males: petiole higher, massive, mean PI : 1.33 (Fig. 74, C). -
Middle Asian Mts M. dshungarica Ruzsky
-. Petiole lower (species' means PIj 1.38...1.40), with longer peduncle, its node
without a dorsal plate, being rather narrowly rounded (e.g. Fig. 119, C). Males:
petiole lower, slender, mean PIX 1.53 (males of M. kryzhanovskii are unknown)
(Fig. 120, C) ,..,... 13
13(12). Scape with long suberect hairs, length of the longest hairs not shorter than maxi-
mum diameter of scape (Fig. 144, B). - Middle Asian Mts .'
M. kryzhanovskii Arnoldi
-, Scape with shorter subdecumbent hairs, length of the longest hairs distinctly shorter
than maximum diameter of scape (e.g. Fig. 119, B) M. juglandeti Arnoldi

5.5. A key for the identification of the Myrmica species of the Tibetan
Plateau (workers)
1. Frontal carinae strongly curved at their anterior third, frontal lobes strongly extended,
wide and subsquare, FLI > 1.20 (Fig. 284, A). - Tibet M. tibetana Mayr

735
 -. Frontal carinae feebly curved along whole length, frontal lobes not extended, rela-
tively narrow, FLI < 1.15 (e.g. Fig. 31, A) 2
2(1). Big and robust species, HW > 1.10, AL > 1.75 mm. Petiole with very short pedun-
cle (mean Plj 1.11), its anterior face steep, straight or at most very slightly concave
(Fig. 143, C). -Tibet, Himalaya M. kozlovi Ruzsky
-. Smaller and more slender species, HW < 0.95, AL < 1.55 mm. Petiole with very dis-
tinct peduncle (species' means Plj 1.29...1.32), its anterior face distinctly concave
(Figs 31, C; 228, C) 3
3(2). Scape longer (SI2 > 0.93), with more abundant suberect hairs. - Tibet
M. bactriana Ruzsky
-, Scape somewhat shorter (SI2 < 0.91), with less abundant subdecumbent hairs.
. - Tibet M. ruzskyana Radchenko et Elmes

5.6. A key for the identification of the Myrmica species of the Himalaya
and Afghanistan (workers*)

* M. ereptrix and M. petita are known only from the holotype queens
- M. ereptrix (India: Kashmir) (Fig. 79) is a socially-parasitic species and well differs from
any other species of this region by its very wide petiole and postpetiole that have con-
spicuous, well developed, large ventral lobes.
- M. petita (India: Kashmir) (Fig. 192) is unusually small (HW 0.82, AL 1.58 mm), smaller
than all other known Myrmica queens from the Himalayan region. It is assumed to be
a free living species because it has none of the normal parasitic morphological characters,
in which case it is probably the smallest queen known for any free-living, Old World
Myrmica species.

1. Basal third of first gastral tergite distinctly punctato-striated (Fig. 43, D). - NE
Pakistan, Nepal M. brancuccii Radchenko et Elmes
-. Gastral tergites smooth and shiny 2
2(1). Whole alitrunk dorsum, or at least part of pronotal dorsum, with transversal rugae
(Figs 110, C, D; 186, C, D; 305, C, D; 306, C, D) 3
-. Alitrunk dorsum never with transversal rugae (e.g. Figs 42, C, D; 44, C, D; 143, C,
D; 172, C, D; 207, C, D; 210, C, D; 219, C, D; 281, C, D; 301, D; 308, C, D) . . . . 6
3(2). Only'pronotal and propodeal dorsum partly with transversal rugae, other part of
alitrunk dorsum with longitudinal rugosity and reticulation (Fig. 305, C,'D). - India:
Kashmir M. varisculpta Radchenko et Rigato
-. Whole alitrunk dorsum with transversal rugae (sculpture on propodeal dorsum may
be reduced) (Figs 110, C, D; 186, C, D; 306, C, D) '.'.' 4
4(3). Lateral margins of head with short decumbent hairs, a few long suberect hairs may
be present only on the occipital margin and genae (Fig. 186, A). - India: Kashmir,
Nepal, Bhutan M. pachei Forel
-. Head margins with long, abundant standing hairs (Figs 110, A; 306, A) .5
5(4). Whole alitrunk with coarse rugosity, petiolar node and postpetiole with coarse
transversally-concentric rugae; petiole with very thin and long peduncle,
Plj > 1.60 (Fig. 110, C, D). Body colour blackish-brown to black. - India: Himahal
Pradesh M. inezae Forel

736
 -. Alitrunk with fine rugulosity, sculpture on propodeal dorsum often reduced;
petiole with much thicker and shorter peduncle, PI, < 1.50 (Fig. 306, C, D).
Colour lighter, head dorsum dark reddish brown, alitrunk and gaster brownish
red. - Bhutan M. villosa Radchenko et Elmes
6(2). Anterior clypeal margin shallowly but distinctly notched medially (e.g. Fig. 52, A).
Propodeal lobes pointed apically (except forM. alperti) (e.g. Fig. 52, C) 7
-. Anterior clypeal margin rounded or pointed, never notched medially (e.g. Figs 219,
A). Propodeal lobes rounded apically (e.g. Fig. 219, C) 13
7(6). Head and alitrunk very coarsely rugose, surface between rugae on head dorsum
smooth (e.g. Figs 301, A, C, D; 312, A, C, D) 8
-. Head and alitrunk much more finely rugulose or even striated, surface between rugu-
lae on head dorsum distinctly punctated (e.g. Figs 42, A, C, D; 52, A, C, D) 11
8(7). Lateral margins of head above the eyes with numerous standing hairs (Fig. 301, A).
- India: Meghalaya M. urbanii Radchenko et Elmes
-. Lateral margins of heads above the eyes without or at most with a few standing
hairs (e.g. Figs 107, A; 312, A) 9
9(8). Smaller species: mean HW 0.92, mean AL 1.68. Posterior half of head dorsum
(above the level of eyes) with coarse reticulation, remainder part of head dorsum
longitudinally rugose (Fig. 312, A). Propodeal spines very slender, not widened at
their base (Fig. 312, C, D). Head and alitrunk black to blackish-brown, gaster dark
brown, while appendages are distinctly lighter, brownish-yellow, contrasting with
dark colour of body. - India: West Bengal (Darjeeling), Nepal, Bhutan
M. weberi Elmes et Radchenko.
-. Bigger species: species' means HW > 1.10, means AL > 1.90. Head dorsum mostly
longitudinally rugose, reticulation present only on occiput and temples (Figs 16,
A; 107, A). Propodeal spines more robust, somewhat widened at their base (Figs
16, C, D; 107, C, D). Body reddish-brown to black, appendages somewhat lighter,
but not contrasting with the colour of body 10
10(9). Frontal carinae curved outwards to merge with the ruga, which surround anten-
nal sockets (Fig. 16, A). Petiolar node dorsum reticulated (Fig. 16, D). Propodeal
lobes rounded apically (Fig. 16, C). - Nepal M. alperti Elmes et Radchenko
-. Frontal carinae not curved outwards and do not merge with the ruga, which sur-
round antennal sockets (Fig. 107, A). Petiolar node dorsum rugose (Fig. 107, D).
Propodeal lobes pointed apically (Fig. 107, C). - India: West Bengal (Darjeeling),
Nepal, Bhutan M. indica Weber
11(7). Surface between rugulae on head dorsum finely punctated, appears more or less
shiny; alitrunk dorsum with irregular striae; petiolar node and postpetiole very
finely superficially punctated or even smooth, appears shiny (Fig. 52, A, C, D).
- Bhutan M. collingwoodi Radchenko et Elmes
-. Surface between rugulae on head dorsum densely and coarsely punctated, appears
dull; alitrunk dorsum finely reticulated; petiolar node and postpetiole densely
punctated, submat (Fig. 42, A, C, D; 165, A, C, D) . 12
12(11). Propodeal spines very long, ESLI 0.52-0.58, directed mainly backwards and slightly
curved downwards, subparallel to dorsal surface of propodeum, and faintly diverg-
ing, ESDI < 1.10 (Fig. 42, C, D). - Nepal M. boltoni Radchenko et Elmes
-. Propodeal spines relatively shorter, ESLI 0.44-0.47, straight, directed upwards
at an angle of about 45°, and distinctly diverging, ESDI > 1.25 (Fig. 165, C, D).
- Nepal M. martensi Radchenko et Elmes

737
 13(6). Frontal carinae strongly curved in their anterior third, frontal lobes strongly
extended, FLI > 1.30 (Fig. 281, A). - Tajikistan, north-eastern Afghanistan . . . .
M. tenuispina Forel
-. Frontal carinae more or less straight, frontal lobes not extended, FLI < 1.15 (e.g.
Fig. 219, A) 14
14(13). Promesonotal dorsum coarsely reticulate, sides of pronotum with slightly sinuous
longitudinal rugulae, surface of both very finely punctated; remainder of alitrunk,
petiole and postpetiole very densely punctated, have no rugae (only fine stria-
tion presents on the lower parts of mesopleura and sides of propodeum). Pro­
podeal spines very short, dentiform, but sharp, ESLI 0.18 (Fig. 318, C, D).
- India: Kashmir M. williamsi Radchenko et Elmes
-. Whole alitrunk rugulose, rugose or reticulate, punctated or not, or sculpture
reduced and alitrunk at least partly appears smooth and shiny; propodeum
and mesopleura never only densely punctated. Propodeal spines of different shape
(e.g. Figs 44, С, D; 143, С, D; 172, С, D; 207, С, D; 210, С, D; 219, С, D; 308,
С, D) 15
15(14). Body sculpture strongly reduced, alitrunk, petiole and postpetiole appear smooth
and shiny, at most partly very finely superficially striated (Fig. 172, C, D). Body
colour yellowish-red. - India: Kashmir M. nitida Radchenko et Elmes
-. Alitrunk, petiole and postpetiole differently sculptured, but never appear smooth
and shiny (e.g. Figs 44, C, D; 143, C, D; 207, C, D; 210, C, D; 219, C, D; 308, C,
D); if sculpture partly reduced, body colour dark reddish brown 16
16(15). Petiole with very thin and long peduncle, PI t 1.65. Propodeal spines very
long, ESLI 0.52, and thin (Fig. 210, C, D). - north-eastern Pakistan
M. rigatoi Radchenko et Elmes
-. Petiole with shorter peduncle, Pl, < 1.60. Propodeal spines shorter, ESLI < 0.45,
and stouter (e.g. Figs 44, C, D; 143, C, D; 207, C, D; 219, C, D; 308, C, D) . . . 17
17(16). Antennal sockets not surrounded by rugae. Head and alitrunk with coarse, almost
straight rugae, without reticulation, surface between rugae appears smooth and
shiny; petiolar and postpetiolar nodes concentrically rugose. Petiole with long
peduncle and sharply concave anterior surface, mean PIj 1.43 (Fig. 308, A, C, D)
- north-eastern Pakistan M. vittata Radchenko et Elmes
-. Antennal sockets always surrounded by concentric rugae or at least by striae (e.g.
Fig. 219, A). Body with various sculpture, but never with only coarse longitudinal
rugae. Petiole with shorter peduncle, means PIj < 1.40 (e.g. Figs 44, C, D; 143, C,
D; 207, C, D; 219, C, D 18
18(17).Large and robust species, HW > 1.20, AL > 2.0 mm, head relatively short,
CI < 1.15; head dorsum with very dense, not coarse longitudinal rugulosity and
reticulation, number of rugulae between frontal carinae level with the eyes > 25,
surface between rugulae densely punctated (Fig. 143, A). Petiole massive, with
very short, thick peduncle, its anterior surface slightly concave or even almost
straight (Fig. 143, C). - Tibet, India: Sikkim, Nepal M. kozlovi Ruzsky
-, Smaller, more slender species, HW < 1.15, AL < 2.0 mm, with relatively longer
head, CI > 1.20. Head dorsum with not so dense sculpture, number of rugae be­
tween frontal carinae level with the eyes < 25 (e.g. Fig. 219, A). Petiole with dis­
tinct peduncle, anterior petiolar margin sharply concave (e.g. Figs 44, C, D; 207,
C, D; 219, C, D) 19

738
19(18). Promesonotal dorsum (seen in profile) distinctly impressed transversely, some-
times saddle-shape; mesonotum abruptly curves down to propodeum to form dis-
tinct, deep and wide metanotal groove (e.g. Figs 44, C; 207, C) 20
-. Promesonotal dorsum (seen in profile) forms more or less regular arch; mesono-
tum not impressed transversely; mesonotum gently curves down to propodeum
to form a metanotal groove, which is generally shallow, but can be quite deep
(e.g. Figs 219, C) 23
20(19). Petiolar node and postpetiole with coarse longitudinally-concentric rugae, surface
between them smooth, not punctated (Fig. 207, C, D). Males: scape long, SI, 0.90.
- India: Kashmir M. rhytida Radchenko et Elmes
-. Petiolar node and postpetiole only punctated, without rugosity, at most with
very fine short striation, sculpture may be reduced at different extent (e.g. Fig.
44, C, D). Males: scape shorter, SI, < 0.60 (males of M. ordinaria are un-
known) 21
21(20). Sides of head with extremely short decumbent pubescence, which is visible under
magnification > 70 x, only occipital margin with a few long erect hairs (Fig. 44, A).
- North-eastern Pakistan, India: Kashmir M. cachmiriensis Forel
-. Sides and occipital margin of head with much longer subdecumbent hairs, which
are well visible under magnification < 30x (Figs 181, A; 309, A) 22
22(21). Surface of head dorsum between rugae at most very finely superficially punctated, ap-
pears shiny (Fig. 181, A). - India: Kashmir M. ordinaria Radchenko et Elmes
-. Surface of head dorsum between rugae densely and coarsely punctated, ap-
pears dull (Fig. 309, A). - North-eastern Pakistan, India: Kashmir, Himahal
Pradesh M. wardi Radchenko et Elmes
23(19). Frontal carinae curved outwards to merge with the rugae that surround antennal
sockets1 (e.g. Fig. 88, A) 24
-. Frontal carinae merging with rugae that extend to the occipital margin, they do not
curve outwards and do not merge with rugae that surround antennal sockets (e.g.
Fig. 219, A) : 26
24(23). Sides of alitrunk with reduced irregular longitudinal rugulae. or even striation,
petiolar node and postpetiole finely punctated only, without rugosity (Fig. 319,
C, D). - North-eastern Pakistan, India: Himahal Pradesh
M. whittmeri Radchenko et Elmes
-. Sides of alitrunk with well developed longitudinal rugosity, petiolar node and post-
petiole at least partly rugose (e.g. Fig. 219, C, D) 25
25(24). Postpetiolar dorsum with numerous, not coarse longitudinally-concentric rugae,
petiolar node dorsum coarsely punctated (Fig. 262, C, D). India: Himahal Pradesh,
Uttarakhand, Nepal; possibly elsewhere in Himalaya . . . . . M. smythiesii ¥ oxe\
-. Postpetiolar dorsum with a few coarse longitudinal rugae, petiolar node dorsum
with irregular, short sinuous rugae and fine reticulation (Fig. 88, C, D). - India:
Kashmir M. fortior Forel
26(23). Petiolar node and postpetiole distinctly, often coarsely, rugose (e.g. Fig. 225,
C, D) ' 27
-. Petiolar node and postpetiole densely punctated, without coarse rugosity, occa-
sionally with short, fine rugulae or striation (e.g. Fig. 219, C, D) 29
27(26). Surface of head dorsum and alitrunk between rugae densely and quite coarsely
punctated, appears completely dull (Fig. 87, A, C, D). - India: "Himalaya, Pach-
marhi, Shurr" M. foreliana Radchenko et Elmes

739
 -. Surface of head dorsum and alitrunk between rugae smooth or at most with
fine superficially micro-punctated, appears more or les shiny (e.g. Fig. 225,
A, C, D) 28
28(27). More hairy species: head margins with long, abundant erect to suberect hairs, the
longest hairs on scape distinctly longer than maximum diameter of scape (Fig. 8,
A-C). Head dorsum mostly with longitudinal rugae, only its lateral parts and
occiput with reticulation (Fig. 8, A). - North-eastern Afghanistan . .
M. afghanica Radchenko et Elmes
-. Less hairy species: head margins with short subdecumbent hairs, the longest hairs
on scape not longer than maximum diameter of scape (Fig. 225, A, B). Only
frons with longitudinal rugae, remainder parts with reticulation (Fig. 225, A).
- North-eastern Afghanistan, India: Kashmir, Himahal Pradesh, Nepal, Bhutan;
possibly somewhere else in Himalaya M. rupestris Forel
29(26). Head dorsum mainly with longitudinal rugae, reticulate sculpture may present only
on occiput and lateral parts of head dorsum above the eyes. Sides of pronotum
mainly with longitudinal rugosity (Fig. 219, A, C). - India: Kashmir, Nepal,
Bhutan; possibly very widespread in Himalaya M. rugosa Mayr
-. Only frons with longitudinal rugae, remainder parts of head dorsum with reticu-
lation. Sides of pronotum with sinuous rugae and reticulation (Figs 10, A, C;
96, A, C) 30
30(29). Mandibles with 7-9 teeth. Propodeal spines somewhat shorter, mean ESLI 0.62.
- North-eastern Afghanistan, north-eastern Pakistan, India: Kashmir, Himahal
Pradesh, West Bengal (Darjeeling), Sikkim, Meghalaya, Nepal, Bhutan
M. aimonissabaudiae Menozzi
-. Mandibles with 10-13 teeth. Propodeal spines somewhat longer, mean ESLI 0.71.
- India: Kashmir, Himahal Pradesh, West Bengal (Darjeeling), Nepal
M. hecate Weber

5.7. A key for the identification of the Myrmica species of South and South
East Asia (workers")
* M. mixta (Fig. 168) is known only from the holotype queen collected in Sichuan Prov.
of China; it has a very unusual combination of morphological features, and generally seems
to be intermediate between the ritae- and inezae species-groups. Its quite coarse body sculp-
ture, very long propodeal spines and fig-shaped postpetiole are all features of the ritae-group.
On the other hand, the not-notched anterior clypeal margin, relatively short scape, not
pointed propodeal lobes, not well marked and not sharp upper latero-ventral corners of the
head and another shape of petiole contradict with the characters of the n'tee-group species.

1. Scape strongly angularly curved at the base, with small vertical lobe (Fig. 258, B-E).
- China: Sichuan Prov M. sinoschencki Radchenko et Elmes
-. Scape gradually curved at the base, without any trace of ridge or lobe (e.g. Figs 58,
B; 167, B) : 2
2(1). Alitrunk dorsum partly with transversal rugosity (e.g. Figs 250, C, D; 324, C, D) . . . 3
-. Alitrunk dorsum with various sculpture, but never with transversal rugosity (e.g.
Figs. 58, C, D; 211, C, D) 13

740
 3(2). Lateral and occipital margins of head with long numerous suberect to erect hairs
. (e.g. Figs 198, A; 314, A; 324, A) 4
-. Lateral margins of head either with no pilosity or with short decumbent hairs; if any
long erect to suberect hairs present, they are restricted to the occipital margin and
genae (e.g. Figs 201, A; 250, A; 277, A) , 8
4(3). Head slightly longer than broad, CI < 1.15, subsquare, with distinctly marked
occipital corners (Fig. 198, A). - China: Yunnan Prov
M. pleiorhytida Radchenko et Elmes
-. Head distinctly elongate, CI > 1.20, suboval, with barely marked occipital corners
(e.g. Fig. 314, A) 5
5(4). Mesonotal and propodeal dorsum with relatively coarse transversal rugosity, num-
ber of rugae on this area < 20 (e.g. Fig. 314, C, D) 6
-. Mesonotal and propodeal dorsum with finer transversal rugosity, number of rugae
on this area > 25 (e.g. Fig. 324, C, D) 7
6(5). Petiole low, V\x 1.68, its node with elongate flattened dorsum (Fig. 314, C). Body
colour reddish-brown. - China: Shaanxi Prov M. weii Radchenko et Zhou
-. Petiole higher, PIj < 1.55, its node with a short, slightly convex, gradually sloping
posteriorly dorsal surface (Fig. 169, C). Body colour blackish-brown. - China:
Shaanxi Prov M. multiplex Radchenko et Elmes
7(5). Frontal carinae merge with the rugae that extend back to the posterior head mar-
gin, do not curve outwards and do not merge with the rugae that surround anten-
nal sockets. Frons wider, FI 0.44-0.45. Surface of head dorsum between rugae
appears shiny, only finely superficially micropunctated (Fig. 324, A). - China:
Yunnan Prov. M. yunnanensis Radchenko et Elmes
-. Frontal carinae curved outwards to merge with the rugae that surround antennal
sockets; frons narrower, FI 0.39. Surface of the head dorsum between rugae
coarsely and densely punctated, appears dull (Fig. 102, A) - China: Yunnan Prov.
M. heterorhytida Radchenko et Elmes
8(3). Basal third of first gastral tergite densely punctated and longitudinally ruguloso-striated;
this sculpture gradually reducing posteriorly, the rest of surface of the tergite with
well visible superficial hexagonal microsculpture (seen under magnification > 50x)
(Fig. 250, D). - China: Sichuan Prov M. sculptiventris Radchenko et. Elmes
-. Whole surface of first gastral tergite smooth (seen under magnification > 50 x) . . . 9
9(8). Propodeal spines long, ESLI > 0.45, massive, strongly widened at the base and
often curved down on their distal third (Fig. 277, C, D). - China: Shaanxi
Prov M. taibaensis Wei, Zhou et Lìu
-. Propodeal spines shorter, ESLI < 0.40, slender and straight (e.g. Figs 201, C, D;
249, C, D) 10
10(9). Head dorsum posterior to the eyes with reticulation, rest of head dorsum with
longitudinal rugosity (Fig. 201, A). - China: Yunnan Prov
-. M. polyglypta Radchenko et Rigato
-. Head dorsum posterior to the eyes with longitudinal rugosity; reticulation, if
present, restricted to temples and/or posterior part of occiput (Figs 106, A; 193, A;
249, A) 11
11(10). Lateral margins of head posterior to the eyes with abundant short decumbent hairs.
Frontal carinae curve outwards to merge with the rugae that surround the antennal
sockets (Fig. 249, A). - China: Shaanxi Prov M. schulzi Radchenko et Elmes

741
 -. Lateral margins of head posterior to the eyes without short decumbent hairs.
Frontal carinae merge with the rugae that extend back to the posterior head mar-
gin, do not curve outwards and do not merge with the rugae that surround anten-
nal sockets (Figs 106, A; 193, A) 12
12(11). Rugosity on the head dorsum partly reduced. Frons (level with the eyes) having
< 15 fine, slightly sinuous longitudinal rugae, with only some of them extending
unbroken to the occipital margin; surface between rugae densely and quite
coarsely punctated, appearing dull (Fig. 193, A). Propodeal dorsum with trans-
versal rugosity (Fig. 193, C, D). Scape longer than head width. - China: Shaanxi
Prov M. phalacra Radchenko et Elmes
-. Rugosity on the head dorsum not reduced. Frons (level with the eyes) having > 20
longitudinal rugae that extend unbroken to the occipital margin; surface be-
tween rugae very finely superficially punctated but appearing more or less shiny
(Fig. 106, A). Propodeal dorsum with short, slightly sinuous longitudinal rugae
(Fig. 106, C, D). Scape shorter than head width. - China: Sichuan Prov
M. hlavaci Radchenko et Elmes
13(2). Anterior clypeal margin not-notched medially; frontal carinae curved outwards to
merge with the rugae that surround antennal sockets (e.g. Fig. 167, A). Propodeal
lobes rounded apically; body not slender, scape, propodeal spines and petiole
shorter (SIj < 0.90, ESLI < 0.40, Pl: < 1.55) (e.g. Fig. 167, C, D) 14
-. Anterior clypeal margin distinctly notched medially; frontal carinae merging with
rugae that extend to the occipital margin, they do not curve outwards and do not
merge with rugae that surround antennal sockets (e.g. Fig. 211, A). Propodeal
lobes pointed apically; body slender, scape, propodeal spines and petiole longer
(SI1 > 0.95, usually > 1.00, ESLI > 0.45, PIj > 1.60; indices do not apply to
M. gigantea) (e.g. Fig. 211, C, D) 16
14(13). Very large, one of the largest known Myrmica species, HW > 1.60, AL > 2.50 mm.
Very hairy species, number of standing hairs on petiolar node > 20 (Fig. 167, C,
D).• - Taiwan M. mirabilis Elmes et Radchenko
-. Smaller species, HW < 1.25, AL < 2.30 mm. Less hairy, number of standing hairs
on petiolar node < 15 (e.g. Fig. 58, C, D) 15
15(14). Mandibles with 11-13 teeth. - China: Sichuan, Hunan and Yunnan Prov
M. curiosa Radchenko et Elmes
-. Mandibles with 7-9 teeth. - Taiwan M. arisana Wheeler
16(13). Surface between rugae on head and dorsal surface of waist is smooth or at most
very finely superficially micropunctated, appearing shiny; head, alitrunk and waist
usually with a similar coarse rugosity (e.g. Figs 164, A, C, D; 211, A, C, D) . . . 17
-. Surface between rugae on waist always distinctly and often coarsely punctated, ap-
pearing dull; surface of head usually is similarly punctated (except for M. angu-
lata); rugosity on head dorsum often is noticeably finer than that on alitrunk (e.g.
Figs 20, A, C, D; 69, A, C, D; 246 A, C, D) 25
17(16).Whole head dorsum with almost straight, subparallel rugae, completely lacking
reticulation, number of rugae between frontal carinae level with the eyes > 6 (Figs
189, A; 211, A) 18
-. Head dorsum with distinctly sinuous rugae and reticulation, if reticulation not
developed, number of rugae between frontal carinae level with the eyes < 6 (e.g.
Figs 78, A; 164, A; 201, A) 19

742
18(17). Occipital and lateral margins of head with long standing hairs (Fig. 189, A). Seen
at magnification 100 x surface of head weakly micropunctated. Pronotum coarsely
reticulate (Fig. 189, C, D). Head brownish-red, not strongly contrasting with the
darker alitrunk. - China: Sichuan Prov M. pararitae Radchenko et Elmes
-. Only occipital margin of head with long erect hairs (Fig. .211, A). Seen at magni­
fication lOOx microsculpture on surface of head completely invisible. Pronotum
with longitudinal rugae (Fig. 211, C, D). Head yellow, strongly contrasting with the
darker alitrunk. - Burma, Thailand M. ritae Emery
19(17). Very large, the largest known Myrmica species, HW > 1.80, AL > 3.30 mm.
- N Vietnam M. titanica Radchenko et Elmes
-. Smaller species, HW < 1.65, AL < 2.60 mm 20
20(19). Scape, petiole and propodeal spines relatively short, SI : 0.90, Plj 1.50, ESLl 0.35
(Fig. 95, B, C). Quite big species, HW 1.62,' AL 2.58 mm. - Burma
M. gigantea (Collingwood)
-. Scape, petiole and propodeal spines distinctly longer, SIj > 1.00, PIj > 1.60, usu­
ally > 2.00, ESLI > 0.50 (e.g. Fig. 164, B-D). Relatively smaller species,
HW < 1.50, AL < 2.55 mm 21
21(20). Long standing hairs present only on occipital margin, genae and temples occa­
sionally with an only few hairs. Frons with very coarse sculpture, having only four
rugae between frontal carinae level with the eyes (Figs 78, A; 164, A) 22
-. Both occipital and lateral margins of head with numerous long standing hairs.
Frons with various sculpture (Figs 202, A; 251, A; 257, A) 23
22(21). Alitrunk dorsum with numerous long standing hairs (Fig. 78, C). Lateral sides of
head slightly convex and occipital corners narrowly rounded, so that head .appears
roughly subrectangular (Fig. 78, A). - Isl. Pulo Laut (SE of Borneo)
M. emeryi Radchenko et Elmes
-. Alitrunk dorsum with a few scattered standing hairs (Fig. 164, C). Lateral sides of
head strongly convex and occipital corners widely rounded, barely marked, so that
head appears roughly suboval (Fig. 164, A). - Burma . . . . . M. margaritae Emery
23(21). Posterior half of head dorsum (from above the level of the eyes) with coarse reti­
culation, remainder with sinuous rugae; frons level with the eyes between frontal
carinae with only 4 coarse rugae (Fig. 257, A). Petiolar node and post­
petiole, as well as sides of alitrunk, with coarse reticulation (Fig. 257, C, D).
- China: Guangxi Prov M. sinensis Radchenko, Zhou et Elmes
-, Head dorsum with sinuous rugae; coarse reticulation, if present, restricted to
occiput; frons with various sculpture (Figs 202, A; 251, A). Petiolar node .and
. postpetiole, as well as sides of alitrunk, with coarse rugosity (Figs 202, C, D; 251,
C, D) 24
24(23). Frons between frontal carinae level with the eyes with 4 very coarse rugae (Fig.
202, A). - Taiwan M. pulchella Santschi
, -. Frons between-frontal carinae level with the eyes with > 6 finely rugae (Fig. 251,
A). - Taiwan, southern China, Northern Vietnam M. serica Wheeler
25(16). Waist with fine sculpture, distinctly contrasting with much coarser sculpture on
alitrunk (Figs 69, С, D; 246, С, D) 26
-. Waist with coarser sculpture, similar to that on alitrunk (e.g. Fig. 20, C, D) . . . .'27
26(25). Head shorter, mean CI 1.11, with narrowly rounded, well-marked occipital cor­
ners, appears subrectangular (Fig. .69, A). Scape shorter, SIa < 1.10. - China:
Guangxi Prov M. draco Radchenko, Zhou et Elmes

743
 -. Head longer, mean CI 1.19, with widely rounded, barely marked occipital
corners, appears suboval (Fig. 246, A). Scape longer, SIj > 1.20. - Northern
Vietnam M. schoedli Radchenko, Elmes et Viet
27(25). Surface of head dorsum between rugae not punctated, appears shiny, only waist
with coarse rugosity and densely punctated, appears dull (Fig. 20, A, C, D).
- China: Guangxi and Hubei Prov M. angulata Radchenko, Zhou et Elmes
-. Surface of both head dorsum and waist between rugae punctated, appears dull
(Fig. 200, A, C, D; 323, A, C, D) 28
28(27). Sculpture on head dorsum distinctly finer than that on alitrunk, frons between
frontal carinae level with the eyes with 9 rugae, head shorter, CI 1.11 (Fig. 200, A).
Petiole shorter, PIj 1.69; propodeal spines directed mainly backward, nearly hori-
zontal (Fig. 200, C). - China: Sichuan Prov M. poldii Radchenko et Rigato
-. Sculpture on head dorsum similar to that on alitrunk, frons between frontal cari-
nae level with the eyes with 6 rugae, head longer, CI > 1.15 (Fig. 323, A). Petiole
longer, PIj > 2.00; propodeal spines directed backward and upward, inclined at an
angle more than 45° (Fig. 323, C). - Northern Vietnam
M. yamanei Radchenko et Elmes

5.8. A key for the identification of the fossil Myrmica species

1. Propodeum with short, blunt teeth; head and alitrunk with fine reticulation; hind
tibiae with distinct spur (Fig. 330) . . . M. paradoxa Radchenko, Dlussky et Elmes
-. Propodeum with long spines; head and alitrunk with very coarse sinuous rugosity
and reticulation (Figs 328, 329, 331); hind tibiae with or without spur 2
2(1). Propodeal spines directed backward and upward, slightly curved down at the apex
(Fig. 331); hind tibia without spur M. rudis (Mayr)
-, Propodeal spines directed backward, straight or slightly curved inward (in dorsal
view) (Figs 328, 329); hind tibiae with or without spurs 3
3(2).Propodeal spines straight, strongly divergent (in dorsal view) (Fig. 328); hind
tibiae with spur; larger species, body length ca. 6.5 mm
M. eocenica Radchenko, Dlussky et Elmes
-. Propodeal spines subparallel, not divergent (in dorsal view) (Fig. 329) ; hind tibiae
with or without spurs; smaller species, body length ca. 5 mm or less 4
4(3).Hind tibiae with spurs. Propodeal lobes rounded, not sharply pointed apically
M. longispinosa Mayr
-. Hind tibiae without spurs. Propodeal lobes sharply pointed apically
M. intermedia (Wheeler)

744
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St. Petersburg University Press, St Petersburg, pp. 97-112.
Akino, T., J. J. Knapp, J. A. Thomas and G. W. Elmes. 1999. Chemical mimicry and host specificity in
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780
INDEX OF THE ALL LATIN NAMES

Myrmica names recognized here as good species

The names of good extant Old World Myrmica species marked by italic, of good extinct
Myrmica species - italic underlined; first page of the species data'in Chapter 3.3, 3.4
marked by bold; that/those for Figs (Chapters 3.3, 3.4) - b y bold italic, that/those for names
in the Key (s) (Chapter 5) - by bold italic underlined.

ademonia 63, 72, 79, 80, 81,120, 296, 298, cagnianti 67, 72, 108, 109, 161, 164, 373,
331, 332, 333, 677, 679, 233 374, 679, 716
afghanica 68, 72, 81, 82, 305, 334, 335, caucasicola 72, 109, 110, 118, 207, 223,
679. 740 269, 270, 278, 375, 376, 577. 723
aimonissabaudiae 66, 72, 73, 78, 82, 83, collingwoodi 65, 73, 10 5, 111, 378, 379,
84, 108, 130, 243, 248, 321, 336, 337, 679. 737
338, 679, 740 commarginata 63, 73, 112, 113, 380, 679,
aloba 67, 72, 76, 78, 84, 85, 86, 109, 161, 732
259, 264, 324, 339, 340, 341, 679, 716 constricta 67, 73, 75, 113, 114, 145, 146,
alperti 64, 72, 86, 87, 152, 321, 322, 342, 147, 148, 156, 245, 246, 324, 381, 382,
679. 737 383, 679, 716, 724
anatolica 62, 72, 87, 88, 343, 344, 345, 679, curiosa 68, 73, 115, 116, 384, 385, 674,
723 679. 742
angulata 64, 72, 88, 89, 200, 306, 346, 347, deplanata 67, 73, 75, 76, 78,116,117,118,
674, 679, 742, 744 119, 256, 270, 278, 291, 293, 298, 386,
angulinodis 63, 72, 74, 78, 89, 90, 91, 92, 387, 388, 658, 662, 663, 675, 679, 686,
95, 121, 122, 127, 134, 135, 161, 162, 704, 715, 723, 734
167, 258, 348, 349, 350, 676, 679, 685, displicentia 63, 72, 73, 112, 119, 120, 156,
686, 688. 733 298, 389, 390, 391, 733
arisana 65, 72, 77, 78, 92, 93, 94, 180, 351, divergens 67, 73, 92, 112, 121, 122, 167,
352, 353, 674, 679, 701, 742 392, 393, 394, 676, 679, 733
arnoldii 20, 30, 52, 59, 60, 72, 94, 95, 145, draco 65, 73, 122, 123, 219, 272, 329, 395,
146, 205, 311, 354, 355, 356, 676, 679, 396, 397, 679, 743
6 8 3 , 6 9 7 , 6 9 9 , 7 0 7 , 7 0 8 , 732 dshungarica 59, 60, 65, 68, 73, 74, 75, 76,
bactriana 68, 72, 73, 78, 96, 97, 249, 306, 77, 78, 82,123, 124,125, 126,127,133,
357, 358, 679, 736 159, 179, 258, 283, 398, 399, 400, 663,
bakurianica 67, 72, 98, 99, 100, 359, 360, 664, 675, 679, 683, 697, 700, 701, 215.
361, 679, 725, 728 eidmanni 62, 73, 74, 78, 127, 128, 158;
bergi 66, 67, 72, 74, 75, 78, 101, 102, 103, 401, 402, 403, 662, 663, 679, 686, 732
104, 121, 138, 139, 140, 155, 175, 257, emeryi 31, 64, 73, 128, 129, 130, 199, 221,
291, 362, 363, 364, 663, 675, 679, 683, 222, 227, 279, 404, 671; 673, 674, 679,
686, 724, 735 689, 743
bibikoffi 30, 67, 72, 104, 105, 150, 151, eocenica 75. 655. 655, 744
251, 252, 316, 365, 366, 367, 679, 716, ereptrix 66, 73, 130, 131, 248, 405, 680,
720. 721 710, 736
boltoni 65, 72,105, 106, 111, 123, 200, 329, excelsa 62, 72, 73, 77, 78, 131, 132, 133,
368, 673, 679, 737 167, 302, 309, 406, 407, 408, 677, 680,
brancuccii 68, 72, 106, 107, 275, 369, 673, 683. 729
679. 736 ferganensis 60, 73, 125,133, 134, 159,179,
cachmiriensis 65, 66, 72, 75, 78, 107, 108, 303, 409, 680, 735
206, 210, 224, 225, 320, 321, 370, 371, forcipata 62, 73, 90, 91, 92, 134, 135, 168,
372, 679, 739 410, 411, 412, 680, 730

78t
foreliana 66, 72, 73, 135, 136, 248, 413, kirghisorum 62, 73, 74, 79, 167, 168, 169,
680. 739 257, 457, 458, 459, 675, 680, 734
fortior Forel 68, 73, 102,136, 137, 179, 180, koreana 67, 74, 118, 169, 170, 209, 269,
283, 414, 415, 416, 680, 739 270, 278, 460, 461, 462, 662, 676, 680,
gallienii 67, 73, 74, 75, 78, 102, 103, 121, 685, 686, 704, 221, 731
138, 139, 140, 156, 165, 166, 179, 181, kotokui 19, 20, 26, 65, 74, 75, 79, 92, 170,
245, 287, 294, 299, 417, 418, 419, 663, 171, 172, 179, 180, 232, 236, 238, 242,
664, 680, 686, 216, 724 463, 464, 465, 680, 701, 750
georgica 67, 73, 141, 142, 143, 172, 173, kozakorum 31, 67, 74, 142, 172, 174, 175,
174, 260, 264, 313, 420, 680, 226, 228 313, 466, 467, 468, 680, 226, 729
gigantea 30, 64, 73,143, 421, 680, 742, 743 kozlovi 61, 74, 75, 77, 78, 79,175, 176, 177,
hecate 66, 73, 83, 116, 144, 165, 243, 248, 178, 179, 180, 220, 469, 673, 675, 680,
422, 423, 424, 673, 680, 740 684, 236, 738
hellenica 67, 74, 76, 79,113,114,145,146, kryzhanovskii 60, 74, 178, 179, 470, 680,
147, 148, 156, 287, 288, 324, 425, 426, 735
427, 680, 211, 726, 729 kurokii 59, 61, 73, 74, 76, 79, 92, 139, 171,
heterorhytida 64, 74, 148, 149, 323, 428, 175, 176, 177, 179, 180, 181, 236, 237,
680, 741 239, 471, 472, 473, 675, 677, 680, 683,
hirsuta 25, 26, 27, 32, 33, 67, 74, 104, 149, 697. 700. 701. 733
150, 151, 182, 184, 251, 252, 253, 300, laurae 27, 28, 52, 57, 59, 60, 61, 74, 76, 79,
316, 429, 430, 431, 680, 710, 211, 720, 150, 163, 165, 181, 182, 183, 252, 474,
721, 724 475, 611, 680, 683, 697, 706, 707, 720.
hlavaci 64, 74, 151, 152, 216, 317, 432, lemasnei 30, 56, 59, 61, 74, 161, 164, 183,
680, 742 184, 185, 252, 476, 477, 680, 220, 721
indica 64, 74, 86, 123,152, 153, 321, 322, lobicornis 27, 32, 59, 62, 72, 73, 74, 75, 77,
433, 434, 435, 680, 737 78, 79, 81, 88, 90, 112, 116, 117, 118,
inezae 59, 60, 61, 74, 116, 153, 154, 202, 119, 120, 121, 126, 127, 128, 132, 133,
203, 212, 226, 301, 322, 323, 436, 437, 134, 156, 157, 158, 162, 166, 167, 168,
673, 674, 679, 680, 683, 697, 698, 710, 185, 186, 187, 188, 189, 190, 191, 192,
736. 740 193, 194, 204, 206, 208, 218, 223, 225,
intermedia Wheeler 74, 257, 653, 661, 656, 230, 250, 251, 252, 254, 255, 256, 257,
744 258, 264, 266, 268, 269, 270, 281, 291,
invicta 67, 74,154, 155, 206, 279, 438, 678, 293, 296, 298, 302, 309, 323, 324, 327,
680. 722 330, 478, 479, 480, 658, 660, 661, 662,
jennyae 68, 74, 155, 156, 439, 440, 441, 663, 675, 676, 677, 678, 680, 683, 684,
680. 724 685, 686, 692, 697, 705, 706, 709, 715.
jessensis 62, 74, 79, 127, 128, 157, 158, 223, 731
442, 443, 444, 680, 731 lobulicornis 27, 62, 72, 75, 76, 79,186, 188,
juglandeti 60, 72, 74, 77, 79, 125, 133,158, 189, 191, 192, 193, 194, 204, 205, 252,
159, 179, 445, 446, 680, 735 324, 481, 482, 483, 680, 686, 709, 715
kabylica 56, 61, 74, 160, 161, 184, 447, lonae 27, 67, 75, 150, 165, 166, 194, 195,
448, 680, 220, 721 196, 197, 242, 250, 251, 252, 254, 255,
kamtschatica 63, 72, 74, 79, 91, 161, 162, 291, 484, 485, 486, 680, 686, 21Â, 719.
163, 240, 298, 449, 450, 451, 680, 686, 725. 728
733 longispinosa 75, 653, 654, 655, 2M
karavajevi 25, 27, 52, 56, 57, 59, 61, 73, 74, luteola Kupyanskaya 59, 63, 160, 197, 198,
75, 78, 79, 160,161,163, 164, 165, 166, 201, 275, 487, 488, 489, 674, 677, 680,
183, 184, 246, 252, 253, 452, 453, 680, 683, 693, 697, 708, 730
685, 697. 709. 720. 721 margaritae 28, 64, 75, 88, 89, 129, 199,
kasczenkoi 62, 63, 74, 90, 91, 112, 120, 200, 221, 222, 275, 279, 490, 663, 680,
131, 133, 156, 166, 167, 218, 258, 454, 743
455, 456, 664, 676, 680, 683, 732, martensi 65, 75, 105, 111, 200, 491, 492,
733 680. 737

782
mirabilis 63, 75, 198, 201, 493, 674, 681, rubra 3, 7, 8, 9, 16, 17, 18, 20, 21, 24, 25,
695. 708. 742 26, 27, 28, 32, 52, 57, 59, 60, 65, 68, 73,
mixta 61, 75, 202, 203, 494, 674, 681, 740 74, 75, 76, 78, 79, 83, 93, 94, 101, 116,
multiplex 64, 75, 203, 322, 323, 495, 496, 124, 125, 133, 137, 140, 145, 146, 151,
681. 741 159, 165, 168, 170, 171, 177, 179, 180,
myrmicoxena 29, 59, 60, 63, 75, 204, 205, ' 184, 185, 191, 194, 196, 228, 229, 230,
497, 677, 681, 683, 697, 699, 708, 709, 231, 232, 233, 234, 235, 236, 237, 238,
710. 720, 721 239, 240, 241, 242, 244, 247, 250, 251,
nitida 68, 75, 154, 205, 206, 498, 499, 500, 252, 259, 261, 264, 265, 266, 268, 276,
673,681,238 283, 294, 299, 300, 303, 304, 305, 321,
obscura 67, 75, 189, 195, 206, 207, 208, 327, 539, 540, 541, 663, 674, 675, 676,
266, 269, 270, 278, 291, 501, 502, 503, 677, 678, 681, 683, 684, 685, 686, 688,
681. 714 693, 697, 700, 701, 702, 710, 711, 714,
onoyamai 67, 75, 208, 209, 277, 278, 279, 722. 730. 734
504, 505, 506, 681, 731 rudis 76, 653, 655, 657, 744
ordinaria 66, 75, 209, 210, 224, 507, 508, ruginodis 18, 19, 20, 25, 26, 27, 28, 32, 33,
681. 739 59, 64, 65, 73, 75, 76, 77, 78, 79, 84, 94,
orthostyla 68, 75, 210, 211, 509, 510, 511, 133, 137, 147, 163, 164, 166, 170, 171,
675. 681, 734 176, 180, 191, 196, 197, 229, 230, 231,
pachei 59, 63, 64, 75, 148, 151, 203, 211, 232, 235, 236, 237, 238, 239, 240, 241,
212, 216, 218, 220, 274, 275, 301, 317, 242, 264, 265, 270, 276, 294, 295, 296,
319, 322, 329, 512, 513, 514, 673, 674, 297, 298, 542, 543, 544, 659, 660, 663,
679, 681, 683, 697, 698, 699, 710, 676, 677, 678, 681, 685, 686, 688, 701,
736 710.711. 714. 722, 730
paradoxa 75, 653, 654, 656, 744 rugosa 59, 65, 66, 76, 83, 92, 93, 94, 101,
pararitae 64, 75, 213, 276, 515, 516, 681, 108, 123, 124, 130, 136, 144, 154, 176,
743 180, 206, 210, 224, 225, 242, 243, 244,
pelops 67, 75, 213, 214, 215, 278, 517, 681, 246, 247, 248, 258, 282, 320, 545, 546,
714 547, 663, 664, 673, 679, 681, 683, 692,
petita 68, 76, 215, 518, 681,236 693, 697, 698, 699, 700, 710, 740
phalacra 64, 76, 151, 216, 317, 519, 520, rugulosa 27, 59, 66, 67, 73, 76, 77, 79, 113,
681. 742 114, 115, 124, 125, 138, 139, 140, 145,
pisarskii 62, 76, 216, 217, 218, 258, 521, 146, 156, 165, 244, 245, 246, 259, 260,
522,523,676.681. 731 284, 285, 287, 289, 548, 549, 550, 663,
pleiorhytida 64, 76, 2Г8, 219, 322, 524, 664, 681, 683, 716, 724
525.681. 741 rupestris 66, 73, 76, 79, 136, 244, 246, 247,
poldii 65, 76, 219, 220, 526, 681, 214 248, 551, 552, 553, 681, 740
polyglypta 64, 76, 220, 221, 527, 681, 741 ruzskyana 68, 73, 76, 96, 249, 554, 555,
pulchella 64, 73, 76, 129, 199, 221, 222, 681. 736
276, 279, 528, 529, 681, 743 sabuleti 27, 29, 32, 59, 66, 67, 76, 79, 86,
ravasinii 67, 76, 110, 214, 223, 224, 268, 104, 142, 147, 149, 150, 151, 165, 166,
278, 530, 531, 532, 681, 714, 722 173, 174, 175, 181, 182, 183, 184, 185,
rhytida 68, 76, 206, 218, 224, 225, 321, 194, 195, 196, 197, 240, 250, 251, 252,
533. 534. 535. 681. 739 253, 254, 259, 260, 261, 262, 263, 265,
rigatoi 61, 76, 202, 225, 226, 536, 681, 738 271, 278, 287, 288, 289, 290, 291, 312,
ritae 14, 20, 28, 30, 58, 59, 61, 64, 65, 76, 313, 316, 556, 557, 558, 657, 660, 681,
86, 89, 105, 111, 116, 123, 128, 129, 683; 686, 718, 719, 726, 728
130, 143, 152, 153, 197, 199, 200, 202, salina 67, 76, 98, 194, 196, 254, 255, 256,
213, 219, 221, 226, 227, 272, 273, 275, 280, 281, 287, 559, 560, 561, 681, 727.
276, 279, 306, 315, 321, 329, 537, 538, 729
653, 673, 674, 679, 681, 683, 687, 692, saposhnikovi 62, 72, 76, 79, 90, 217, 218,
693, 695, 696, 697, 698, 699, 700, 708, 256, 257, 258, 562, 563, 564, 661, 675,
710. 740. 743 681. 734

783
scabrinodis 27, 28, 32, 34, 59, 66, 67, 72, 591, 657, 658, 675, 682, 683, 686, 703,
76, 78, 79, 84, 85, 86, 88, 89, 90, 99, 718. 719. 727. 729
100, 102, 104, 109, 113, 116, 118, 119, spinosior 27, 67, 77, 104, 183, 185, 250,
121, 127, 139, 141, 142, 145, 146, 150, 278, 288, 289, 290, 291, 592, 593, 594,
156, 157, 163, 164, 165, 166, 167, 173, 682, 706,118, 719
174, 175, 181, 182, 183, 185, 189, 191, stangeana 67, 74, 75, 77, 79, 116,118, 291,
194, 195, 196, 197, 211, 230, 231, 232, 292, 293, 595, 596, 597, 682, 221, 728
233, 234, 244, 245, 250, 251, 252, 253, sulcinodis 19, 20, 27, 59, 62, 63, 73, 75, 77,
254, 255, 256, 257, 258, 259, 260, 261, 78, 79, 81, 84, 120, 138, 141, 156, 244,
262, 263, 264, 265, 266, 267, 268, 269, 245, 293, 294, 295, 296, 297, 298, 299,
270, 271, 283, 284, 286, 287, 288, 289, 323, 324, 327, 598, 599, 600, 659, 675,
291, 292, 293, 294, 295, 298, 303, 306, 676, 677, 682, 683, 685, 686, 688, 711,
310, 315, 316, 319, 320, 324, 326, 565, 715. 723. 732. 733. 735
566, 567, 660, 661, 663, 664, 665, 666, symbiotica 52, 67, 77, 299, 300, 601, 682,
675, 676, 677, 678, 681, 683, 684, 685, 215
686, 692, 693, 697, 702, 703, 705, 706, taibaensis 64, 77, 301, 602, 603, 682, 741
709, 718, 719,721, 728, 730 tamarae 62, 77, 132, 302, 309, 310, 604,
schencki 59, 67, 72, 74, 76, 77, 78, 79, 109, 605, 682, 221
ПО, 116, 118, 121, 154, 169, 170, 185, tenuispina 68, 77, 82, 133, 134, 158, 211,
186, 187, 188, 189, 195, 196, 206, 207, 303, 304, 305, 606, 607, 608, 673, 675,
208, 214, 223, 230, 252, 253, 254, 255, 682, 684, 754, 738
256, 266, 267, 268, 269, 270, 271, 272, tibetana 59, 68, 77, 81, 96, 179, 211, 303,
277, 278, 279, 280, 289, 293, 327, 568, 304, 305, 306, 609, 610, 675, 682, 683,
569, 570, 658, 662, 674, 675, 677, 678, 697, 699, 700, 735
681, 683, 684, 685, 686, 692, 697, 704, titanica 64, 77, 201, 276, 306, 307, 611,
705, 706, 714, 722, ZU, 734 682,143
schoedli 65, 76, 123, 219, 272, 273, 329, tobiasi 67, 77, 287, 307, 308, 612, 613, 614,
571, 572, 573, 681, 744 675. 682. 734
schulzi 64, 77, 273, 274, 275, 574, 681, transsibirica 62, 77, 132, 302, 308, 309,
741 310, 615, 616, 617, 676, 682, 686,721,
sculptiventris 64, 77, 106, 274, 275, 575, 729.
681. 741 tschekanovskii 78,310, 311, 618, 619, 676,
serica 64, 77, 152, 222, 275, 276, 576, 577, 682, 733
578, 681. 743 tulinae 67, 78, 100,142, 175,263, 311, 312,
siciliana 67, 77, 277, 278, 579, 580, 581, 620, 621, 622, 682, 686, 718, 719, 126,
681. 715 728
sinensis 64, 77, 129, 199, 221, 222, 276, turcica 67, 78, 99, 141, 142, 287, 312, 313,
278, 279, 582, 682, 743 314, 623, 624, 625, 682, 221,129
sinoschencki 67, 77, 270, 279, 280, 583, urbanii 64, 78, 143, 314, 315, 626, 682,
674, 682,740 737
slovaca 67, 73, 77, 79, 98, 252, 254, 256, vandeli 59, 66, 67, 78, 86, 251, 261, 315,
280, 281, 282, 584, 585, 586, 682, 686, 316, 317, 627, 628, 629, 682, 686, 707,
716, 724 713, 717, 725
smythiesii 59, 67, 68, 74, 77, 79, 82, 96, varisculpta 64, 78, 317, 318, 630, 682, 736
107, 123, 125, 126, 135, 136, 137, 206, villosa 64, 78, 206, 212, 317, 318, 319, 322,
246, 247, 248, 249, 282, 283, 303, 326, 631, 632, 682, 737
587, 588, 660, 675, 682, 683, 697, 700, vittata 68, 78, 319, 633, 682, 738
739 wardi 66, 78, 83, 108, 210, 224, 225, 320,
specioides 66, 67, 72, 73, 75, 76, 77, 78, 79, 321, 634, 635, 636, 682, 739
99, 100, 142, 145, 146, 147, 173, 174, weberi 64, 78, 152, 321, 322, 637, 638, 682,
175, 253, 256, 259, 260, 261, 263, 264, 737
271, 281, 284, 285, 286, 287, 288, 289, weii 64, 78, 203, 301, 322, 323, 639, 682,
292, 300, 307, 313, 314, 324, 589, 590, 741

784
wesmaeli 62, 78, 323, 324, 325, 640, 641, yamanei 65, 78 ; 123, 219, 220, 272, 328,
642, 682, 715 . . ' 329, 64S, 682, 744
williamsi 68, 78, 325, 326, 643, 682, 738 yunnanensis 64, 78, 148, 322, 323, 329,
wittmeri 68, 78, 215, 326, 327, 644, 682, 649, 682, 741
739 zojae 62, 78, 330, 650, 651, 652, 682, 723,
xavieri 62, 63, 78, 193, 324, 327, 328, 645, 731
646, 647, 682, 683, 715

Other names
aborigenica 72, 79, 161, 162 atomaria 664
acervorum 663, 666 atratula 664
adelaidae 664 Atta 27, 664, 668
aemula 664 baikalensis 12, 7 8 , 9 0 , 9 1
aenescens 308 balcanica 72, 79, 284, 285, 286, 287, 289,
Aeromyrma 689 663
agilis 664 barbarus 665, 668
Agroecomyrmex 665 barchanica 72, 78,101, 102, 104
ahngeri 72, 79, 259, 260, 261, 2 6 3 , 2 8 4 , basalis 664
289 beesoni 664
alajensis 72, 79,158, 159 bergiana 72, 79, 257, 258
alaskensis 708 bessarabica 72, 98, 99, 145, 146, 147, 250,
albuferensis 72, 78, 84, 85 251, 284,285, 287, 288, 657, 658
alcon 86, 240 betuliana 72, 79, 266, 267, 270
algerica 664 bicarinatum 667
algirica 664 bicolor Guérin-Méneville 669
alienus 193, 308 bicolor Heer 119, 664
alpestris 72, 79, 186, 187, 188, 189 bicolor Kupyanskaya 48, 72, 73, 119,120
alpina 72, 79, 176, 191, 192, 193 bidens 664
amaurocyclia 664 bidentata 664
amedeussabaudiae 82, 83 bremii 664
Anergates 664 breviceps 72, 658, 659
Aneuretinae 40, 689 brevispina 72, 126, 663
angustifrons 72, 79,185, 186, 187, 188,189 brevispinosa 77, 79
angustula 664 brunescens 72, 79, 186, 187, 188, 189, 270
apennina 72, 79,192, 193 Bupleuram 160
Aphaenogaster 30,143, 277, 278, 664, 665, bupleuroïdes 160
667, 668, 669, 690, 692 burtshakabramovitshi 72, 79, 185, 186,
archaica 664 187, 188, 189
arduennae 72, 79,185, 186, 187, 189, 191, busoni 72, 663
192 cadusa 72, 78, 131, 132
arenaria 664, 668 caeca 664, 665
arion 252 caespitum 271, 665, 666, 667
Armaniidae 689 Camponotus 8
armatus 160 carbonaria Forel 72, 73, 135, 136
Artemisia 291 . carbonaria Smith F. 135, 665
Asemorhoptrum 667 Cardiocondyla. 666, 669
aspersa Kupyanskaya 48, 72, 79,-81 . carinata Kupyanskaya 48, 72, 77, 308, 309,
aspersa Smith F. 79, 664 310
Astragalus 160 carinata Smith F. 665
atlantica 72; 663 Cataglyphis 43

785
caucasica 67, 72, 145, 146, 147, 679 flavidula 665
Cephalotes 27 flavus 191, 264, 265
chersonensis 72, 139, 140, 663 fodicus 665
chinensis 73, 79, 179, 180, 181 foreli 73, 186, 187, 188, 189
cingulata 665 foreliella 73, 79, 167, 168, 169
clandestina 73, 79, 244, 245 Formica 96, 147, 228, 230, 231, 235, 265,
Clinopodium 278, 299, 664, 665
clypeata 665 Formicidae 40, 51
colax 52 Formicinae 689
communis 160 Formicoxenus 663, 666, 667
concinna 665 formosa 92, 221,222, 665
consternes 665 formosae 73, 76, 221, 222
contigua 665 fortior Crawley 73, 75, 101, 102
corticalis 665 fragilis 665
Cratomyrmex 51 fugax 665
Crematogaster 227, 308, 664, 665, 666, furva 73, 78, 96, 97, 306
668, 669 fusca 8
cursor 73, 659 fuscipennis 665
curvithorax 73, 79, 280, 281, 282 fuscula 665
dagmarae 73, 78, 116, 117, 118, 119 galbula 665
daurica 73, 663 geminata 666, 667
debilior 73, 79, 244, 246, 247, 248 glyciphila 666
debilis 247, 665 gracilescens 666
dentata 73, 126, 663 gracillima 666
denticornis 73, 79, 185, 186, 187, 189, 230, grandis 73, 119, 663
660 granulinodis 27, 73, 250, 251, 659, 660
Dermaptera 45 Harpagoxenus 666, 669
derzhavini 73, 79, 294, 295, 296, 298 helleri 74, 79, 179, 180, 181
destructor 669 hemisphaerica 160
dicaporiaccoi 73, 78, 82, 83, 84, 108 herculeanus 8
diffusa 665 himalayana 74, 79, 282, 283
diligens 665 Hippophaë 104
diluta 27, 73, 79, 236, 237, 238 hirtula 666
Diplorhoptrum 665 Homoptera 45
Diptera 45 Huberia 51
diversus 668 humilis 666
Dodecamyrmica 30, 52, 94, 95, 311 Hylomyrma 51, 690, 691
dolens 73, 79, 284, 285, 286, 289 hyungokae 74, 79,157, 158
Dolichoderinae 689 iessensis 127', 157
domestica 665 impura 666
Drosera 265 incerta 666
duisburgi 665 incompleta 95, 699, 708
Ephebomyrmex 51 incurvata 74, 78, 90, 91
erraticum 307 infuscata 74, 126, 663
Eryngium 143 inornata 74, 663
Euphorbia 160 insolens 666
europea 73, 79, 228, 229, 232 intermedia Kuznetsov-ugamsky 74, 257,
Eutetramorium 51 258,661
everesti 73, 79, 246, 247, 248 interrupta 666
exigua Buckley 249 interruptus 669
exigua Ruzsky 73, 76, 249 iranica 74, 103, 663
eximia73, 79, 294, 295, 296, 297 irritans 666
faniensis 56, 73, 79, 163, 164, 165 iskanderi 74, 78, 124, 125, 126

786
jacobsoni 74, 78, 138, 139, 140 Manica 51, 116, 198, 662, 666, 667, 673,
jucunda 666 690,691,693,708
Juglandacea 159 medvedevi 75, 79, 291, 292
Juniperus 160 megacephala 666, 669
jurinei 666 mekongi 75, 79, 175, 176, 177
kamyschiensis 74, 78, 101, 102, 103 melanocephala 75, 663
khamensis 74, 660 melanocephalum 668
kievensis 74, 79, 186, 187, 188, 189 mellea 667
kirbii Stephens 74, 663 Meranoplus 669
kirbii Sykes 666 Mermis 15, 239, 270, 297, 300
kirgisica 74, 78, 101, 102 Messor 51, 308, 664, 665, 667, 668
kollari 666 microgyna 19, 28, 75, 79, 171, 231, 236,
kutteri 74, 79, 266, 267, 270 237, 238, 239, 241
lacteipennis 666 microrubra 26, 28, 75, 79, 228, 229, 231
lacustris 74, 79, 116, 118, 119, 256, 291, minkii 667
292,293,663 minuta Karawajew 73, 75, 113, 114, 145
laevigata 666 minuta Ruzsky 75, 78, 113, 114, 124, 125,
laevinodis 16, 27, 28, 32, 59, 74, 79, 113, 126, 145
114, 124, 133, 145, 170, 228, 229, 230, minuta Say 113, 114, 124, 145
231, 232, 236, 237, 238, 239, 241, 299, minutum 665
303, 304, 539, 540, 541 mirabile 201
laevinodoruginodis 74, 663 mixtossima 204
laevissima 666 modesta Foerster 667
laeviuscula 666 modesta Smith F. 667
Lasius 24, 147, 191, 193, 233, 246, 264, moesta 667 • '
265,277,278,308 molassica 667
latreillei 664, 669 Molinea 240
leonina 666 Monomarium 668
Leptogenys 227 Monomorium 135, 266, 664, 665, 666, 667,
Leptothorax 124, 125, 190, 284, 285, 288, 668, 669
663, 665, 666, 667, 669 montana 667
levinodis 228 moravica 75, 78, 116, 117, 118, 119
limanica 74,78, 138, 140, 663, 664 muscorum 667
lippula 667 mutata 75, 79, 236, 237, 238, 239
lippulum 667 mutica 667
lissahorensis 74, 79, 186, 187, 188, 189 Myrmeciinae 689
littoralis 74, 78, 127, 128 Myrmecina 664, 669
lobata 75, 663 . myrmecophila 59, 75, 79, 294, 295, 297
lobicornosulcinodis 75 myrmecoxena 204
longiceps 667 Myrmicaria 665, 669
longipes 667 Myrmicidae 51
longiscapus 75, 79, 228, 229, 23.0 Myrmicinae 51, 52, 689, 690, 691
lucidula 75, 663 Myrmicini 51, 52, 689, 690, 691, 692
luctuosa 75, 661, 662 Myrmicites 51, 52
lugubris 235 nausithous 233
lulakeranica 75, 119, 663 nebulosa 667
luteola Coss. et Dur. 75 neglecta 75, 663
lutescens 75, 78, 107, 108 nevodovskii 75, 79, 284, 285, 286, 288
macrocephala 667 niger 193, 233, 246, 295
macrogyna 18, 28, 75, 79, 236, 237, 238, nigra 183
239, 241 nigripes 75, 79, 294, 295, 296, 297
Maculinea 3, 9, 17, 38 nitidula 667
maligna 667 nitidulus 663, 666

787
nodicornis 75, 188, 663 puerilis 76, 79, 284, 285, 286, 289, 663
Nothomyrmica 51, 653, 655 punctata 668
numidicus 160 punica. 668
nylanderi 665, 667 pusilla 668
obensis 75, 78, 138, 139 pyrenaea 76, 79, 191, 192, 193
oblonga 667 quadrispinosa 668
obscurata 667 Quercus 106
obsoleta 667 rebeli 271,272
ominosa 664, 667 recedens 668
opaca 267, 667 rediana 668
orientalis 75, 79, 170, 171, 236 reticulata 76, 79, 259, 260, 261, 263
Orthoptera 45 Rhododendron 106,206, 219
orthostila 210 rhynchophora 76, 662
Pachycondyla 227 Rhyncophora 662
pallida 667 rotondi 76, 78, 84, 85, 86, 138, 259, 260,
pallidinodis .668 263
pallidula 667 Rombonella 667
pallinodis 668 Rosa 321
parallela 667 rubida 662, 666, 667, 691
Paramyrmica 30, 52, 56 rubriceps 668
parvula 667 rufa 668
pechei 163 ruficeps 668
pechei 56, 75, 79, 163, 164, 165 rufitarsis 668
pedestris 668 rufotestacea 668
pellucida 668 rugiceps 668
perelegans 75, 79, 230, 294, 295, 296 rugifrons 668
persiana 73, 75, 78, 101, 102, 103 ruginodiformis 76, 78, 124, 125
pertinax 668 ruginodolaevinodis 76, 79, 133, 236, 237,
pharaonis 665 238, 239, 240
Pheidole 664, 666, 667, 668, 669 rugososmythiesi 76, 246, 247, 248
Pheidolini 51 rugulosoides 76, 79, 84, 145, 259, 260, 262,
Pheidologeton 668 264, 284, 285, 287, 288, 289
Phengaris 3, 4, 9, 17, 38, 86,190, 207, 233, rugulososcabrinodis 76, 79, 145, 146, 147,
240, 252, 271 426, 427
Phragmites 104 rupestria 246
pilinodi 667 ruzskyi 76, 79, 175, 177, 179, 180
pilinodis 667, 668 Salix 104
piloscapa 250 Saloia 143
pilosiscapus 76, 79, 85, 86, 98, 99, 252, samnitica 76, 79, 181, 182, 183, 474, 475
259, 260, 262, 263 sancta 76, 79, 145, 146, 259, 264, 284, 285,
Pinus 183, 327 286, 287, 288, 289
plana 76, 78, 116, 117, 118, 270 scabrinodoides 76, 79, 284, 285, 286, 287,
Plesiomyrmex 51, 690, 692 289, 295
Podomyrma 221, 664, 665, 667 scabrinodolobicornis 76, 79, 194, 196, 250,
Pogonomyrmex 51, 201, 690, 691 251,252
p o l i t a 668 scabrinodorugulosa 76, 663-
Ponerinae 689, 690 scabrinodosabuleti 76, 79, 250, 259, 260,
poneroides 668 261,263
Populus 104 scabrosa 668
pradella 76, 663 scalpturata 668
Pristomyrmex 665, 668, 669 schenckioides 76, 79, 266, 267, 270
Protomyrmica 51, 690, 692 scutellaris 664, 668
provancheri Secostruma 51

788
semipolita 669 svartshevskii 77, 78, 124, 125
Sifolinia 28, 30, 51, 52, 56, 57, 160, 163, sylvestris 327
164, 165, 181, 183 Symbiomyrma 30, 52, 56, 57, 61, 160, 163,
silvestrianum 77, 79, 284, 286, 289 164, 165, 183
silvestrii 77, 79, 179, 181, 236, 237, 239 taediosa 72, 77, 308, 309, 310
simillima 669 Tapinoma 147, 271, 307, 668
simpliciuscula 669 tarbinskii 77, 78, 124, 125
sinica 77, 78, 131, 132,278 tarda 669
slobodensis 77,113, 114,145, 245 Temnothorax 51, 664, 665, 666, 667, 668,
smythiesi 96, 101, 107, 124,135,137, 144, 669
159, 179, 247, 248, 249, 282, 283 terminalis 669
Solenopsis 665, 666, 667 tertiaria 664, 669
Solifugae 45 testaceopilosa 669
Sommimyrma 30, 52, 56 testaceus 77, 663
sontica 77, 79, 236, 237, 238, 239 Tetramoriini 51, 705
sordidula 308, 669 Tetramorium 79, 132, 147, 227, 271, 278,
speciolosa 77, 664 284, 289, 307, 308, 309, 664, 665, 666,
specularis 77, 79, 175, 176, 177, 178 667, 668, 669, 721, 729
Sphecomyrminae 689 thoracica 669
spinosum 160 tianshanica 77, 79, 158, 159
starki 77, 79, 185, 186, 187, 188, 189, 270 tigreensis 669
Stenamma 32, 665, 667 tipuna 77, 78, 92, 93
strandi 77, 139, 140, 664 trachylissa 669
striata 77, 79, 145, 284, 285, 286, 288 trinodis 669
striatula 669 tschuliensis 78, 79
structor 308 Ulex 196
subacuta 77, 126, 664 umbripennis 669
subalpina 77, 79,175, 176, 177, 178 ussuriensis 78, 662
subaltaica 77, 167, 664 vagans 230
subbrevispinosa 77, 79, 175, 176,177 vastator 669
sublaevis 669 venusta 669
subopaca Arnoldi 77, 79, 266, 267, 270 vexator 669
subopaca Smith F. 266, 669 vicaria 78, 79, 294, 295, 296, 298
sulcinodolobicornis 77, 663 Viola 253
sulcinodoruginodis 77, 79, 294, 295, 296, Vollenhovia 667, 668
297 vulgare 253
sulcinodorugulosa 77, 79, 244, 245 westwoodi 665, 667
sulcinodoscabrinodis Forel 77, 79, 294, wheeleri 693
295,297,323 whittmeri 739
sulcinodoscabrinodis Ruzsky 77, 79, 294, winterae 56, 78, 79, 163, 164, 165, 166
295,297,298 yessensis 157, 198, 691
suspiciosa 669 yoshiokai 78, 79, 236, 237, 238, 239

789