Sclerophyll: Difference between revisions
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'''Sclerophyll''' is a type of vegetation that has hard [[leaf|leaves]] and short internodes (the distance between leaves along the stem). The word comes from the Greek ''sclero'' (hard) and ''phyllon'' (leaf). Sclerophyllous plants occur in all parts of the world but are most typical of [[Australia]]. They are also prominent in the [[Mediterranean forests, woodlands, and shrub]] [[biome]] that covers the [[Mediterranean Basin]], [[California chaparral and woodlands|Californian woodlands]], [[Chilean Matorral]], the [[Cape Province]] of [[South Africa]], southern [[Western Australia]], and western [[Southern Australia]]. |
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[[File:Arroyo pescadero 03.JPG|thumb|right|260px|[[Chaparral]] vegetation in [[Southern California]]]] |
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[[File:Banksia integrifolia subsp. monticola leaves (8668633194).jpg|thumb|260px|The hard leaves of a ''[[Banksia integrifolia]]'' (notice the short internodes)]] |
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'''Sclerophyll''' is a type of vegetation that is adapted to long periods of dryness and heat. The plants feature hard [[leaf|leaves]], short [[Internode (botany)|internode]]s (the distance between leaves along the stem) and leaf orientation which is parallel or oblique to direct sunlight. The word comes from the Greek ''sklēros'' (hard) and ''phyllon'' (leaf). The term was coined by [[Andreas Franz Wilhelm Schimper|A.F.W. Schimper]] in 1898 (translated in 1903), originally as a synonym of [[xeromorph]], but the two words were later differentiated.<ref>{{Cite book |url=https://www.cambridge.org/core/books/australian-rainforests/sclerophyll-problem/D63987C538BBFD20772BC543343B194C |title=Australian Rainforests: Islands of Green in a Land of Fire |date=2000 |publisher=Cambridge University Press |isbn=978-0-521-46568-7 |editor-last=Bowman |editor-first=D. M. J. S. |location=Cambridge |pages=48–67 |doi=10.1017/cbo9780511583490.004}}</ref> |
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Sclerophyllous plants occur in many parts of the world,<ref>C. Michael Hogan. 2010. [http://www.eoearth.org/article/Leather_Oak ''Leather Oak, Quercus durata''. Encyclopedia of Earth. National Council for Science and Environment]. Washington, DC</ref> but are most typical of areas with low rainfall or seasonal droughts, such as Australia, Africa, and western North and South America. They are prominent throughout [[Flora of Australia|Australia]], parts of [[Flora of Argentina|Argentina]], the [[Cerrado]] biogeographic region of [[Geography of Bolivia|Bolivia]], [[Geography of Paraguay|Paraguay]] and [[Flora of Brazil|Brazil]], and in the [[Mediterranean forests, woodlands, and scrub|Mediterranean biomes]] that cover the [[Mediterranean Basin]], [[California chaparral and woodlands|California]], [[Chilean Matorral|Chile]], and the [[Cape Province]] of [[South Africa]]. |
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In the [[Mediterranean basin]], [[Quercus ilex|holm oak]], [[cork oak]] and [[olive]]s are typical hardwood trees. In addition, there are several species of pine under the trees in the vegetation zone. The shrub layer contains numerous herbs such as [[rosemary]], [[thyme]] and [[lavender]]. In relation to the potential natural vegetation, around 2% of the Earth's land surface is covered by sclerophyll woodlands, and a total of 10% of all plant species on Earth live there. |
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==Description== |
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[[File:Dehesa_Los_Valhondos_Jamon_BEHER_Bernardo_Hernandez_Salamanca_Pelayos.jpg|250px|thumb|Sclerophyllous woodland in Spain]] |
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Sclerophyll [[woody plant]]s are characterized by their relatively small, stiff, leathery and long-lasting leaves. The sclerophyll vegetation is the result of an adaptation of the flora to the summer dry period of a [[Mediterranean climate|Mediterranean-type climate]]. |
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Plant species with this type of adaptation tend to be [[evergreen]] with great longevity, slow growth and with no loss of leaves during the unfavorable season. As a result, the [[thicket]]s that make up these [[ecosystem]]s are of the persistent evergreen type, in addition to the predominance of plants, even [[herbaceous]] ones, with "hard" leaves, which are covered by a thick leathery layer called the [[cuticle]], that prevents water loss during the dry season. The aerial and underground structures of these plants are modified to make up for water shortages that may affect their survival. |
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The name sclerophyll derives from the highly developed [[sclerenchyma]] from the plant, which is responsible for the hardness or stiffness of the leaves. This structure of the leaves inhibits transpiration and thus prevents major water losses during the dry season. Most of the plant species in the sclerophyll zone are not only insensitive to summer drought, they have also used various strategies to adapt to frequent [[wildfires]], heavy rainfall and [[Soil nutrient|nutrient]] deficiencies.<ref>{{Cite book |title=Farbatlas Ökosysteme der Erde: natürliche, naturnahe und künstliche Land-Ökosysteme aus geobotanischer Sicht |date=1997 |publisher=Ulmer |isbn=978-3-8001-3489-2 |editor-last=Grabherr |editor-first=Georg |location=Stuttgart}}</ref> |
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==Ecology== |
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[[File:San Rossore, Riserva del Paduletto.jpg|thumb|250px|Sclerophyll woodland in [[Tuscany]], Italy]] |
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The [[Type species|type]] of sclerophyllic trees in the [[Palearctic]] flora region include the holm oak (''[[Quercus ilex]]''), myrtle (''[[Myrtus communis]]''), strawberry tree (''[[Arbutus unedo]]''), wild olive (''[[Olea europaea]]''), laurel (''[[Laurus nobilis]]''), mock privet (''[[Phillyrea latifolia]]''), the Italian buckthorn (''[[Rhamnus alaternus]]''), etc.<ref>{{Cite book |last=Pott |first=Richard |title=Allgemeine Geobotanik: Biogeosysteme und Biodiversität |date=2005 |publisher=Springer |isbn=978-3-540-23058-8 |series=Springer-Lehrbuch |location=Berlin Heidelberg |pages=353–398}}</ref> |
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* In central and southern California, the coastal hills are covered in sclerophyll vegetation known as chaparral. The flora of this ecoregion also includes tree species [[Quercus dumosa|Scrub oak]] (''Quercus dumosa''), [[Aesculus californica|California buckeye]] (''Aesculus californica''), [[Dudleya densiflora|San Gabriel Mountain liveforever]] (''Dudlea densiflora''), [[Cercocarpus traskiae|Catalina mahogany]] (''Cercocarpus traskiae''), and the threatened [[Streptanthus albidus|jewelflower]] (''Streptanthus albidus'' ssp. ''Peramoenus'').<ref name="wwf">{{Cite web |title=California Chaparral & Woodlands |url=http://wwf.panda.org/about_our_earth/ecoregions/california_chaparral_woodlands.cfm |url-status=dead |archive-url=https://web.archive.org/web/20121008083831/http://wwf.panda.org/about_our_earth/ecoregions/california_chaparral_woodlands.cfm |archive-date=October 8, 2012 |access-date=2012-06-15 |publisher=World Wildlife Fund}} (material included verbatim under the [https://creativecommons.org/licenses/by-sa/3.0/deed.en CC BY-SA 3.0 license])</ref> |
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* In [[South Africa]], in the Cape region, there are Mediterranean open forests known as [[fynbos]]. The abundance of endemics is so extraordinary (68% of the 8600 vascular plant species in the area) that the South African sclerophyll area, the cape flora, forms the smallest of the six flora kingdoms on earth. Plants include ''[[Elegia (plant)|Elegia]]'', ''[[Thamnochortus]]'', and ''[[Willdenowia (plant)|Willdenowia]]'' and proteas such as king protea (''[[Protea cynaroides]]'') and blushing bride (''[[Serruria florida]]''). |
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* In most of Australia, sclerophyll vegetation such as [[eucalyptus]] trees, [[melaleuca]]s, [[banksia]]s, [[callistemon]]s and [[grevillea]]s dominate the [[Mallee (habit)|mallee]] and woodland areas of its cities, including those lacking a Mediterranean climate, such as [[Sydney]], [[Melbourne]], [[Hobart]] and [[Brisbane]].<ref name="duncanbrown">{{Cite journal |last=Duncan |first=F |last2=Brown |first2=M.J |date=1985 |title=Dry Sclerophyll vegetation in Tasmania |journal=Wildlife Division Technical Report 85/1- National Parks and Wildlife Services}}</ref><ref>{{Cite web |title=Sydney Coastal Dry Sclerophyll Forests |url=http://www.environment.nsw.gov.au/ThreatenedSpeciesApp/VegClass.aspx?vegClassName=Sydney+Coastal+Dry+Sclerophyll+Forests |access-date=September 17, 2012 |publisher=NSW Environment & Heritage}}</ref> |
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* In Chile, south of the desert areas, there is evergreen bushland called [[matorral]]. Typical species include Litre [[Lithraea|''(Lithraea venenosa)'']], Quillay or Soapbark Tree ''([[Quillaja saponaria]])'', and bromeliads of genus ''[[Puya (genus)|Puya]]''.<ref>{{Cite web |last=Hogan |first=C. Michael |date=2012-10-17 |title=Chilean Wine Palm (Jubaea chilensis) |url=http://globaltwitcher.auderis.se/artspec_information.asp?thingid=82831 |url-status=dead |archive-url=https://web.archive.org/web/20121017013207/http://globaltwitcher.auderis.se/artspec_information.asp?thingid=82831 |archive-date=2012-10-17 |access-date=2023-11-19}}</ref> |
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==Climate== |
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[[File:Brenanparkbush.jpg|thumb|250px|A sclerophyll bushland in [[Sydney]] (which falls in the [[humid subtropical]] zone)]] |
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The sclerophyll regions are located in the outer subtropics bordering the temperate zone (also known as the warm-temperate zone). Accordingly, the annual average temperatures are relatively high at {{Convert|12-24|C}}; An average of over {{Convert|18|C}} is reached for at least four months, eight to twelve months it is over {{Convert|10|C}} and no month is below {{Convert|5|C}} on average. [[Frost]] and snow occur only occasionally and the [[growing season]] lasts longer than 150 days and is in the winter half-year.<ref>{{Cite book |last=Pfadenhauer |first=Jörg |title=Vegetation der Erde: Grundlagen, Ökologie, Verbreitung |last2=Klötzli |first2=Frank |date=2014 |publisher=Springer Spektrum |isbn=978-3-642-41949-2 |location=Berlin Heidelberg |pages=289–291}}</ref> The lower limit of the moderate annual precipitation is {{Convert|300|mm|abbr=on}} ([[semi-arid climate]]) and the upper limit {{Convert|900-1000|mm|abbr=on}}. |
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Generally, the summers are dry and hot with a [[dry season]] of a maximum of seven months, but at least two to three months. The winters are rainy and cool. However, not all regions with sclerophyll vegetation feature the classic [[Mediterranean climate]]; parts of eastern Italy, eastern Australia and eastern South Africa, which feature sclerophyll woodlands, tend to have uniform rainfall or even a more summer-dominant rainfall, whereby falling under the [[humid subtropical climate]] zone (''Cfa''/''Cwa''). Furthermore, other areas with sclerophyll flora would grade to the [[oceanic climate]] (''Cfb''); particularly the eastern parts of the [[Eastern Cape]] province in South Africa, and [[Tasmania]], [[Victoria (state)|Victoria]] and southern [[New South Wales]] in Australia.<ref>{{Cite book |last=Schultz |first=Jürgen |title=Handbuch der Ökozonen |date=2000 |publisher=Ulmer |isbn=978-3-8252-8200-4 |series=UTB für Wissenschaft Geographie, Ökologie, Agrar- und Forstwissenschaften |location=Stuttgart |pages=35, 45-46}}</ref> |
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==Soils== |
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Sclerophyll plants are also found in areas with nutrient-poor and acidic soils, and soils with heavy concentrations of aluminum and other metals. Sclerophyll leaves transpire less and have a lower {{CO2}} uptake than malacophyllous or laurophyllous leaves. These lower transpiration rates may reduce the uptake of toxic ions and better provide for C-carboxylation under nutrient-poor conditions, particularly low availability of mineral nitrogen and phosphate. Sclerophyllous plants are found in tropical heath forests, which grown on nutrient-poor sandy soils in humid regions in the Rio Orinoco and the Rio Negro basins of northern South America on quartz sand, in the [[kerangas]] forests of Borneo and on the Malay Peninsula, in coastal sandy areas along the [[Gulf of Guinea]] in Gabon, Cameroon, and Côte d'Ivoire, and in eastern Australia. Since water drains rapidly through these soils, sclerophylly also protects plants against drought stress during dry periods.<ref name="fundamentals">{{Cite book |last=Pfadenhauer |first=Jörg S. |url=https://doi.org/10.1007/978-3-030-49860-3_1 |title=Fundamentals towards Understanding Global Vegetation |last2=Klötzli |first2=Frank A. |date=2020 |publisher=Springer International Publishing |isbn=978-3-030-49859-7 |location=Cham |pages=1–120 |language=en |doi=10.1007/978-3-030-49860-3_1}}</ref><ref name="zonal">{{Cite book |last=Pfadenhauer |first=Jörg S. |url=https://doi.org/10.1007/978-3-030-49860-3_2 |title=Zonal Vegetation of the Tropical Zone with Year-Round Rain |last2=Klötzli |first2=Frank A. |date=2020 |publisher=Springer International Publishing |isbn=978-3-030-49859-7 |location=Cham |pages=121–177 |language=en |doi=10.1007/978-3-030-49860-3_2}}</ref> |
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Sclerophylly's advantages in nutrient-poor conditions may be another factor in the prevalence of sclerophyllous plants in nutrient-poor areas in drier-climate regions, like much of Australia and the [[Cerrado]] of Brazil.<ref name = fundamentals/> |
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==Distribution== |
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The zone of the sclerophyll vegetation lies in the border area between the [[subtropics]] and the [[temperate zone]], approximately between the 30th and 40th degree of [[latitude]] (in the northern hemisphere also up to the 45th degree of latitude). Their presence is limited to the coastal western sides of the continents, but nonetheless can typical in any regions of a continent with scarce annual precipitation or frequent seasonal droughts and poor soils that are heavily leached.<ref>Susanne Heise: Vegetation zones: The zone of the evergreen hardwood plants, in TERRA-Online/Realschule on Klett.de, Klett, Leipzig 2003, version January 26, 2006, accessed on December 17, 2020.</ref> |
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The sclerophyll zone often merges into [[temperate deciduous forest]]s towards the poles, on the coasts also into [[temperate rainforest]]s and towards the equator in hot [[semi-desert]]s or deserts. The Mediterranean areas, which have a very high [[biodiversity]], are under great pressure from the population. This is especially true for the Mediterranean region since ancient times. Through [[overexploitation]] (logging, grazing, agricultural use) and frequent fires caused by people, the original forest vegetation is converted. In extreme cases, the hard-leaf vegetation disappears completely and is replaced by open rock [[heaths]]. |
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[[File:Spiny forest 1, Ifaty, Madagascar.jpg|thumb|250px|Sclerophyll shrubland in southwestern Madagascar]] |
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Some sclerophyll areas are closer to the equator than the Mediterranean zone—for example, the interior of [[Madagascar]], the dry half of [[New Caledonia]], the lower edge areas of the [[Madrean pine-oak woodlands]] of the [[Sierra Madre Occidental|Mexican highlands]] between 800 and 1800/2000 m or around 2000 m high plateaus of the [[Asir Mountains]] on the western edge of the [[Arabian Peninsula]].<ref>{{Cite book |title=Atlas zur Biogeographie |date=1976 |publisher=Bibliograph. Inst |isbn=978-3-411-00303-7 |editor-last=Schmithüsen |editor-first=Josef |series=Meyers großer physischer Weltatlas |location=Mannheim |pages=19, 43}}</ref> |
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===Land use=== |
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While the winter rain areas of America, South Africa and [[Australia]], with an unusually large variety of [[food crops]], were ideal gathering areas for [[hunter gatherer]]s until [[European colonization]], agriculture and [[cattle breeding]] spread in the Mediterranean area since the [[Neolithic]], which permanently changed the face of the landscape. In the sclerophyll regions near the coast, permanent crops such as olive and wine cultivation established themselves; However, the landscape forms that characterize the degenerate shrubbery and shrub heaths [[Macchie]] and [[Garigue]] are predominantly a result of [[grazing]] (especially with goats). |
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In the course of the last millennia, the original vegetation in almost all areas of this vegetation zone has been greatly changed by the influence of humans. Where the plants have not been replaced by [[vineyard]]s and olive [[grove (nature)|grove]]s, the [[maquis shrubland|maquis]] was the predominant form of vegetation on the Mediterranean. The maquis has been degraded in many places to the low shrub heather, the [[garigue]]. Many plant species that are rich in aromatic oils belong to both vegetation societies. The diversity of the original sclerophyll vegetation in the world is high to extremely high (3000–5000 species per ha).<ref>Klaus Müller-Hohenstein: The geo-ecological zones of the Earth. In: ''Geography and School''. Issue 59, Bayreuth 1989.</ref> |
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==Australian bush== |
==Australian bush== |
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{{see also|Flora of Australia}} |
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Arid, largely treeless areas aside, most [[Australia]]n bushland is '''sclerophyll''' forest. Common plants include the [[Proteaceae]] ([[Grevillea]]s, [[Banksia]]s and Proteas), tea-trees, [[Acacia]]s, [[Boronia]]s, and the [[Eucalypt]]s. |
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[[Image:Bush Eagle-Bay WA.jpg|thumb|left|Bush around Eagle Bay, [[Western Australia]] ]] |
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Most areas of the [[Australian continent]] able to support woody plants are occupied by sclerophyll communities as [[forest]]s, [[savanna]]s, or [[Heath (habitat)|heathlands]]. Common plants include the [[Proteaceae]] ([[grevillea]]s, [[banksia]]s and [[hakea]]s), [[Melaleuca|tea-trees]], [[acacia]]s, [[boronia]]s, and [[eucalypt]]s. |
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The most common sclerophyll communities in Australia are [[savanna]]s dominated by grasses with an overstorey of eucalypts and acacias. Acacia (particularly [[Acacia aneura|mulga]]) [[shrubland]]s also cover extensive areas. All the dominant overstorey acacia species and a majority of the understorey acacias have a scleromorphic adaptation in which the leaves have been reduced to [[phyllode]]s consisting entirely of the [[Petiole (botany)|petiole]].<ref>{{Cite book |last=Simmons |first=Marion H. |title=Acacias of Australia |date=1982 |publisher=Nelson |isbn=978-0-17-005779-0 |edition=Reprint |location=Melbourne, Vic}}</ref> |
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The ubiquitous ''dry sclerophyll forests'' covered vast areas of the continent (and to a much lesser extent still do). They have a eucalyptus overstory (10 to 30 metres) and the understory is also hard-leaved. Often seeming drab and barren to the casual European eye, dry sclerophyll forest is highly complex and diverse, with up to twice as many different plant species per square metre as [[rainforest]], and high animal diversity too. (If at first this seems counter-intuitive, remember that, as a broad generalisation, harsher conditions promote speciation and diversity, with every tiny variation in soil, light, or moisture offering a niche, where rich soils and ample water tend to simply favour species which can take advantage of them most rapidly.) |
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Many plants of the sclerophyllous woodlands and shrublands also produce leaves unpalatable to herbivores by the inclusion of toxic and indigestible compounds which assure survival of these long-lived leaves. This trait is particularly noticeable in the eucalypt and ''[[Melaleuca]]'' species which possess oil glands within their leaves that produce a pungent volatile oil that makes them unpalatable to most browsers.<ref>{{Cite book |last=Florence |first=Ross G. |title=Ecology and silviculture of eucalypt forests |date=2007 |publisher=CSIRO Publ |isbn=978-0-643-05799-9 |location=Collingwood}}</ref> These traits make the majority of woody plants in these woodlands largely unpalatable to domestic livestock.<ref>Mott, J. J., and J. C. Tothill. 1984. Tropical and subtropical woodlands. in G. N. Harrington and A. D. Wilson, editors. Management of Australia's Rangelands. CSIRO Publishing, Melbourne</ref> It is therefore important from a grazing perspective that these woodlands support a more or less continuous layer of herbaceous ground cover dominated by grasses. |
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The less extensive ''wet sclerophyll forests'' have a taller eucalyptus overstory, 30 metres or more (typically [[Eucalyptus regnans|Mountain Ash]], [[Eucalyptus delegatensis|Alpine Ash]], [[Messmate Stringybark]] or [[Manna Gum]]), and a soft-leaved, fairly open understory ([[tree fern]]s are common). They require ample rainfall — at least 1000mm (40 inches). |
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Sclerophyll [[forests]] cover a much smaller area of the continent, being restricted to relatively high rainfall locations. They have a eucalyptus overstory (10 to 30 metres) with the understory also being hard-leaved. ''Dry sclerophyll'' forests are the most common forest type on the continent, and although it may seem barren dry sclerophyll forest is highly diverse. For example, a study of sclerophyll vegetation in Seal Creek, [[Victoria (Australia)|Victoria]], found 138 species.<ref>{{Cite journal |last=Parsons R. F. |last2=Cameron D .G. |year=1974 |title=Maximum Plant Species Diversity in Terrestrial Communities |journal=Biotropica |publisher=The Association for Tropical Biology and Conservation |volume=6 |issue=3 |page=202 |doi=10.2307/2989653 |jstor=2989653}}</ref> |
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==History== |
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Sclerophyllous plants are all apart of a specific environmentand are anything but newcomers — the [[Proteaceae]] family dates back 80 million years to the late [[Cretaceous]]—but sclerophyll forests did not start becoming a major part of the Australian landscape until around 15 million years ago. By the time of European settlement, sclerophyll forest accounted for the vast bulk of the forested areas. |
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Even less extensive are ''wet sclerophyll'' forests. They have a taller eucalyptus overstory than dry sclerophyll forests, {{Convert|30|m|abbr=}} or more (typically [[Eucalyptus regnans|mountain ash]], [[Eucalyptus delegatensis|alpine ash]], [[Eucalyptus grandis|rose gum]], [[Eucalyptus diversicolor|karri]], [[Eucalyptus obliqua|messmate stringybark]], or [[Eucalyptus viminalis|manna gum]], and a soft-leaved, fairly dense understory ([[tree fern]]s are common). They require ample rainfall—at least 1000 mm (40 inches). |
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Most of the wooded parts of present-day Australia have become sclerophyll forest as a result of the extreme age of the continent. Deep [[weathering]] of the crust over many millions of years leached chemicals out of the rock, leaving Australian soils deficient in nutrients, particularly [[phosphorus]]. (Small areas, mostly in the south, have benefited from relatively recent volcanic activity and have fairly rich soils as a result.) |
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===Evolution=== |
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Sclerophyllous plants generally resist dry conditions well, making them successful in areas of [[mediterranean climate]] with rainy winters but hot dry summers. In Australia, however, they evolved in response to the low level of phosphorus in the soil — indeed, many Australian native plants cannot tolerate higher levels of phosphorus and will die if fertilised incorrectly. The leaves are hard due to [[lignin]], which prevents wilting and allows plants to grow even when there isn't enough phosphorus for substantial new cell growth (Major, 2003). |
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[[File:Yellow_box_2.jpg|thumb|250px|Yellow Box (''[[Eucalyptus melliodora]]'') in a sclerophyll woodland, [[Melbourne]] (''[[Oceanic climate|Cfb]]'' climate)]] |
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Sclerophyllous plants are all part of a specific environment and are anything but newcomers. By the time of European settlement, sclerophyll forest accounted for the vast bulk of the forested areas. |
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Most of the wooded parts of present-day Australia have become sclerophyll dominated as a result of the extreme age of the continent combined with [[Indigenous Australians|Aboriginal]] fire use. Deep [[weathering]] of the crust over many millions of years leached chemicals out of the rock, leaving Australian soils deficient in nutrients, particularly [[phosphorus]]. Such nutrient deficient soils support non-sclerophyllous plant communities elsewhere in the world and did so over most of Australia prior to European arrival. However such deficient soils cannot support the nutrient losses associated with frequent fires and are rapidly replaced with sclerophyllous species under traditional Aboriginal burning regimens. With the cessation of traditional burning non-sclerophyllous species have re-colonized sclerophyll habitat in many parts of Australia.{{Citation needed|date=November 2014}} |
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==References== |
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* {{cite web|url = http://www.amonline.net.au/factsheets/sclerophyll_forests.htm | title = Sclerophyll forests | author = R. Major | year = 2003 | publisher = Australian Museum | accessdate = 2005-02-14 }} |
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The presence of toxic compounds combined with a high carbon : nitrogen ratio make the leaves and branches of scleromorphic species long-lived in the litter, and can lead to a large build-up of litter in woodlands.<ref>{{Cite book |title=The greening of Gondwana |date=1988 |publisher=Reed Books |isbn=978-0-7301-0154-3 |editor-last=White |editor-first=Mary E. |edition=Repr |location=French's Forest |editor-last2=Frazier |editor-first2=Jim}}</ref><ref>{{Cite book |last=Flannery |first=Tim F. |title=The future eaters: an ecological history of the Australasian lands and people |date=2002 |publisher=Grove Press |isbn=978-0-8021-3943-6 |edition=1 |location=New York, NY}}</ref> The toxic compounds of many species, notably Eucalyptus species, are volatile and flammable and the presence of large amounts of flammable litter, coupled with an herbaceous understorey, encourages fire.<ref>{{Cite journal |last=Burrows |first=W. H. |last2=Carter |first2=J. O. |last3=Scanlan |first3=J. C. |last4=Anderson |first4=E. R. |date=1990 |title=Management of Savannas for Livestock Production in North-East Australia: Contrasts Across the Tree-Grass Continuum |url=https://www.jstor.org/stable/2845383 |journal=Journal of Biogeography |volume=17 |issue=4/5 |pages=503–512 |doi=10.2307/2845383 |issn=0305-0270}}</ref> |
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[[Category:Ecology]] |
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All the Australian sclerophyllous communities are liable to be burnt with varying frequencies and many of the woody plants of these woodlands have developed adaptations to survive and minimise the effects of fire.<ref>Harrington, G. N., M. H. Friedel, K. C. Hodgkinson, and J. C. Noble. 1984. Vegetation ecology and management. in G. N. Harrington and A. D. Wilson, editors. Management of Australia's Rangelands. CSIRO Publishing, Melbourne.</ref> |
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[[de:Hartlaubvegetation]] |
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[[ja:硬葉樹林]] |
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Sclerophyllous plants generally resist dry conditions well, making them successful in areas of seasonally variable rainfall. In Australia, however, they evolved in response to the low level of phosphorus in the soil—indeed, many native Australian plants cannot tolerate higher levels of phosphorus and will die if fertilised incorrectly. The leaves are hard due to [[lignin]], which prevents [[wilting]] and allows plants to grow, even when there is not enough phosphorus for substantial new cell growth.<ref>{{Cite web |last=R. Major |year=2003 |title=Sclerophyll forests |url=http://www.amonline.net.au/factsheets/sclerophyll_forests.htm |access-date=February 14, 2005 |publisher=Australian Museum}}</ref> |
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==Regions== |
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[[File:כסא אליהו בהר מירון.jpg|thumb|250px|Sclerophyll vegetation on [[Mount Meron]] in [[Israel]]]] |
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[[File:High atlas mountains.jpg|thumb|250px|Hard-leaved vegetation in the [[High Atlas]] mountains, [[Morocco]]]] |
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These are the biomes or ecoregions in the world that feature an abundance of, or are known for having, sclerophyll vegetation: |
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*[[Cumberland Plain Woodland]] |
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*[[Sydney Sandstone Ridgetop Woodland]] |
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*[[Eastern Suburbs Banksia Scrub]] |
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*[[Tasmanian dry sclerophyll forests]] |
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*[[Aegean and Western Turkey sclerophyllous and mixed forests]] |
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*[[California chaparral and woodlands]] |
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*[[California coastal sage and chaparral]] |
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*[[Chilean Matorral]] |
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*[[Mallee Woodlands and Shrublands]] |
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*[[Italian sclerophyllous and semi-deciduous forests]] |
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*[[Eastern Mediterranean conifer–sclerophyllous–broadleaf forests]] |
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*[[Southwest Iberian Mediterranean sclerophyllous and mixed forests]] |
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*[[Tyrrhenian–Adriatic sclerophyllous and mixed forests]] |
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*[[Canary Islands dry woodlands and forests]] |
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*[[Mediterranean acacia–argania dry woodlands]] |
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*[[Mediterranean dry woodlands and steppe]] |
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*[[Southeastern Iberian shrubs and woodlands]] |
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*[[Cyprus Mediterranean forests]] |
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*[[Crete Mediterranean forests]] |
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*[[Cape Floristic Region]] |
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*[[Southern Anatolian montane conifer and deciduous forests]] |
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*[[Albany thickets]] |
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*[[Northwest Iberian montane forests]] |
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==See also== |
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*[[Mediterranean forests, woodlands, and scrub]] |
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*[[Chaparral]] |
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*[[Fynbos]] |
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*[[Maquis shrubland]] |
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*[[Garrigue]] |
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*[[Kwongan]] |
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*[[Matorral]] |
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*[[Barren vegetation]] |
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==References== |
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{{reflist|2}}{{Vegetation}} |
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[[Category:Mediterranean forests, woodlands, and scrub]] |
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[[Category:California chaparral and woodlands]] |
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[[Category:Flora of the Chilean Matorral]] |
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[[Category:Mallee Woodlands and Shrublands]] |
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[[Category:Ecology]] |
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[[Category:Sclerophyll forests]] |
Latest revision as of 07:27, 30 March 2024
![](http://upload.wikimedia.org/wikipedia/commons/thumb/7/77/Banksia_integrifolia_subsp._monticola_leaves_%288668633194%29.jpg/260px-Banksia_integrifolia_subsp._monticola_leaves_%288668633194%29.jpg)
Sclerophyll is a type of vegetation that is adapted to long periods of dryness and heat. The plants feature hard leaves, short internodes (the distance between leaves along the stem) and leaf orientation which is parallel or oblique to direct sunlight. The word comes from the Greek sklēros (hard) and phyllon (leaf). The term was coined by A.F.W. Schimper in 1898 (translated in 1903), originally as a synonym of xeromorph, but the two words were later differentiated.[1]
Sclerophyllous plants occur in many parts of the world,[2] but are most typical of areas with low rainfall or seasonal droughts, such as Australia, Africa, and western North and South America. They are prominent throughout Australia, parts of Argentina, the Cerrado biogeographic region of Bolivia, Paraguay and Brazil, and in the Mediterranean biomes that cover the Mediterranean Basin, California, Chile, and the Cape Province of South Africa.
In the Mediterranean basin, holm oak, cork oak and olives are typical hardwood trees. In addition, there are several species of pine under the trees in the vegetation zone. The shrub layer contains numerous herbs such as rosemary, thyme and lavender. In relation to the potential natural vegetation, around 2% of the Earth's land surface is covered by sclerophyll woodlands, and a total of 10% of all plant species on Earth live there.
Description
[edit]![](http://upload.wikimedia.org/wikipedia/commons/thumb/2/28/Dehesa_Los_Valhondos_Jamon_BEHER_Bernardo_Hernandez_Salamanca_Pelayos.jpg/250px-Dehesa_Los_Valhondos_Jamon_BEHER_Bernardo_Hernandez_Salamanca_Pelayos.jpg)
Sclerophyll woody plants are characterized by their relatively small, stiff, leathery and long-lasting leaves. The sclerophyll vegetation is the result of an adaptation of the flora to the summer dry period of a Mediterranean-type climate.
Plant species with this type of adaptation tend to be evergreen with great longevity, slow growth and with no loss of leaves during the unfavorable season. As a result, the thickets that make up these ecosystems are of the persistent evergreen type, in addition to the predominance of plants, even herbaceous ones, with "hard" leaves, which are covered by a thick leathery layer called the cuticle, that prevents water loss during the dry season. The aerial and underground structures of these plants are modified to make up for water shortages that may affect their survival.
The name sclerophyll derives from the highly developed sclerenchyma from the plant, which is responsible for the hardness or stiffness of the leaves. This structure of the leaves inhibits transpiration and thus prevents major water losses during the dry season. Most of the plant species in the sclerophyll zone are not only insensitive to summer drought, they have also used various strategies to adapt to frequent wildfires, heavy rainfall and nutrient deficiencies.[3]
Ecology
[edit]![](http://upload.wikimedia.org/wikipedia/commons/thumb/2/27/San_Rossore%2C_Riserva_del_Paduletto.jpg/250px-San_Rossore%2C_Riserva_del_Paduletto.jpg)
The type of sclerophyllic trees in the Palearctic flora region include the holm oak (Quercus ilex), myrtle (Myrtus communis), strawberry tree (Arbutus unedo), wild olive (Olea europaea), laurel (Laurus nobilis), mock privet (Phillyrea latifolia), the Italian buckthorn (Rhamnus alaternus), etc.[4]
- In central and southern California, the coastal hills are covered in sclerophyll vegetation known as chaparral. The flora of this ecoregion also includes tree species Scrub oak (Quercus dumosa), California buckeye (Aesculus californica), San Gabriel Mountain liveforever (Dudlea densiflora), Catalina mahogany (Cercocarpus traskiae), and the threatened jewelflower (Streptanthus albidus ssp. Peramoenus).[5]
- In South Africa, in the Cape region, there are Mediterranean open forests known as fynbos. The abundance of endemics is so extraordinary (68% of the 8600 vascular plant species in the area) that the South African sclerophyll area, the cape flora, forms the smallest of the six flora kingdoms on earth. Plants include Elegia, Thamnochortus, and Willdenowia and proteas such as king protea (Protea cynaroides) and blushing bride (Serruria florida).
- In most of Australia, sclerophyll vegetation such as eucalyptus trees, melaleucas, banksias, callistemons and grevilleas dominate the mallee and woodland areas of its cities, including those lacking a Mediterranean climate, such as Sydney, Melbourne, Hobart and Brisbane.[6][7]
- In Chile, south of the desert areas, there is evergreen bushland called matorral. Typical species include Litre (Lithraea venenosa), Quillay or Soapbark Tree (Quillaja saponaria), and bromeliads of genus Puya.[8]
Climate
[edit]![](http://upload.wikimedia.org/wikipedia/commons/thumb/f/f8/Brenanparkbush.jpg/250px-Brenanparkbush.jpg)
The sclerophyll regions are located in the outer subtropics bordering the temperate zone (also known as the warm-temperate zone). Accordingly, the annual average temperatures are relatively high at 12–24 °C (54–75 °F); An average of over 18 °C (64 °F) is reached for at least four months, eight to twelve months it is over 10 °C (50 °F) and no month is below 5 °C (41 °F) on average. Frost and snow occur only occasionally and the growing season lasts longer than 150 days and is in the winter half-year.[9] The lower limit of the moderate annual precipitation is 300 mm (12 in) (semi-arid climate) and the upper limit 900–1,000 mm (35–39 in).
Generally, the summers are dry and hot with a dry season of a maximum of seven months, but at least two to three months. The winters are rainy and cool. However, not all regions with sclerophyll vegetation feature the classic Mediterranean climate; parts of eastern Italy, eastern Australia and eastern South Africa, which feature sclerophyll woodlands, tend to have uniform rainfall or even a more summer-dominant rainfall, whereby falling under the humid subtropical climate zone (Cfa/Cwa). Furthermore, other areas with sclerophyll flora would grade to the oceanic climate (Cfb); particularly the eastern parts of the Eastern Cape province in South Africa, and Tasmania, Victoria and southern New South Wales in Australia.[10]
Soils
[edit]Sclerophyll plants are also found in areas with nutrient-poor and acidic soils, and soils with heavy concentrations of aluminum and other metals. Sclerophyll leaves transpire less and have a lower CO2 uptake than malacophyllous or laurophyllous leaves. These lower transpiration rates may reduce the uptake of toxic ions and better provide for C-carboxylation under nutrient-poor conditions, particularly low availability of mineral nitrogen and phosphate. Sclerophyllous plants are found in tropical heath forests, which grown on nutrient-poor sandy soils in humid regions in the Rio Orinoco and the Rio Negro basins of northern South America on quartz sand, in the kerangas forests of Borneo and on the Malay Peninsula, in coastal sandy areas along the Gulf of Guinea in Gabon, Cameroon, and Côte d'Ivoire, and in eastern Australia. Since water drains rapidly through these soils, sclerophylly also protects plants against drought stress during dry periods.[11][12]
Sclerophylly's advantages in nutrient-poor conditions may be another factor in the prevalence of sclerophyllous plants in nutrient-poor areas in drier-climate regions, like much of Australia and the Cerrado of Brazil.[11]
Distribution
[edit]The zone of the sclerophyll vegetation lies in the border area between the subtropics and the temperate zone, approximately between the 30th and 40th degree of latitude (in the northern hemisphere also up to the 45th degree of latitude). Their presence is limited to the coastal western sides of the continents, but nonetheless can typical in any regions of a continent with scarce annual precipitation or frequent seasonal droughts and poor soils that are heavily leached.[13]
The sclerophyll zone often merges into temperate deciduous forests towards the poles, on the coasts also into temperate rainforests and towards the equator in hot semi-deserts or deserts. The Mediterranean areas, which have a very high biodiversity, are under great pressure from the population. This is especially true for the Mediterranean region since ancient times. Through overexploitation (logging, grazing, agricultural use) and frequent fires caused by people, the original forest vegetation is converted. In extreme cases, the hard-leaf vegetation disappears completely and is replaced by open rock heaths.
![](http://upload.wikimedia.org/wikipedia/commons/thumb/4/4a/Spiny_forest_1%2C_Ifaty%2C_Madagascar.jpg/250px-Spiny_forest_1%2C_Ifaty%2C_Madagascar.jpg)
Some sclerophyll areas are closer to the equator than the Mediterranean zone—for example, the interior of Madagascar, the dry half of New Caledonia, the lower edge areas of the Madrean pine-oak woodlands of the Mexican highlands between 800 and 1800/2000 m or around 2000 m high plateaus of the Asir Mountains on the western edge of the Arabian Peninsula.[14]
Land use
[edit]While the winter rain areas of America, South Africa and Australia, with an unusually large variety of food crops, were ideal gathering areas for hunter gatherers until European colonization, agriculture and cattle breeding spread in the Mediterranean area since the Neolithic, which permanently changed the face of the landscape. In the sclerophyll regions near the coast, permanent crops such as olive and wine cultivation established themselves; However, the landscape forms that characterize the degenerate shrubbery and shrub heaths Macchie and Garigue are predominantly a result of grazing (especially with goats).
In the course of the last millennia, the original vegetation in almost all areas of this vegetation zone has been greatly changed by the influence of humans. Where the plants have not been replaced by vineyards and olive groves, the maquis was the predominant form of vegetation on the Mediterranean. The maquis has been degraded in many places to the low shrub heather, the garigue. Many plant species that are rich in aromatic oils belong to both vegetation societies. The diversity of the original sclerophyll vegetation in the world is high to extremely high (3000–5000 species per ha).[15]
Australian bush
[edit]![](http://upload.wikimedia.org/wikipedia/commons/thumb/f/f0/Bush_Eagle-Bay_WA.jpg/220px-Bush_Eagle-Bay_WA.jpg)
Most areas of the Australian continent able to support woody plants are occupied by sclerophyll communities as forests, savannas, or heathlands. Common plants include the Proteaceae (grevilleas, banksias and hakeas), tea-trees, acacias, boronias, and eucalypts.
The most common sclerophyll communities in Australia are savannas dominated by grasses with an overstorey of eucalypts and acacias. Acacia (particularly mulga) shrublands also cover extensive areas. All the dominant overstorey acacia species and a majority of the understorey acacias have a scleromorphic adaptation in which the leaves have been reduced to phyllodes consisting entirely of the petiole.[16]
Many plants of the sclerophyllous woodlands and shrublands also produce leaves unpalatable to herbivores by the inclusion of toxic and indigestible compounds which assure survival of these long-lived leaves. This trait is particularly noticeable in the eucalypt and Melaleuca species which possess oil glands within their leaves that produce a pungent volatile oil that makes them unpalatable to most browsers.[17] These traits make the majority of woody plants in these woodlands largely unpalatable to domestic livestock.[18] It is therefore important from a grazing perspective that these woodlands support a more or less continuous layer of herbaceous ground cover dominated by grasses.
Sclerophyll forests cover a much smaller area of the continent, being restricted to relatively high rainfall locations. They have a eucalyptus overstory (10 to 30 metres) with the understory also being hard-leaved. Dry sclerophyll forests are the most common forest type on the continent, and although it may seem barren dry sclerophyll forest is highly diverse. For example, a study of sclerophyll vegetation in Seal Creek, Victoria, found 138 species.[19]
Even less extensive are wet sclerophyll forests. They have a taller eucalyptus overstory than dry sclerophyll forests, 30 metres (98 ft) or more (typically mountain ash, alpine ash, rose gum, karri, messmate stringybark, or manna gum, and a soft-leaved, fairly dense understory (tree ferns are common). They require ample rainfall—at least 1000 mm (40 inches).
Evolution
[edit]![](http://upload.wikimedia.org/wikipedia/commons/thumb/4/44/Yellow_box_2.jpg/250px-Yellow_box_2.jpg)
Sclerophyllous plants are all part of a specific environment and are anything but newcomers. By the time of European settlement, sclerophyll forest accounted for the vast bulk of the forested areas.
Most of the wooded parts of present-day Australia have become sclerophyll dominated as a result of the extreme age of the continent combined with Aboriginal fire use. Deep weathering of the crust over many millions of years leached chemicals out of the rock, leaving Australian soils deficient in nutrients, particularly phosphorus. Such nutrient deficient soils support non-sclerophyllous plant communities elsewhere in the world and did so over most of Australia prior to European arrival. However such deficient soils cannot support the nutrient losses associated with frequent fires and are rapidly replaced with sclerophyllous species under traditional Aboriginal burning regimens. With the cessation of traditional burning non-sclerophyllous species have re-colonized sclerophyll habitat in many parts of Australia.[citation needed]
The presence of toxic compounds combined with a high carbon : nitrogen ratio make the leaves and branches of scleromorphic species long-lived in the litter, and can lead to a large build-up of litter in woodlands.[20][21] The toxic compounds of many species, notably Eucalyptus species, are volatile and flammable and the presence of large amounts of flammable litter, coupled with an herbaceous understorey, encourages fire.[22]
All the Australian sclerophyllous communities are liable to be burnt with varying frequencies and many of the woody plants of these woodlands have developed adaptations to survive and minimise the effects of fire.[23]
Sclerophyllous plants generally resist dry conditions well, making them successful in areas of seasonally variable rainfall. In Australia, however, they evolved in response to the low level of phosphorus in the soil—indeed, many native Australian plants cannot tolerate higher levels of phosphorus and will die if fertilised incorrectly. The leaves are hard due to lignin, which prevents wilting and allows plants to grow, even when there is not enough phosphorus for substantial new cell growth.[24]
Regions
[edit]![](http://upload.wikimedia.org/wikipedia/commons/thumb/e/e3/%D7%9B%D7%A1%D7%90_%D7%90%D7%9C%D7%99%D7%94%D7%95_%D7%91%D7%94%D7%A8_%D7%9E%D7%99%D7%A8%D7%95%D7%9F.jpg/250px-%D7%9B%D7%A1%D7%90_%D7%90%D7%9C%D7%99%D7%94%D7%95_%D7%91%D7%94%D7%A8_%D7%9E%D7%99%D7%A8%D7%95%D7%9F.jpg)
![](http://upload.wikimedia.org/wikipedia/commons/thumb/8/81/High_atlas_mountains.jpg/250px-High_atlas_mountains.jpg)
These are the biomes or ecoregions in the world that feature an abundance of, or are known for having, sclerophyll vegetation:
- Cumberland Plain Woodland
- Sydney Sandstone Ridgetop Woodland
- Eastern Suburbs Banksia Scrub
- Tasmanian dry sclerophyll forests
- Aegean and Western Turkey sclerophyllous and mixed forests
- California chaparral and woodlands
- California coastal sage and chaparral
- Chilean Matorral
- Mallee Woodlands and Shrublands
- Italian sclerophyllous and semi-deciduous forests
- Eastern Mediterranean conifer–sclerophyllous–broadleaf forests
- Southwest Iberian Mediterranean sclerophyllous and mixed forests
- Tyrrhenian–Adriatic sclerophyllous and mixed forests
- Canary Islands dry woodlands and forests
- Mediterranean acacia–argania dry woodlands
- Mediterranean dry woodlands and steppe
- Southeastern Iberian shrubs and woodlands
- Cyprus Mediterranean forests
- Crete Mediterranean forests
- Cape Floristic Region
- Southern Anatolian montane conifer and deciduous forests
- Albany thickets
- Northwest Iberian montane forests
See also
[edit]- Mediterranean forests, woodlands, and scrub
- Chaparral
- Fynbos
- Maquis shrubland
- Garrigue
- Kwongan
- Matorral
- Barren vegetation
References
[edit]- ^ Bowman, D. M. J. S., ed. (2000). Australian Rainforests: Islands of Green in a Land of Fire. Cambridge: Cambridge University Press. pp. 48–67. doi:10.1017/cbo9780511583490.004. ISBN 978-0-521-46568-7.
- ^ C. Michael Hogan. 2010. Leather Oak, Quercus durata. Encyclopedia of Earth. National Council for Science and Environment. Washington, DC
- ^ Grabherr, Georg, ed. (1997). Farbatlas Ökosysteme der Erde: natürliche, naturnahe und künstliche Land-Ökosysteme aus geobotanischer Sicht. Stuttgart: Ulmer. ISBN 978-3-8001-3489-2.
- ^ Pott, Richard (2005). Allgemeine Geobotanik: Biogeosysteme und Biodiversität. Springer-Lehrbuch. Berlin Heidelberg: Springer. pp. 353–398. ISBN 978-3-540-23058-8.
- ^ "California Chaparral & Woodlands". World Wildlife Fund. Archived from the original on October 8, 2012. Retrieved June 15, 2012. (material included verbatim under the CC BY-SA 3.0 license)
- ^ Duncan, F; Brown, M.J (1985). "Dry Sclerophyll vegetation in Tasmania". Wildlife Division Technical Report 85/1- National Parks and Wildlife Services.
- ^ "Sydney Coastal Dry Sclerophyll Forests". NSW Environment & Heritage. Retrieved September 17, 2012.
- ^ Hogan, C. Michael (October 17, 2012). "Chilean Wine Palm (Jubaea chilensis)". Archived from the original on October 17, 2012. Retrieved November 19, 2023.
- ^ Pfadenhauer, Jörg; Klötzli, Frank (2014). Vegetation der Erde: Grundlagen, Ökologie, Verbreitung. Berlin Heidelberg: Springer Spektrum. pp. 289–291. ISBN 978-3-642-41949-2.
- ^ Schultz, Jürgen (2000). Handbuch der Ökozonen. UTB für Wissenschaft Geographie, Ökologie, Agrar- und Forstwissenschaften. Stuttgart: Ulmer. pp. 35, 45–46. ISBN 978-3-8252-8200-4.
- ^ a b Pfadenhauer, Jörg S.; Klötzli, Frank A. (2020). Fundamentals towards Understanding Global Vegetation. Cham: Springer International Publishing. pp. 1–120. doi:10.1007/978-3-030-49860-3_1. ISBN 978-3-030-49859-7.
- ^ Pfadenhauer, Jörg S.; Klötzli, Frank A. (2020). Zonal Vegetation of the Tropical Zone with Year-Round Rain. Cham: Springer International Publishing. pp. 121–177. doi:10.1007/978-3-030-49860-3_2. ISBN 978-3-030-49859-7.
- ^ Susanne Heise: Vegetation zones: The zone of the evergreen hardwood plants, in TERRA-Online/Realschule on Klett.de, Klett, Leipzig 2003, version January 26, 2006, accessed on December 17, 2020.
- ^ Schmithüsen, Josef, ed. (1976). Atlas zur Biogeographie. Meyers großer physischer Weltatlas. Mannheim: Bibliograph. Inst. pp. 19, 43. ISBN 978-3-411-00303-7.
- ^ Klaus Müller-Hohenstein: The geo-ecological zones of the Earth. In: Geography and School. Issue 59, Bayreuth 1989.
- ^ Simmons, Marion H. (1982). Acacias of Australia (Reprint ed.). Melbourne, Vic: Nelson. ISBN 978-0-17-005779-0.
- ^ Florence, Ross G. (2007). Ecology and silviculture of eucalypt forests. Collingwood: CSIRO Publ. ISBN 978-0-643-05799-9.
- ^ Mott, J. J., and J. C. Tothill. 1984. Tropical and subtropical woodlands. in G. N. Harrington and A. D. Wilson, editors. Management of Australia's Rangelands. CSIRO Publishing, Melbourne
- ^ Parsons R. F.; Cameron D .G. (1974). "Maximum Plant Species Diversity in Terrestrial Communities". Biotropica. 6 (3). The Association for Tropical Biology and Conservation: 202. doi:10.2307/2989653. JSTOR 2989653.
- ^ White, Mary E.; Frazier, Jim, eds. (1988). The greening of Gondwana (Repr ed.). French's Forest: Reed Books. ISBN 978-0-7301-0154-3.
- ^ Flannery, Tim F. (2002). The future eaters: an ecological history of the Australasian lands and people (1 ed.). New York, NY: Grove Press. ISBN 978-0-8021-3943-6.
- ^ Burrows, W. H.; Carter, J. O.; Scanlan, J. C.; Anderson, E. R. (1990). "Management of Savannas for Livestock Production in North-East Australia: Contrasts Across the Tree-Grass Continuum". Journal of Biogeography. 17 (4/5): 503–512. doi:10.2307/2845383. ISSN 0305-0270.
- ^ Harrington, G. N., M. H. Friedel, K. C. Hodgkinson, and J. C. Noble. 1984. Vegetation ecology and management. in G. N. Harrington and A. D. Wilson, editors. Management of Australia's Rangelands. CSIRO Publishing, Melbourne.
- ^ R. Major (2003). "Sclerophyll forests". Australian Museum. Retrieved February 14, 2005.