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{{shortShort description|Species of plant}}
{{Speciesbox
| image = Pinus pinasterpinea and ruins on Palatine Hill.jpg
| image2 = Pinus pinaster02.jpg
| status = LC
| status_system = IUCN3.1
| status_ref = <ref name=IUCN"iucn status 19 November 2021">{{Citecite journaliucn | author = Farjon, A. |date=2013 |title = ''Pinus pinaster'' | journal = [[The IUCN Red List of Threatened Species]] | volume = 2013 | page = e.T42390A2977079 | publisher = [[IUCN]] | date = 2013 | url = http://www.iucnredlist.org/details/42390/0 | doi = 10.2305/IUCN.UK.2013-1.RLTS.T42390A2977079.en | access-date =19 9 JanuaryNovember 20182021}}</ref>
| display_parents = 3
| genus = Pinus
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}}
 
'''''Pinus pinaster''''', the '''maritime pine'''<ref name=GRIN>{{GRIN|name=Pinus pinaster Aiton|id=28525|accessdateaccess-date=4 November 2013}}</ref><ref>{{Cite web|url=http://www.wood-database.com/lumber-identification/softwoods/maritime-pine/|title=Maritime Pine &#124; the Wood Database - Lumber Identification (Softwood)}}</ref> or '''cluster pine''',<ref name=GRIN/> is a [[pine]] native to the south [[MediterraneanAtlantic Europe]] region and parts of the western [[Mediterranean]]. It is a hard, fast growing pine containingbearing small seeds with large wings.
 
==Distribution==
Its range is in the western [[Mediterranean Basin]], extending from [[Portugal]] and [[Northern Spain]] (especially in [[Galicia (Spain)|Galicia]]) to southern and [[Geography of France|Western France]], east to western Italy, [[Croatia]] and south to northern [[Tunisia]], [[Algeria]] and northern [[Morocco]].<ref name=Moran>{{cite book|first=V.C.|last=Moran|trans-title=X International Symposium on Biological Control of Weeds | title=Biological Control of Alien, Invasive Pine Trees (''Pinus'' species) in South Africa|year=2000|pages=941–953}}</ref> It favors a [[Mediterranean climate]], which is one that has cool, rainy winters and hot, dry summers.<ref name=Richardson>{{cite book|first=M|last=Richardson|trans-title=Journal of Vegetation Science | title=Assessing the risk of invasive success in ''Pinus'' and ''Banksia'' in South African mountain fynbos|year=1990|edition=1st|pages=629–642}}</ref>
 
It generally occurs at low to moderate altitudes, mostly from sea level to 600 m, but up to 2,000 m in the south of its range in Morocco. The high degree of fragmentation in the current natural distribution is caused by two factors: the discontinuity and altitude of the mountain ranges causing isolation of even close populations, and human activity.<ref name=alia/>
 
==History==
''Pinus pinaster'' is a popular topic in [[ecology]] because of its problematic growth and spread in [[South Africa]] for the past 150 years after being imported into the region at the end of the 17th century (1685–1693).<ref name=Moran /> It was found spreading in the [[Cape Peninsula]] by 1772.<ref name=Moran /> Towards the end of the 18th century (1780), ''P. pinaster'' was widely planted, and at the beginning of the 19th century (1825–1830), ''P. pinaster'' was planted commercially as a timber resource and for the [[forestry]] industry.<ref name=Moran /> The pine tree species invades large areas and more specifically [[fynbos]] [[vegetation]]. Fynbos vegetation is a fire-prone [[shrubland]] vegetation that is found in the southern and southwest cape of South Africa. It is found in greater abundance close to [[watercourse]]s.<ref name=Richardson /> [[Biological dispersal|Dispersal]], [[habitat]] loss, and [[fecundity]] are all factors that affect spread rate. The species favors [[Acidic soil|acidic soils]] with medium to high-density vegetation,<ref name=Richardson /> but it can also grow in basic soils and even in sandy and poor soils, where only few commercial species can grow.<ref name=alia>{{citation |author=Alía, R. |author2=Martín, S. |last-author-amp=yes|title=Maritime pine – ''Pinus pinaster'': Technical guidelines for genetic conservation and use|date=2003|page=6 pp |url=http://www.euforgen.org/fileadmin//templates/euforgen.org/upload/Publications/Technical_guidelines/857_Technical_guidelines_for_genetic_conservation_and_use_for_maritime_pine__Pinus_pinaster_.pdf |publisher=[[European Forest Genetic Resources Programme]]}}</ref>
 
==Description==
[[File:Pinus pinaster decoration cones.jpg|left|alt=|thumb|The [[Cone (botany)|cone]] of ''P. pinaster'']]
''Pinus pinaster'' is a medium-size [[tree]], reaching {{Convert|20–35 |m|abbr=off}} tall and with a [[Trunk (botany)|trunk]] diameter of up to {{Convert|1.2 |m|ft|abbr=on|frac=2}}, exceptionally {{Convert|1.8 |m|ft|abbr=on|frac=2}}.
[[File:Pinus pinaster Cones.jpg|thumb|Pinus pinaster Cones]]
The [[Bark (botany)|bark]] is orange-red, thick, and deeply fissured at the base of the trunk, somewhat thinner in the upper [[Crown (botany)|crown]].
 
The [[leaf|leaves]] ('needles') are in pairs, very stout ({{Convert|2&nbsp;|mm|abbr=off|disp=or|frac=16}} broad), up to {{Convert|25&nbsp;|cm|abbr=on|frac=2}} long,<ref name=Rushforth>{{cite book|first=Keith |last=Rushforth |trans-title=Pocket Guide to Trees | title=Bäume |language=Germande |location=[[Bern]] |publisher=Hallwag AG |origyearorig-year=1980 |year=1986 |edition=2nd |isbn=3-444-70130-6 |page=63}}</ref> and bluish-green to distinctly yellowish-green. The maritime pine features the longest and most robust needles of all European pine species.<ref name=Rushforth/>
The bark is orange-red, thick, and deeply fissured at the base of the trunk, somewhat thinner in the upper [[Crown (botany)|crown]].
 
The [[conifer cone|cones]] are conic, {{Convert|10–20&nbsp;|cm|abbr=on|frac=2}} long<ref name=Rushforth/> and {{Convert|4–6&nbsp;|cm|abbr=on|frac=2}} broad at the base when closed, green at first, ripening glossy red-brown when 24 months old. They open slowly over the next few years, or after being heated by a [[wildfire|forest fire]], to release the seeds, opening to {{Convert|8–12&nbsp;|cm|abbr=on|frac=2}} broad.
The [[leaf|leaves]] ('needles') are in pairs, very stout (2&nbsp;mm broad), up to 25&nbsp;cm long,<ref name=Rushforth>{{cite book|first=Keith |last=Rushforth |trans-title=Pocket Guide to Trees | title=Bäume |language=German |location=[[Bern]] |publisher=Hallwag AG |origyear=1980 |year=1986 |edition=2nd |isbn=3-444-70130-6 |page=63}}</ref> and bluish-green to distinctly yellowish-green. The maritime pine features the longest and most robust needles of all European pine species.<ref name=Rushforth/>
 
The seeds are {{Convert|8–10 |mm|abbr=on|frac=16}} long, with a {{Convert|20–25 |mm|abbr=on|frac=16}} wing, and are wind-[[Seed dispersal|dispersed]].
The [[conifer cone|cones]] are conic, 10–20&nbsp;cm long<ref name=Rushforth/> and 4–6&nbsp;cm broad at the base when closed, green at first, ripening glossy red-brown when 24 months old. They open slowly over the next few years, or after being heated by a [[wildfire|forest fire]], to release the seeds, opening to 8–12&nbsp;cm broad.
 
The seeds are 8–10 mm long, with a 20–25 mm wing, and are wind-dispersed.
 
=== Similar species ===
Maritime pine is closely related to [[Turkish pine]], [[Canary Island pine]], and [[Aleppo pine]], which all share many features with it. It is a relatively non-variable species, with constant [[Morphology (biology)|morphology]] over the entire range.
 
== Distribution and habitat ==
==Ecology==
Its range is in the western [[Mediterranean Basin]] and the southern Atlantic coast of Europe, extending from central [[Portugal]] and [[Northern Spain]] (especially in [[Galicia (Spain)|Galicia]]) to southern and [[Geography of France|Western France]], east to western Italy, [[Croatia]] and south to northern [[Tunisia]], [[Algeria]] and northern [[Morocco]].<ref name=Moran>{{cite booksfnp|first=V.C.|last=Moran|trans-title=X International Symposium on Biological Control of Weeds Hoffmann|Donnelly|van title=Biological Control of Alien, Invasive Pine Trees (''Pinus'' species) in South AfricaWilgen|year=2000|pages=941–953}}</ref> It favorsfavours a [[Mediterranean climate]], which is one that has cool, rainy winters and hot, dry summers.<ref name="Richardson">{{cite book|first=M |last=Richardson |trans-titlefirst=Journal of Vegetation ScienceM | title=Assessing the risk of invasive success in ''Pinus'' and ''Banksia'' in South African mountain fynbos |year=1990 |edition=1st |pages=629–642 |trans-title=Journal of Vegetation Science}}</ref>
[[Larva]]e of the moth ''[[Dioryctria sylvestrella]]'' feed on this pine. Their boring activity causes large quantities of [[resin]] to flow from the wounds which weakens the tree and allows fungi and other pathogens to gain entry.<ref name=Ciesla>{{cite book|author=Ciesla, William |title=Forest Entomology: A Global Perspective|url=https://books.google.com/books?id=PkUPKWoRt6YC&pg=PT546 |year=2011 |publisher=John Wiley & Sons|isbn=978-1-4443-9788-8 |page=546}}</ref>
 
It generally occurs at low to moderate altitudes, mostly from sea level to {{Convert|600|m|abbr=on}}, but up to {{Convert|2,000|m|abbr=on}} in the south of its range in Morocco. The high degree of fragmentation in the current natural distribution is caused by two factors: the discontinuity and altitude of the mountain ranges causing isolation of even close populations, and human activity.<ref name="alia">{{citation |author=Alía, R. |title=Maritime pine – ''Pinus pinaster'': Technical guidelines for genetic conservation and use |date=2003 |url=http://www.euforgen.org/fileadmin//templates/euforgen.org/upload/Publications/Technical_guidelines/857_Technical_guidelines_for_genetic_conservation_and_use_for_maritime_pine__Pinus_pinaster_.pdf |page=6 pp |archive-url=https://web.archive.org/web/20170119050303/http://www.euforgen.org/fileadmin//templates/euforgen.org/upload/Publications/Technical_guidelines/857_Technical_guidelines_for_genetic_conservation_and_use_for_maritime_pine__Pinus_pinaster_.pdf |publisher=[[European Forest Genetic Resources Programme]] |access-date=2017-01-18 |name-list-style=amp |archive-date=2017-01-19 |author2=Martín, S. |url-status=dead}}</ref>
==Uses==
[[File:Pinheiro bravo Parque Litoral.jpg|thumb|Maritime pine forest in the Coastal Park in [[Esposende]], Northern Portugal as seen from the foredunes.]]
''Pinus pinaster'' is widely planted for timber in its native area, being one of the most important trees in [[forestry]] in France, Spain and Portugal. [[Landes forest]] in southwest France is the largest man-made maritime pine forest in Europe. ''Pinus pinaster'' [[resin]] is also a useful source of [[turpentine]] and [[rosin]].<ref>{{cite web|title=Interview: ''Pinus pinaster'' resin industry in Portugal (portuguese)|url=http://www.agrotec.pt/noticias/resinagem-em-portugal-um-foco-de-esperanca/|publisher=AGROTEC|accessdate=12 August 2015}}</ref>
 
== Ecology ==
In addition to industrial uses, maritime pine is also a popular ornamental tree, often planted in parks and gardens in areas with warm [[temperateness|temperate]] climates. It has become [[naturalised]] in parts of southern England, Argentina, South Africa and Australia.<ref>{{cite web|title=''Pinus pinaster''|url=http://apps.rhs.org.uk/plantselector/plant?plantid=6298|publisher=Royal Horticultural Society|accessdate=23 July 2013}}</ref>
''Pinus pinaster'' is a popular topic in [[ecology]] because of its problematic growth and spread in [[South Africa]] for the past 150 years after being imported into the region at the end of the 17th century (1685–1693).<ref name={{sfnp|Moran|Hoffmann|Donnelly|van />Wilgen|2000}} It was found spreading in the [[Cape Peninsula]] by 1772.<ref name=Moran /> Towards the end of the 18th century (1780), ''P. &nbsp;pinaster'' was widely planted, and at the beginning of the 19th century (1825–1830), ''P. &nbsp;pinaster'' was planted commercially as a timber resource and for the [[forestry]] industry.<ref name={{sfnp|Moran|Hoffmann|Donnelly|van />Wilgen|2000}} The pine tree species invades large areas and more specifically [[fynbos]] [[vegetation]]. Fynbos vegetation is a fire-prone [[shrubland]] vegetation that is found in the southern and southwest cape of South Africa. It is found in greater abundance close to [[watercourse]]s.<ref name="Richardson" /> [[Biological dispersal|Dispersal]], [[habitat]] loss, and [[fecundity]] are all factors that affect spread rate. The species favors [[Acidicacidic soil|acidic soils]]s with medium to high-density vegetation,<ref name="Richardson" /> but it can also grow in basic soils and even in sandy and poor soils, where only few commercial species can grow.<ref name="alia>{{citation" |author=Alía, R. |author2=Martín, S. |last-author-amp=yes|title=Maritime pine – ''Pinus pinaster'': Technical guidelines for genetic conservation and use|date=2003|page=6 pp |url=http://www.euforgen.org/fileadmin//templates/euforgen.org/upload/Publications/Technical_guidelines/857_Technical_guidelines_for_genetic_conservation_and_use_for_maritime_pine__Pinus_pinaster_.pdf |publisher=[[European Forest Genetic Resources Programme]]}}</ref>
 
''Pinus pinaster'' is a diagnostic species of the vegetation class ''Pinetea halepensis''.<ref>{{Cite journal|last1=Bonari|first1=Gianmaria|last2=Fernández‐González|first2=Federico|last3=Çoban|first3=Süleyman|last4=Monteiro‐Henriques|first4=Tiago|last5=Bergmeier|first5=Erwin|last6=Didukh|first6=Yakiv P.|last7=Xystrakis|first7=Fotios|last8=Angiolini|first8=Claudia|last9=Chytrý|first9=Kryštof|last10=Acosta|first10=Alicia T.R.|last11=Agrillo|first11=Emiliano|date=January 2021|editor-last=Ewald|editor-first=Jörg|title=Classification of the Mediterranean lowland to submontane pine forest vegetation|url=https://onlinelibrary.wiley.com/doi/10.1111/avsc.12544|journal=Applied Vegetation Science|language=en|volume=24|issue=1|doi=10.1111/avsc.12544|hdl=10400.5/21923|s2cid=228839165|issn=1402-2001|hdl-access=free}}</ref>
It is also used as a source of [[Flavonoid|flavonoids]], [[catechins]], [[proanthocyanidin]]s, and [[phenolic acid]]s. A [[dietary supplement]] derived from extracts from ''Pinus pinaster'' bark called [[Pycnogenol]] is marketed with claims it can treat many conditions; however, according to a 2012 Cochrane review, the evidence is insufficient to support its use for the treatment of any chronic disorder.<ref>{{cite journal|last=Schoonees|first=A|author2=Visser, J |author3=Musekiwa, A |author4= Volmink, J |title=Pycnogenol (extract of French maritime pine bark) for the treatment of chronic disorders|journal=Cochrane Database of Systematic Reviews|year=2012|issue=7|pages=CD008294|doi=10.1002/14651858.CD008294.pub4|pmid=22513958}}</ref>
 
[[Larva]]e of the moth ''[[Dioryctria sylvestrella]]'' feed on this pine. Their boring activity causes large quantities of [[resin]] to flow from the wounds which weakens the tree and allows fungi and other pathogens to gain entry.<ref name=Ciesla>{{cite book|author=Ciesla, William |title=Forest Entomology: A Global Perspective|url=https://books.google.com/books?id=PkUPKWoRt6YC&pg=PT546 |year=2011 |publisher=John Wiley & Sons|isbn=978-1-4443-9788-8 |page=546}}</ref>
==Invasiveness==
 
=== Invasiveness ===
===Results of ''P. pinaster'' invasion===
''Pinus pinaster'' is a successful [[invasive species]] in South Africa. One of the results of its invasion in South Africa is a decrease in the [[biodiversity]] of the native environment.<ref name=Moran /> The increase of [[extinction]] rates of the native species is correlated with the introduction of these species to South Africa. Invasive species occupy [[habitats]] of native species often forcing them to extinction or endangerment. For example, invasive species have the potential to decrease the diversity of native plants by 50–86% in the [[Cape Peninsula]] of South Africa.<ref name=Higgins>{{cite book|first=S|last=Higgins|trans-title=Conservation Biology | title=Predicting the Landscape-Scale Distribution of Alien Plants and Their Threat to Plant Diversity|year=1999|pages=303–313}}</ref> As discussed above, ''P. pinaster'' is a tree species found in shrubland in South Africa. When compared to other [[Environment (biophysical)|environment]], shrublands have the largest decline of species richness when invaded by an invasive species (Z=–1.33, p<0.001).<ref name=Gaertner>{{cite book|first=M|last=Gaertner|trans-title=Progress in Physical Geography | title=Impacts of alien plant invasions on species richness in Mediterranean-type ecosystems: a meta-analysis|year=2009|edition=33rd|pages=319–338}}</ref> Compared to [[graminoids]]; trees, annual [[herbs]], and creepers have a larger effect on decline of [[species richness]] (Z=–3.78; p<0.001).<ref name=Gaertner /> Lastly, compared to other countries, South Africa had the largest species richness decline when faced with invasive species.<ref name=Gaertner /> South Africa is not home to many insects and diseases that limit the population of ''P. pinaster'' back in its native [[habitat]].<ref name=Moran /> Not only is there evidence that alien plant invasions decrease biodiversity, but there is also evidence that the location of ''P. pinaster'' increases its negative effect on the species richness.
 
==== Results of invasion ====
In addition, depending on the regions ''P. pinaster'' invades, ''P. pinaster'' has the potential to dramatically alter the quantity of water in the environment. If ''P. pinaster'' invades an area covered with grasses and shrubs, the water level of the streams in this area would lower significantly because ''P. pinaster'' are [[evergreen trees]] that take up considerably more water than grasses and shrubs all year around.<ref name=Carbon>{{cite book|first=B.A.|last=Carbon|trans-title=Journal of Hydrology | title=Deep drainage and water use of forests and pastures grown on deep sands in a Mediterranean environment|year=1982|pages=53–63}}</ref> They deplete [[Surface run off|run-off]] in [[Drainage basin|catchment areas]] and water flow in rivers. This depletes the resources available for other species in the environment. ''P. pinaster'' tends to grow rapidly in [[riparian zones]], which are areas with abundant water where trees and plants grow twice as fast and invade. ''P. pinaster'' takes advantage of the water available and consequently reduces the amount of water in the area available for other species.<ref name=Carbon /> The fynbos catchments on the [[Western Cape]] of South Africa are a habitat negatively affected by ''P. pinaster''. Twenty-three years after planting the pines, there was a 55% decrease in streamflow in this area.<ref name=Moran /> Similarly, in [[KwaZulu-Natal]] Drakensberg there was an 82% reduction in streamflow 20 years after introducing ''P. pinaster'' to the area. In the [[Mpumalanga Province]], 6 streams completely dried up 12 years after [[grasslands]] were replaced with pines.<ref name=Moran /> To reinforce that, there is a negative effect from the invasive species ''P. pinaster'', these areas of dense ''P. pinaster'' were thinned and the number of trees in the area decreased. As a result, the [[streamflow]] in the fynbos catchments of the Western Cape increased by 44%. The streamflow in the Mpumalanga Province increased by 120%.<ref name=Moran /> As a result of ''P. pinaster'' growth, there is often less understory vegetation for livestock grazing. Once again there was a positive effect when some of the pines were removed and agreeable range grasses were planted. The grazing conditions for the sheep of the area were greatly improved when the ''P. pinaster'' plantation was thinned to 300 trees per hectare. The invasion of ''P. pinaster'' leads to the decrease of [[understory]] vegetation and therefore a decrease in [[livestock]].<ref name=Papanastasis>{{cite book|first=V.|last=Papanastasis|trans-title=School of Forestry and Natural Environment | title=Effects of thinning, fertilisation and sheep grazing on the understory vegetarion of ''Pinus pinaster'' plantations|year=1995|pages=181–189}}</ref>
''Pinus pinaster'' is a successful [[invasive species]] in South Africa. One of the results of its invasion in South Africa is a decrease in the [[biodiversity]] of the native environment.<ref name={{sfnp|Moran|Hoffmann|Donnelly|van /> Wilgen|2000}} The increase of [[extinction]] rates of the native species is correlated with the introduction of these species to South Africa. Invasive species occupy [[habitats]] of native species often forcing them to extinction or endangerment. For example, invasive species have the potential to decrease the diversity of native plants by 50–86% in the [[Cape Peninsula]] of South Africa.<ref name="Higgins">{{cite book|first=S |last=Higgins |trans-titlefirst=Conservation BiologyS | title=Predicting the Landscape-Scale Distribution of Alien Plants and Their Threat to Plant Diversity |year=1999 |pages=303–313 |trans-title=Conservation Biology}}</ref> As discussed above, ''P. &nbsp;pinaster'' is a tree species found in shrubland in South Africa.; Whenwhen compared to other [[EnvironmentBiophysical (biophysical)environment|environmentenvironments]], shrublands have the largest decline of species richness when invaded by an invasive species (Z=–1.33, p<0.001).<ref name="Gaertner">{{cite book|first=M |last=Gaertner |trans-titlefirst=Progress in Physical GeographyM | title=Impacts of alien plant invasions on species richness in Mediterranean-type ecosystems: a meta-analysis |year=2009 |edition=33rd |pages=319–338 |trans-title=Progress in Physical Geography}}</ref> Compared to [[graminoids]]; trees, annual [[herbs]], and creepers have a larger effect on decline of [[species richness]] (Z=–3.78; p<0.001).<ref name="Gaertner" /> Lastly, compared to other countries, South Africa had the largest species richness decline when faced with invasive species.<ref name="Gaertner" /> South Africa is not home to many insects and diseases that limit the population of ''P. &nbsp;pinaster'' back in its native [[habitat]].<ref name={{sfnp|Moran|Hoffmann|Donnelly|van /> Wilgen|2000}} Not only is there evidence that alien plant invasions decrease biodiversity, but there is also evidence that the location of ''P. &nbsp;pinaster'' increases its negative effect on the species richness.
 
In addition, depending on the regions ''P. pinaster'' invades, ''P. &nbsp;pinaster'' has the potential to dramatically alter the quantity of water in the environment. If ''P. &nbsp;pinaster'' invades an area covered with grasses and shrubs, the water level of the streams in this area would lower significantly because ''P. pinaster'' are [[evergreen trees]] that take up considerably more water than grasses and shrubs all year around.<ref name="Carbon">{{cite bookjournal |last=Carbon |first=B.A. |last=Carbon|trans-titleyear=Journal of Hydrology1982 | title=Deep drainage and water use of forests and pastures grown on deep sands in a Mediterranean environment |yeartrans-title=1982Journal of Hydrology |journal=Journal of Hydrology |volume=55 |issue=1 |pages=53–63 |bibcode=1982JHyd...55...53C |doi=10.1016/0022-1694(82)90120-2}}</ref> They deplete [[Surface run off|run-off]] in [[Drainage basin|catchment areas]] and water flow in rivers. This depletes the resources available for other species in the environment. ''P. &nbsp;pinaster'' tends to grow rapidly in [[riparian zones]], which are areas with abundant water where trees and plants grow twice as fast and invade. ''P. &nbsp;pinaster'' takes advantage of the water available and consequently reduces the amount of water in the area available for other species.<ref name="Carbon" /> The fynbos catchments on the [[Western Cape]] of South Africa are a habitat negatively affected by ''P. &nbsp;pinaster''. Twenty-three years after planting the pines, there was a 55% decrease in streamflow in this area.<ref name={{sfnp|Moran|Hoffmann|Donnelly|van /> Wilgen|2000}} Similarly, in [[KwaZulu-Natal]] Drakensberg there was an 82% reduction in streamflow 20 years after introducing ''P. pinaster'' to the area. In the [[Mpumalanga Province]], 6 streams completely dried up 12 years after [[grasslands]] were replaced with pines.<ref name=Moran /> To reinforce that, there is a negative effect from the invasive species ''P. &nbsp;pinaster'', these areas of dense ''P. &nbsp;pinaster'' were thinned and the number of trees in the area decreased. As a result, the [[streamflow]] in the fynbos catchments of the Western Cape increased by 44%. The streamflow in the Mpumalanga Province increased by 120%.<ref name={{sfnp|Moran|Hoffmann|Donnelly|van />Wilgen|2000}} As a result of ''P. &nbsp;pinaster'' growth, there is often less understory vegetation for livestock grazing. Once again there was a positive effect when some of the pines were removed and agreeable range grasses were planted. The grazing conditions for the sheep of the area were greatly improved when the ''P. &nbsp;pinaster'' plantation was thinned to 300 trees per hectare. The invasion of ''P. &nbsp;pinaster'' leads to the decrease of [[understory]] vegetation and therefore a decrease in [[livestock]].<ref name="Papanastasis">{{cite book|first=V. |last=Papanastasis |trans-titlefirst=School of Forestry and Natural EnvironmentV. | title=Effects of thinning, fertilisation and sheep grazing on the understory vegetarion of ''Pinus pinaster'' plantations |year=1995 |pages=181–189 |trans-title=School of Forestry and Natural Environment}}</ref>
 
It is sporadically naturalizing in Oakland and San Leandro in northern California.
 
==== Ecological interactions ====
[[Image:Pinus pinaster JPG2.jpg|thumb|right|Bark of ''P. pinaster''.|upright=1.11]]
''Pinus pinaster'' is particularly successful in regions with fynbos vegetation because it is [[adapted]] to high-intensity fires, thus allowing it to outcompete other species that are not as well adapted to high-intensity fires. In areas of fire-prone shrubland, the cones of ''P. &nbsp;pinaster'' will release seeds when in a relatively high-temperature environment for [[germination]] as a recovery mechanism. This adaptation increases the competitive ability of ''P. &nbsp;pinaster'' amongst other species in the fire-prone shrubland.<ref name="Richardson" /> In a 3-year observational study done in [[Spain|Northwestern Spain]], ''P. &nbsp;pinaster'' showed a naturally high [[Regeneration (biology)|regeneration]] rate.<ref name="Calvo">{{cite book|first=L |last=Calvo |trans-titlefirst=Plant EcologyL | title=Post-fire natural regeneration of a ''Pinus pinaster'' forest in NW Spain |year=2008 |volume=197 |pages=81–90 |trans-title=Plant Ecology}}</ref> Observations showed a mean of 25.25 seedlings per square metre within the first year and then slowly decreased the next two years due to [[intraspecific competition]].<ref name="Calvo" /> So not only does ''P. &nbsp;pinaster'' compete with other species, they also compete within their own species as well. When the height of ''P. &nbsp;pinaster'' increased there was a negative correlation with the number of ''P. &nbsp;pinaster'' [[seedlings]], results showed a decrease in ''P. &nbsp;pinaster'' seedlings (r=–0.41, p<0.05).<ref name="Calvo" />
 
Several other characteristics contribute to their success in the regions they have invaded, including their ability to grow rapidly and to produce small seeds with large [[Conifer cone|wings]]. Their ability to grow quickly with short juvenile periods allows them to [[Competition (biology)|outcompete]] many native species while their small seeds aids in their dispersal. The small seeds with large wings are beneficial for wind dispersal, which is the key to reaching new areas in regions with fynbos vegetation.<ref name="Richardson" /> [[Vertebrate]] [[Seed dispersal|seed dispersers]] are not commonly found in mountain fynbos vegetation; therefore those species that require the aid of vertebrate dispersal would be at a disadvantage in such an environment. For this reason, the small seed, low seed wing loading, and high winds found in mountainous regions all combine to provide a favorable situation for the dispersal of ''P. &nbsp;pinaster'' seeds.<ref name="Richardson" /> Without this efficient dispersal strategy, ''P. &nbsp;pinaster'' would not have been able to reach and invade areas, such as South Africa, that are suitable for its growth. Its dispersal ability is one of the key factors that have allowed ''P. pinaster'' to become such a successful invasive species.<ref name="Richardson" />
 
In addition to being an efficient disperser, ''P. &nbsp;pinaster'' is known to produce [[Resins|oleoresins]], such as oily [[terpenes]] or [[fatty acids]], which can inhibit other species within the community from growing.<ref name="Santalla">{{cite book|first=L |last=Calvo |trans-titlefirst=Forest Ecology and ManagementL | title=Regeneration after wildfire in communities dominated by ''Pinus pinaster'', an obligate seeder, and in others dominated by ''Quercus pyrenaica'', typical resprouter |year=2003 |volume=184 |pages=209–223 |trans-title=Forest Ecology and Management}}</ref> These [[resins]] are produced as a defense mechanism against [[insect]] predators, such as the large [[pine weevil]]. According to an experiment done in [[Spain]], the [[resin canal]] density was twice as high in the ''P. &nbsp;pinaster'' seedlings attacked by the weevils compared to the unattacked seedlings. Since ''P. &nbsp;pinaster'' has the ability to regulate their production of defense mechanisms, it can protect itself from predatory in an energy-efficient manner. The resins make the ''P. &nbsp;pinaster'' less vulnerable to damage from insects, but they are only produced in high concentrations when ''P. &nbsp;pinaster'' is under attack. In other words, ''P. &nbsp;pinaster'' does not waste energy producing resins in safe conditions, so the conserved energy can be used for growth or reproduction. These characteristics enhance the ability of ''P. &nbsp;pinaster'' survive and flourish in the areas it invades.<ref name="Krebs">{{cite book|first=C |last=Krebs |trans-titlefirst=PearsonC | title=Ecology |year=2009 |trans-title=Pearson}}</ref> Both the traits of ''P. &nbsp;pinaster'' and the habitat in South Africa are conducive to the success of ''P. pinaster'' in this region of the world.
 
==== Options for biological control ====
[[Insects]] and [[mites]] that feed on the seeds and cones of ''P. &nbsp;pinaster'' can be effective [[biological control]] options. An insect or mite that acts as an ideal biological control should have a high [[Total fertility rate|reproductive rate]] and be [[Host (biology)|host-specific]], meaning that it preys specifically on ''P. &nbsp;pinaster''. The [[Biological life cycle|life cycle]] of the predator should also match that of its specific host. Two key characteristics the predator should also exhibit are [[Self-limiting (biology)|self-limitation]] and the ability to survive in the presence of a declining prey population.<ref name="Krebs" /> Seed feeding insects are an effective control because they have high reproductive rates and target the seeds without diminishing the positive effect of the plant on the environment. Controlling the spread of ''P. &nbsp;pinaster'' seeds in the region is the key to limiting the growth and spread of this species because ''P. pinaster'' has the ability to produce a large number of seeds that are capable of dispersing very efficiently.<ref name={{sfnp|Moran|Hoffmann|Donnelly|van />Wilgen|2000}} One possible option is Trisetacus, an [[Eriophyidae|eriophyid]] mite. The main advantage to using this mite to control the population of ''P. &nbsp;Pinaster'' is its specificity to ''P. pinaster''; it can effectively control the population of ''P. &nbsp;pinaster'' by destroying the growing conelets in ''P. &nbsp;pinaster'' while limiting its impact to only this species. Another possible option is [[Pissodes|''Pissodes validirostris'']], a cone-feeding weevil that lays eggs in developing cones. When the [[larvae]] hatch, they feed on the growing seed tissue, preventing ''P. &nbsp;pinaster'' seeds from forming and dispersing. Although the adults feed on the trees as well, they do not do any damage to the seeds and only feed on the shoots of the tree, so they do not appear to negatively impact the growth of the trees. Different forms of ''P. &nbsp;validirostris'' have diverged to become host-specific to different pine trees. The type of ''P. &nbsp;validirostris'' that originated from [[Portugal]] appears to have specialized to ''P. &nbsp;pinaster''; therefore, this insect may be used in the future to control the spread of ''P. &nbsp;pinaster'' in South Africa.<ref name="Hoffmann">{{cite book|first=J |last=Hoffmann |trans-titlefirst=African EntomologyJ | title=Prospects for the biological control of invasive ''Pinus'' species (''Pinaceae'') in South Africa |year=2011 |pages=393–401 |trans-title=African Entomology}}</ref> The uncertainties regarding the host-specificity of different types of ''P. &nbsp;validirostris'', however, require more research to be completed before the introduction of the weevils into South Africa. An introduction of a species that is not host-specific to ''P. &nbsp;pinaster'' can lead to detrimental effects on both the environment and [[Industry|commercial (economics)|industries]] that are dependent on certain tree species. Two other biological control possibilities include the [[pyralid]] moth species [[Dioryctria|''Dioryctria mendasella'']] and ''D. &nbsp;mitatella'', but these species attack the vegetative [[Tissue (biology)|tissue]] instead of just the seeds of ''P. &nbsp;pinaster'', harming the plant itself.<ref name={{sfnp|Moran|Hoffmann|Donnelly|van />Wilgen|2000}} As of now, the eriophyid mite and cone-feeding weevil seem to hold the most potential to controlling the spread of ''P. &nbsp;pinaster'' in the regions it has invaded because they destroy the reproductive structures of the target invasive species.
 
==Uses==
[[File:Pinheiro bravo Parque Litoral.jpg|thumb|Maritime pine forest in the Coastal Park in [[Esposende]], Northern Portugal as seen from the foredunes.]]
''Pinus pinaster'' is widely planted for timber in its native area, being one of the most important trees in [[forestry]] in France, Spain and Portugal. [[Landes forest]] in southwest France is the largest man-made maritime pine forest in Europe. It has also been cultivated in Australia as plantation tree, to provide softwood timber.<ref>{{cite news |title=Anglesea Plantation |url=https://trove.nla.gov.au/newspaper/article/232257607 |access-date=2020-10-08 |work=The Geelong Advertiser |date=1926-05-01}}</ref> ''Pinus P.&nbsp;pinaster'' [[resin]] is also a useful source of [[turpentine]] and [[rosin]].<ref>{{cite web|title=Interview: ''Pinus pinaster'' resin industry in Portugal (portuguese)|url=http://www.agrotec.pt/noticias/resinagem-em-portugal-um-foco-de-esperanca/|publisher=AGROTEC|accessdateaccess-date=12 August 2015}}</ref>
 
In addition to industrial uses, maritime pine is also a popular ornamental tree, often planted in parks and gardens in areas with warm [[temperateness|temperate]] climates. It has become [[naturalised]] in parts of southern England, Uruguay, Argentina, South Africa and Australia.<ref>{{cite web|title=''Pinus pinaster''|url=http://apps.rhs.org.uk/plantselector/plant?plantid=6298|archive-url=https://archive.today/20121224035202/http://apps.rhs.org.uk/plantselector/plant?plantid=6298|url-status=dead|archive-date=24 December 2012|publisher=Royal Horticultural Society|accessdateaccess-date=23 July 2013}}</ref>
 
It is also used as a source of [[Flavonoid|flavonoidsflavonoid]]s, [[catechins]], [[proanthocyanidin]]s, and [[phenolic acid]]s. A [[dietary supplement]] derived from extracts from ''Pinus P.&nbsp;pinaster'' bark called [[Pycnogenol]] is marketed with claims it can treat many conditions; however, according to a 2012 Cochrane review, the evidence is insufficient to support its use for the treatment of any chronic disorder.<ref>{{cite journal|last=Schoonees|first=A|author2=Visser, J |author3=Musekiwa, A |author4= Volmink, J |title=Pycnogenol (extract of French maritime pine bark) for the treatment of chronic disorders|journal=Cochrane Database of Systematic Reviews|year=2012|issue=7|pages=CD008294|doi=10.1002/14651858.CD008294.pub4|pmid=22513958}}</ref>
 
==Pests==
''[[Pestalotiopsis|Pestalotiopsis pini]]'' (a species of [[ascomycete]] fungi), was found as an emerging pathogen on ''[[Pinus pinea]]'' {{Au|L.}} (Stone pine) and also on ''Pinus pinaster'' in Portugal. Evidence of shoot blight and stem necrosis were found in 2020. The fungus was found on needles, shoots and trunks of the pines.<ref>{{cite journal |last1=Silva |first1=Ana Cristina |last2=Diogo |first2=Eugénio |last3=Henriques |first3=Joana |last4=Ramos |first4=Ana Paula |last5=Sandoval-Denis |first5=Marcelo |last6=Crous |first6=Pedro W. |last7=Bragança |first7=Helena |title=''Pestalotiopsis pini'' sp. nov., an Emerging Pathogen on Stone Pine (''Pinus pinea'' L.) |journal=Forests |date=2020 |volume=11 |issue=8 |page=805 |doi=10.3390/f11080805|doi-access=free |hdl=10400.5/20420 |hdl-access=free }}</ref>
 
==References==
{{Reflist|2}}
 
== Sources ==
* {{cite journal
| date = 2000
|last1=Moran |first1=V.C. | last2=Hoffmann | first2=J.H. |last3=Donnelly | first3= D. | last4=van Wilgen | first4=B.W. |last5=Zimmermann | first5=H.G.
|title=Biological Control of Alien, Invasive Pine Trees (''Pinus'' species) in South Africa
|pages=941–953
|journal=Proceedings of the X International Symposium on Biological Control of Weeds
| url = https://www.invasive.org/publications/xsymposium/proceed/13pg941.pdf
}}
 
==Further reading==
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{{Commons|Pinus pinaster}}
* [http://www.euforgen.org/species/pinus-pinaster/ ''Pinus pinaster''] – distribution map, genetic conservation units and related resources. [[European Forest Genetic Resources Programme]] (EUFORGEN)
* {{PFAF|Pinus pinaster}}
* {{CalPhotos|Pinus|pinaster}}
 
{{Taxonbar|from=Q271582}}
{{Authority control}}
 
[[Category:Flora of Algeria]]
[[Category:Flora of Malta]]
[[Category:Invasive plant species in South Africa]]
[[Category:Least concern plants]]
[[Category:Pinus|pinaster]]
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[[Category:Trees of Morocco]]
[[Category:Trees of Europe]]
[[Category:Flora of the Mediterranean Basin]]