{{Short description|None}} {{DISPLAYTITLE:List of ''Pinus'' species}} '''''[[Pine|Pinus]]''''', the '''pines''', is a genus of approximately 110–120 [[Neontology|extant]] tree and shrub species. The genus is currently split into two [[subgenus|subgenera]], subgenus '''''Pinus''''' ('''hard pines'''), and subgenus '''''Strobus''''' ('''soft pines'''). Each of the subgenera have been further divided into [[section (botany)|sections and subsections]] based in the past on morphology, ecology and biogeography, and more recently increasingly from [[chloroplast]] DNA sequencing{{sfn|Gernandt|López|García|Liston|2005}} and whole [[plastid]] genomic analysis.{{sfn|Zeb|Dong|Zhang|Wang|2019}} While the genetic analysis has given robust results at the higher levels, they often give conflicting results lower in the phylogenetic trees, with species allocated to different subsections (and sometimes different sections) by different studies or even within a study.<ref name="Parks2012">{{cite journal | last1=Parks | first1=Matthew | last2=Cronn | first2=Richard | last3=Liston | first3=Aaron | title=Separating the wheat from the chaff: mitigating the effects of noise in a plastome phylogenomic data set from ''Pinus'' L. (Pinaceae) | journal=BMC Evolutionary Biology | volume=12 | issue=1 | date=2012 | issn=1471-2148 | pmid=22731878 | pmc=3475122 | doi=10.1186/1471-2148-12-100 | doi-access=free | page=100}}</ref> Within subsections, the genetic relationships between species can be even more complex and conflicting; in one study, three samples of the very distinctive and morphologically constant ''[[Pinus lambertiana]]'' were placed in three different [[clade]]s of the subsection ''Strobus'', and similar problems with many other species with widespread nonmonophyly.<ref name="Syring07">{{cite journal | last1=Syring | first1=John | last2=Farrell | first2=Kathleen | last3=Businský | first3=Roman | last4=Cronn | first4=Richard | last5=Liston | first5=Aaron | title=Widespread Genealogical Nonmonophyly in Species of ''Pinus'' Subgenus ''Strobus'' | journal=Systematic Biology | volume=56 | issue=2 | date=2007-04-01 | issn=1076-836X | doi=10.1080/10635150701258787 | doi-access=free | pages=163–181 | url=https://academic.oup.com/sysbio/article-pdf/56/2/163/26562340/10635150701258787.pdf | access-date=2025-11-01}}</ref>
{{clade |label1=''Pinus'' |1={{clade |label1=Subgenus ''Pinus'' |1={{clade |label1=Section ''Trifoliae'' |1={{clade |1=Subsection ''Ponderosae'' |2={{clade |1=Subsection ''Contortae'' |2=Subsection ''Australes'' }} }} |label2=Section ''Pinus'' |2={{clade |1=Subsection ''Pinus'' |2=Subsection ''Pinaster'' }} }} |label2=Subgenus ''Strobus'' |2={{clade |label1=Section ''Quinquefoliae'' |1={{clade |1=Subsection ''Gerardianae'' |2={{clade |1=Subsection ''Krempfianae'' |2=Subsection ''Strobus'' }} }} |label2=Section ''Parrya'' |2={{clade |1=Subsection ''Nelsonianae'' |2={{clade |1=Subsection ''Balfourianae'' |2=Subsection ''Cembroides'' }} }} }} }} }}
Several features are used to distinguish the subgenera, sections, and subsections of pines; the number of leaves (needles) per [[fascicle (botany)|fascicle]], whether the [[fascicle sheath]]s are [[deciduous]] or [[Glossary of botanical terms#persistent|persistent]], the number of [[fibrovascular bundle]]s per needle (two in ''Pinus'', one in ''Strobus''), the position of the [[resin duct]]s in the needles (internal or external), the presence or shape of the seed wings (rudimentary or effective, articulate or adnate), and the position of the [[Glossary of botanical terms#umbo|umbo]] (dorsal or terminal) and presence of a prickle on the scales of the [[seed cone]]s.{{sfn|Gernandt|López|García|Liston|2005|p=35}}
The two subgenera are thought to have ancient divergence from each other, having diverged at some point between the late [[Jurassic]] (around 160 million years ago),<ref name="Stull-2021">{{Cite journal |last1=Stull |first1=Gregory W. |last2=Qu |first2=Xiao-Jian |last3=Parins-Fukuchi |first3=Caroline |last4=Yang |first4=Ying-Ying |last5=Yang |first5=Jun-Bo |last6=Yang |first6=Zhi-Yun |last7=Hu |first7=Yi |last8=Ma |first8=Hong |last9=Soltis |first9=Pamela S.|last10=Soltis|first10=Douglas E. |last11=Li |first11=De-Zhu |date=19 July 2021 |title=Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms |url=https://www.nature.com/articles/s41477-021-00964-4 |journal=Nature Plants |language=en |volume=7 |issue=8 |pages=1015–1025 |doi=10.1038/s41477-021-00964-4 |pmid=34282286 |bibcode=2021NatPl...7.1015S |s2cid=236141481 |issn=2055-0278|url-access=subscription }}</ref> the mid [[Cretaceous]] (around 125 million years ago),<ref name="Saladin2017">{{cite journal | last1=Saladin | first1=Bianca | last2=Leslie | first2=Andrew B. | last3=Wüest | first3=Rafael O. | last4=Litsios | first4=Glenn | last5=Conti | first5=Elena | last6=Salamin | first6=Nicolas | last7=Zimmermann | first7=Niklaus E. | title=Fossils matter: improved estimates of divergence times in Pinus reveal older diversification | journal=BMC Evolutionary Biology | volume=17 | issue=1 | date=2017 | issn=1471-2148 | pmid=28376717 | pmc=5381128 | doi=10.1186/s12862-017-0941-z | doi-access=free | page=}}</ref> to the late Cretaceous (around 100 million years ago).<ref name="Jin2021">{{cite journal | last1=Jin | first1=Wei-Tao | last2=Gernandt | first2=David S. | last3=Wehenkel | first3=Christian | last4=Xia | first4=Xiao-Mei | last5=Wei | first5=Xiao-Xin | last6=Wang | first6=Xiao-Quan | title=Phylogenomic and ecological analyses reveal the spatiotemporal evolution of global pines | journal=Proceedings of the National Academy of Sciences | volume=118 | issue=20 | date=2021-05-18 | issn=0027-8424 | pmid=33941644 | pmc=8157994 | doi=10.1073/pnas.2022302118 | doi-access=free | page=}}</ref>
==Subgenus ''Pinus'' {{anchor|Subgenus Pinus - yellow or hard pines}} == Subgenus ''[[Pinus subg. Pinus|Pinus]]'' includes the yellow and hard pines. Pines in this subgenus have two to five needles per fascicle (rarely as many as eight, in ''[[Pinus durangensis|P. durangensis]]'').{{sfn|Gernandt|López|García|Liston|2005|pp=35, 38}} They have two [[fibrovascular bundle]]s per needle, and the fascicle sheaths are [[Glossary of botanical terms#persistent|persistent]], except in ''[[Pinus leiophylla|P. leiophylla]]'' and ''[[Pinus lumholtzii|P. lumholtzii]]''. The cone scales are thicker and more rigid than those of subgenus ''Strobus'', and have a resin sealing band before opening (resulting in the cones opening with an audible crack); the [[pine cone|cones]] either open soon after they mature or are [[serotinous]].{{sfn|Gernandt|López|García|Liston|2005|p=38}}
===Section ''Pinus''=== '''Section ''[[Pinus sect. Pinus|Pinus]]''''' has two or three needles per fascicle. The cones have moderately thin to thick scales; in most they open at maturity, but are weakly serotinous in some species in subsection ''Pinaster''. Species in this section are native to [[Europe]], [[Asia]], and the [[Mediterranean region|Mediterranean]], except for ''P. resinosa'' in northeastern [[North America]] and ''P. tropicalis'' in western [[Cuba]].{{sfn|Gernandt|López|García|Liston|2005|p=38}}
{{anchor|Subsection Incertae sedis}} '''Subsection ''[[Incertae sedis]]''''' *†''[[Pinus driftwoodensis|P. driftwoodensis]]'' – Early [[Eocene]], [[British Columbia]], [[Canada]]{{sfn|Stockey|1983}}
====Subsection ''Pinus''==== [[File:Utsjoki, Lapland, Finland1.jpg|thumb|right|''Pinus sylvestris'']] [[File:2024-07-24 Pinus heldreichii, Park Prirode Blidinje, Jablanica, Herzegovina 1.jpg|thumb|right|''Pinus heldreichii'']] All but two species ('''''P. resinosa''''' and '''''P. tropicalis''''') in '''Subsection ''Pinus''''' are native to Eurasia.{{sfn|Gernandt|López|García|Liston|2005|p=38}} The cones have moderate thickness scales, and are characterised by a slightly off-centre prickle ('excentromucronate') on the umbo.<ref>{{cite journal | last=Klaus | first=Wilhelm | title=Neue Beobachtungen zur Morphologie des Zapfens von ''Pinus'' und ihre Bedeutung für die Systematik, Fossilbestimmung, Arealgestaltung und Evolution der Gattung / New Observations on the Morphology of ''Pinus''-Cones and Their Bearing on Taxonomy, Fossil Determination, Distribution, and Evolution of the Genus | journal=Plant Systematics and Evolution | publisher=Springer | volume=134 | issue=3/4 | year=1980 | issn=03782697 | jstor=23642506 | pages=137–171 | url=http://www.jstor.org/stable/23642506 | access-date=2025-11-01}}</ref> *''[[Pinus densata|P. densata]]'' – Sikang pine *''[[Pinus densiflora|P. densiflora]]'' – Japanese red pine or Korean red pine *''[[Pinus henryi|P. henryi]]'' – Henry's pine *''[[Pinus hwangshanensis|P. hwangshanensis]]'' – Huangshan pine *''[[Pinus kesiya|P. kesiya]]'' – Khasi pine *''[[Pinus luchuensis|P. luchuensis]]'' – Luchu pine *''[[Pinus massoniana|P. massoniana]]'' – Masson's pine *''[[Pinus mugo|P. mugo]]'' – mountain pine **''P. m.'' subsp. ''mugo'' **''P. m.'' subsp. ''uncinata'' (treated as a separate species ''P. uncinata'' by POWO,<ref>{{cite POWO |id=263396-1 |title=''Pinus uncinata'' Ramond ex DC. |access-date=1 November 2025}}</ref> but with minimal genetic distinction from ''P. mugo''<ref>{{cite journal | last1=Sokołowska | first1=Joanna | last2=Fuchs | first2=Hanna | last3=Celiński | first3=Konrad | title=New Insight into Taxonomy of European Mountain Pines, ''Pinus mugo'' Complex, Based on Complete Chloroplast Genomes Sequencing | journal=Plants | volume=10 | issue=7 | date=2021-06-29 | issn=2223-7747 | pmid=34209970 | pmc=8309040 | doi=10.3390/plants10071331 | doi-access=free | page=1331}}</ref>) *''[[Pinus nigra|P. nigra]]'' – Austrian pine *''[[Pinus resinosa|P. resinosa]]'' – red pine *''[[Pinus sylvestris|P. sylvestris]]'' – Scots pine *''[[Pinus tabuliformis|P. tabuliformis]]'' – Chinese red pine *''[[Pinus taiwanensis|P. taiwanensis]]'' – Taiwan red pine *''[[Pinus thunbergii|P. thunbergii]]'' – Japanese black pine *''[[Pinus tropicalis|P. tropicalis]]'' – tropical pine *''[[Pinus yunnanensis|P. yunnanensis]]'' – Yunnan pine *†''[[Pinus prehwangshanensis|P. prehwangshanensis]]'' *†[[Pinus yorkshirensis|''P. yorkshirensis'']]
====Subsection ''[[Incertae sedis]]''==== This pine is commonly resolved into subsection ''Pinaster'' by genetic studies, but this is in strong conflict with its morphology and ecology, which is much closer to subsection ''Pinus'', or alternatively placed in its own subsection ''Leucodermes'' close to subsection ''Pinus''.<ref name="Businský2008">{{cite book | last=Businský | first=Roman | title=The genus ''Pinus'' L., pines : contribution to knowledge | date=2008 | isbn=978-80-85116-60-1 | language=cs }}</ref> *''[[Pinus heldreichii|P. heldreichii]]'' – Bosnian pine
====Subsection ''[[Incertae sedis]]''==== These pines are placed in subsection ''Pinus'' by some genetic studies,<ref name="Jin2021"/> but in subsection ''Pinaster'' by others.{{sfn|Zeb|Dong|Zhang|Wang|2019}} In morphology and ecology, they belong in the latter group.<ref>{{cite journal | last=Frankis | first=Michael | title=Plate 367. Pinus brutia | journal=Curtis's Botanical Magazine | volume=16 | issue=3 | date=1999 | issn=1355-4905 | doi=10.1111/1467-8748.00216 | pages=173–184 | url=https://onlinelibrary.wiley.com/doi/10.1111/1467-8748.00216 | access-date=2025-11-01}}</ref> The subsection ''Merkusia'' has also been proposed for them.<ref name="Businský2008"/> *''[[Pinus latteri|P. latteri]]'' – Tenasserim pine *''[[Pinus merkusii|P. merkusii]]'' – Sumatran pine *''[[Pinus ustulata|P. ustulata]]'' – Philippine pine (recently split from ''P. merkusii'')<ref name="Businský2014">{{cite journal |last1=Businský |first1=Roman |title=The ''Pinus merkusii'' agg. (Pinaceae): Literature Review, Taxa Delimitation and Typifications |journal=Phyton |date=2014 |volume=54 |issue=1 |pages=1–26 |doi=10.12905/0380.phython54(1)2014-0001}}</ref>
==== Subsection ''Pinaster''==== [[File:Pinus roxburghii Dharamsala 1.jpg|thumb|right|''Pinus roxburghii'']] '''Subsection ''[[Pinus subsect. Pinaster|Pinaster]]''''' contains species native to warm climates in the Mediterranean region at low altitudes, as well as ''[[Pinus roxburghii|P. roxburghii]]'' from the [[Himalayas]]. The cones are thick-scaled and orange-brown to red-brown, and the cone scales are glossy and lack umbo spines.{{sfn|Gernandt|López|García|Liston|2005|p=35}} It is named after ''P. pinaster''. *''[[Pinus brutia|P. brutia]]'' – Turkish pine *''[[Pinus canariensis|P. canariensis]]'' – Canary Islands pine *''[[Pinus halepensis|P. halepensis]]'' – Aleppo pine *''[[Pinus pinaster|P. pinaster]]'' – maritime pine *''[[Pinus pinea|P. pinea]]'' – stone pine *''[[Pinus roxburghii|P. roxburghii]]'' – chir pine
===Section ''Trifoliae''=== '''Section ''Trifoliae''''' (American hard pines), despite its name (which means "three-leaved"), has two to five needles per fascicle, or rarely eight. The cones of most species open at maturity, but a few are [[serotinous]]. All but two American hard pines belong to this section.{{sfn|Gernandt|López|García|Liston|2005|p=38}}
The timing of divergences within this section is disputed, with subsections ''Australes'' and ''Ponderosae'' having diverged during the mid [[Cretaceous]] (about 110 million years ago) according to one study,<ref name="Stull-2021" /> but not until the mid [[Oligocene]] (about 30–35 million years ago) in others.<ref name="Saladin2017"/><ref name="Jin2021"/>
====Subsection ''Attenuatae''==== [[File:Pinus attenuata, San Luis Obispo, California 2.jpg|thumb|right|''Pinus attenuata'']] The three closed-cone (serotinous) and fire adapted species of California and Baja California form a small subsection;{{sfn|Earle 2025}} closely related to, and often included within subsection ''Australes''. Some studies suggest ''Pinus glabra'' may also belong here (despite its morphological dissimilarity){{sfn|Zeb|Dong|Zhang|Wang|2019}} though others include it in subsection ''Australes''.<ref name="Jin2021"/>
*''[[Pinus attenuata|P. attenuata]]'' – knobcone pine *''[[Pinus muricata|P. muricata]]'' – Bishop pine *''[[Pinus radiata|P. radiata]]'' – Monterey pine
====Subsection ''Australes''==== [[File:Pinus elliottii forest3.jpg|thumb|right|''Pinus elliottii'']] This subsection is native to [[North America|North]] and [[Central America]] and islands in the [[Caribbean]].{{sfn|Gernandt|López|García|Liston|2005|p=35}}<ref>{{GRIN |id=28439 |name=Pinus cubensis}}</ref><ref>{{GRIN |id=28511 |name=Pinus occidentalis}}</ref> It has 26 living species.{{sfn|Earle 2025}}
*''[[Pinus caribaea|P. caribaea]]'' – Caribbean pine *''[[Pinus cubensis|P. cubensis]]'' – Cuban pine *''[[Pinus echinata|P. echinata]]'' – shortleaf pine *''[[Pinus elliottii|P. elliottii]]'' – slash pine *''[[Pinus glabra|P. glabra]]'' – spruce pine *''[[Pinus georginae|P. georginae]]'' *''[[Pinus greggii|P. greggii]]'' – Gregg's pine *''[[Pinus herrerae|P. herrerae]]'' – Herrera's pine *''[[Pinus hondurensis|P. hondurensis]]'' – Honduras pine (treated as a variety of ''P. caribaea'' by POWO,<ref>{{cite POWO |id=314861-2 |title=''Pinus hondurensis'' Sénécl. |access-date=1 November 2025}}</ref> but considered distinct by Businský<ref name="Businský2008"/>) *''[[Pinus jaliscana|P. jaliscana]]'' – Jalisco pine *''[[Pinus lawsonii|P. lawsonii]]'' – Lawson's pine *''[[Pinus leiophylla|P. leiophylla]]'' – Chihuahua pine *''[[Pinus lumholtzii|P. lumholtzii]]'' – Lumholtz's pine *''[[Pinus luzmariae|P. luzmariae]]''<ref name=WeheMariGonzAgui20>{{Cite journal |last1=Wehenkel |first1=C. |last2=Mariscal-Lucero |first2=S. |last3=González-Elizondo |first3=M.S. |last4=Aguirre-Galindo |first4=V.A. |last5=Fladung |first5=M. |last6=López-Sánchez |first6=C.A. |date=2020 |title=Tall ''Pinus luzmariae'' trees with genes from ''P. herrerae'' |journal=PeerJ |volume=8 |article-number=e8648 |doi=10.7717/peerj.8648 |pmid=32149029 |pmc=7049253 |name-list-style=amp |doi-access=free }}</ref> *''[[Pinus occidentalis|P. occidentalis]]'' – Hispaniolan pine *''[[Pinus oocarpa|P. oocarpa]]'' – egg-cone pine *''[[Pinus palustris|P. palustris]]'' – longleaf pine *''[[Pinus patula|P. patula]]'' – patula pine *''[[Pinus praetermissa|P. praetermissa]]'' – McVaugh's pine *''[[Pinus pringlei|P. pringlei]]'' – Pringle's pine *''[[Pinus pungens|P. pungens]]'' – table mountain pine *''[[Pinus rigida|P. rigida]]'' – pitch pine *''[[Pinus serotina|P. serotina]]'' – pond pine *''[[Pinus taeda|P. taeda]]'' – loblolly pine *''[[Pinus tecunumanii|P. tecunumanii]]'' – Tecun Uman's pine *''[[Pinus teocote|P. teocote]]'' – ocote pine *''[[Pinus vallartensis|P. vallartensis]]'' *†''[[Pinus foisyi|P. foisyi]]'' – extinct
====Subsection ''Contortae''==== The ''Contortae'' are native to North America and Mexico.{{sfn|Gernandt|López|García|Liston|2005|p=35}} It contains four accepted species.{{sfn|Earle 2025}}
*''[[Pinus banksiana|P. banksiana]]'' – jack pine *''[[Pinus clausa|P. clausa]]'' – sand pine *''[[Pinus contorta|P. contorta]]'' (infraspecific taxa traditionally cited at varietal rank,<ref>{{cite POWO |id=30000492-2 |title=''Pinus contorta'' Douglas ex Loudon |access-date=22 October 2025}}</ref> but with marked divergence in morphology and ecology, better treated at subspecific rank<ref name="Earle contorta">{{cite web | title=Pinus contorta (lodgepole pine) description | website=The Gymnosperm Database | date=2025-02-10 | url=https://www.conifers.org/pi/Pinus_contorta.php | access-date=2025-11-01}}</ref>) **''P. c.'' subsp. ''contorta'' – shore pine **''P. c.'' subsp. ''latifolia'' – lodgepole pine **''P. c.'' subsp. ''murrayana'' – tamarack pine *''[[Pinus virginiana|P. virginiana]]'' – Virginia pine *†''[[Pinus matthewsii|P. matthewsii]]'' – extinct [[Pliocene]] species from [[Yukon Territory]], Canada{{sfn|McKown|Stockey|Schweger|2002}}
====Subsection ''Ponderosae''==== [[File:Pinus gordoniana trees, Cerro de San Juan, Nayarit.jpg|thumb|right|''Pinus gordoniana'']] [[File:Pinus jeffreyi cones BigBearLake.jpg|thumb|right|''Pinus jeffreyi'']] This subsection is native to [[Central America]], [[Mexico]], the western [[United States]], and southwestern [[Canada]],{{sfn|Gernandt|López|García|Liston|2005|p=35}}<ref>{{GRIN |id=409709 |name=Pinus hartwegii}}</ref> although its former range was possibly much wider as evidenced by upper Miocene fossils belonging to this subsection found in Japan <ref name="Yamada2015">{{Cite journal |last1=Yamada |first1=Toshihiro |last2=Yamada |first2=Mariko |last3=Tsukagoshi |first3=Minoru |date=2015-12-16 |title=Taxonomic revision of ''Pinus fujiii'' (Yasui) Miki (Pinaceae) and its implications for the phytogeography of the Section Trifoliae in east Asia |journal=PLOS ONE |language=en |volume=10 |issue=12 |article-number=e0143512 |doi=10.1371/journal.pone.0143512 |issn=1932-6203 |pmc=4689564 |pmid=26673795|bibcode=2015PLoSO..1043512Y |doi-access=free }}</ref> It contains at least 13 living species and may contain five more if the disputed species become widely accepted.{{sfn|Earle 2025}}
*''P. apulcensis'' (syn. ''P. oaxacana''; currently treated as a variety of ''P. pseudostrobus'' by POWO<ref>{{cite POWO |id=309646-2 |title=''Pinus apulcensis'' Lindl. |access-date=2 November 2025}}</ref>) *''[[Pinus arizonica|P. arizonica]]'' – Arizona pine *''[[Pinus cooperi|P. cooperi]]'' – Cooper's pine *''[[Pinus devoniana|P. devoniana]]'' – Michoacan pine *''[[Pinus durangensis|P. durangensis]]'' – Durango pine *''[[Pinus engelmannii|P. engelmannii]]'' – Apache pine * ''P. estevezii'' – (not currently accepted by POWO)<ref>{{cite POWO |id=1102546-2 |title=''Pinus estevezii'' (Martínez) J.P.Perry |access-date=22 October 2025}}</ref> *''[[Pinus gordoniana|P. gordoniana]]'' – Gordon's pine *''[[Pinus hartwegii|P. hartwegii]]'' – Hartweg's pine *''P. martinezii'' – (not accepted)<ref>{{cite POWO |id=197020-2 |title=''Pinus martinezii'' E.Larsen |access-date=22 October 2025}}</ref> *''[[Pinus maximinoi|P. maximinoi]]'' – thinleaf pine *''[[Pinus montezumae|P. montezumae]]'' – Montezuma pine *''[[Pinus ponderosa|P. ponderosa]]'' – ponderosa pine **''P. p.'' subsp. ''benthamiana'' – Pacific ponderosa<ref name="GRIN P ponderosa">{{GRIN |id=28530 |name=Pinus ponderosa}}</ref> **[[Pinus ponderosa subsp. brachyptera|''P. p.'' subsp. ''brachyptera'']] – Southwestern ponderosa (disputed, ''P. brachyptera'' in [[Plants of the World Online|POWO]] and a subspecies in [[Germplasm Resources Information Network|GRIN]])<ref>{{cite POWO |id=307170-2 |title=''Pinus brachyptera'' Engelm. |access-date=22 October 2025}}</ref> **''P. p.'' subsp. ''ponderosa'' – Columbia Basin ponderosa **''P. p.'' subsp. ''readiana'' – central high plains ponderosa **[[Pinus ponderosa subsp. scopulorum|''P. p.'' subsp. ''scopulorum'']] – Rocky Mountain ponderosa (disputed, ''P. scopulorum'' in POWO and a subspecies in GRIN)<ref>{{cite POWO |id=263310-1 |title=''Pinus scopulorum'' (Engelm.) Lemmon |access-date=22 October 2025}}</ref> *''[[Pinus pseudostrobus|P. pseudostrobus]]'' – smooth-bark Mexican pine *''[[Pinus stormiae|P. stormiae]]'' – Storm's pine *''[[Pinus yecorensis|P. yecorensis]]'' – Yecora pine *†''[[Pinus fujiii|P. fujiii]]''<ref name="Yamada2015"/> *†''[[Pinus johndayensis|P. johndayensis]]'' – [[Oligocene]]
====Subsection ''Sabinianae''==== These are pines of the western United States and Mexico with four existing species. Within the subsection the Coulter pine is closely related with the Jeffery pine and the gray pine is likewise paired with the Torrey pine.{{sfn|Earle 2025}}
*''[[Pinus coulteri|P. coulteri]]'' – Coulter pine *''[[Pinus jeffreyi|P. jeffreyi]]'' – Jeffrey pine *''[[Pinus sabineana|P. sabiniana]]'' – gray pine *''[[Pinus torreyana|P. torreyana]]'' – Torrey pine
==Subgenus ''Strobus'' {{anchor|Subgenus Strobus: white or soft pines}} == Subgenus ''Strobus'' includes the white and soft pines. Pines in this subgenus have one to five needles per fascicle and one [[fibrovascular bundle]] per needle, and the fascicle sheaths are [[deciduous]], except in ''[[Pinus nelsonii|P. nelsonii]]'', where they are persistent. Cone scales are thinner and more flexible than those of subgenus ''Pinus'', except in some species like ''[[Pinus maximartinezii|P. maximartinezii]]'', and [[pine cone|cones]] usually open soon after they mature.{{sfn|Gernandt|López|García|Liston|2005|p=38}}
===Section ''Nelsonia''=== [[File:Pinus nelsonii, General Zaragoza, Nuevo León, Mexico 1.jpg|thumb|right|''Pinus nelsonii'']] This unique pine is remarkable in differing from all other pines in multiple cone and foliage characters. Genetically, its position has varied between and within studies; three nuclear genes and chloroplast DNA placed it as [[Sister group|sister taxon]] to the rest of Section ''Parrya'', while a late embryogenesis abundant (LEA)-like gene (IFG8612) resolved it as sister to Section ''Quinquefoliae'';<ref name="Syring07"/> another study resolved it as sister to Subsection ''Balfourianae''.<ref name="Jin2021"/> It has probably evolved separately from the rest of the genus since the late [[Eocene]], with its [[most recent common ancestor]] around 37 million years ago.<ref name="Jin2021"/> These unique characters warrant recognition as a monotypic section.<ref name="Businský2008"/> ====Subsection ''Nelsonianae''==== '''Subsection ''Nelsonianae''''' is native to northeastern Mexico. It consists of the single species with persistent fascicle sheaths. *''[[Pinus nelsonii|P. nelsonii]]'' – Nelson's pinyon
===Section ''Parrya''=== '''Section Parrya''' has one to five needles per fascicle. The seeds have [[Glossary of botanical terms#articulate|articulate]] (jointed) wings; in most, the wing is vestigial, and remains attached to the cone scale when the seed is released apparently wingless. In all species, the fascicle sheaths curl back to form a [[rosette (botany)|rosette]] before falling away. The cones have thick scales (thin in subsection ''Balfourianae'') and release the seeds at maturity. This section is native to the southwestern United States and Mexico.{{sfn|Gernandt|López|García|Liston|2005|p=38}}
====Subsection ''Balfourianae''==== '''[[Pinus subsect. Balfourianae|Subsection Balfourianae]]''' (bristlecone and foxtail pines) is native to southwest [[United States]]. *''[[Pinus aristata|P. aristata]]'' – Rocky Mountains bristlecone pine *''[[Pinus balfouriana|P. balfouriana]]'' – foxtail pine *†''[[Pinus crossii|P. crossii]]''<ref name="Knowlton1923">{{cite report |last1=Knowlton |first1=F.H. |year=1923 |title=Fossil plants from the Tertiary lake beds of South-Central Colorado |series=Professional Paper |publisher=United States Geological Survey |volume=131-G |pages=183–197 |doi=10.3133/pp131G |doi-access=free }}</ref><ref name="WolfeSchorn1990">{{cite report |last1=Wolfe |first1=J.A. |last2=Schorn |first2=H.E. |year=1990 |title=Taxonomic revision of the Spermatopsida of the Oligocene Creede flora, southern Colorado |series=Bulletin |publisher=United States Geological Survey |volume=1923 |pages=1–40 |doi=10.3133/b1923 |doi-access=free }}</ref> - ([[Chattian]]; [[Creede Formation]], Colorado) *''[[Pinus longaeva|P. longaeva]]'' – Great Basin bristlecone pine
====Subsection ''Rzedowskiae''==== The 'big-cone' pinyons, with larger cones than subsection ''Cembroides''.{{sfn|Earle 2025}} *''[[Pinus maximartinezii|P. maximartinezii]]'' – big-cone pinyon *''[[Pinus pinceana|P. pinceana]]'' – weeping pinyon *''[[Pinus rzedowskii|P. rzedowskii]]'' – Rzedowski's pinyon
====Subsection ''Cembroides''==== [[File:Pinus cembroides Chisos 1.jpg|thumb|right|''Pinus cembroides'']] '''[[Pinyon pine|Subsection ''Cembroides'']]''' (pinyons or piñons) is native to [[Mexico]] and the southwestern [[United States]]. *''[[Pinus cembroides|P. cembroides]]'' – Mexican pinyon *''[[Pinus culminicola|P. culminicola]]'' – Potosi pinyon *''[[Pinus discolor|P. discolor]]'' – border pinyon *''[[Pinus edulis|P. edulis]]'' – Colorado pinyon *''[[Pinus johannis|P. johannis]]'' – Johann's pinyon *''[[Pinus monophylla|P. monophylla]]'' – single-leaf pinyon *''[[Pinus orizabensis|P. orizabensis]]'' – Orizaba pinyon *''[[Pinus quadrifolia|P. quadrifolia]]'' – Parry pinyon *''[[Pinus remota|P. remota]]'' – Texas pinyon or papershell pinyon
===Section ''Quinquefoliae''=== '''Section ''Quinquefoliae''''' (white pines), as its name (which means "five-leaved") suggests, has five needles per fascicle except for ''[[Pinus krempfii|P. krempfii]]'', which has two, and ''[[Pinus gerardiana|P. gerardiana]]'' and ''[[Pinus bungeana|P. bungeana]]'', which have three. All species have cones with thin or thick scales that open at maturity or do not open at all; none are serotinous. Species in this section are found in Eurasia and North America, and one species, ''[[Pinus chiapensis|P. chiapensis]]'' reaches Guatemala.{{sfn|Gernandt|López|García|Liston|2005|pp=38–39}}<ref>{{GRIN |id=457536 |name=Pinus chiapensis}}</ref>
====Subsection ''Gerardianae''==== '''Subsection ''[[Pinus subsection Gerardianae|Gerardianae]]''''' is native to East Asia. It has three or five needles per fascicle. *''[[Pinus bungeana|P. bungeana]]'' – lacebark pine *''[[Pinus gerardiana|P. gerardiana]]'' – chilgoza pine *''[[Pinus squamata|P. squamata]]'' – Qiaojia pine
====Subsection ''Krempfianae''==== '''Subsection ''Krempfianae''''' is currently native to Vietnam, with a fossil record extending into the Oligocene. It has two needles per fascicle, and they are atypically flattened. The cone scales are thick and have no prickles. Until 2021, the subsection was considered monotypic, when an Oligocene fossil species was described from Yunnan Province, China. *''[[Pinus krempfii|P. krempfii]]'' – [[Krempf's pine]] *†''[[Pinus leptokrempfii|P. leptokrempfii]]'' – Oligocene<ref>{{cite journal |url=https://www.journals.uchicago.edu/doi/abs/10.1086/713957 |last1=Zhang |first1=Jian-Wei |last2=Wang |first2=Li |last3=D'Rozario |first3=Ashalata |last4=Liang |first4=Xiao-Qing |last5=Huang |first5=Jian |last6=Zhou |first6=Zhe-Kun |year=2021 |title=Pinus leptokrempfii, an Oligocene Relative of the Flat-Needled Pine PINUS krempfii (Pinaceae) from China: Implications for Paleogeographic Origin |journal=International Journal of Plant Sciences |volume=182 |issue=5 |pages=389–400 |doi=10.1086/713957|bibcode=2021IJPlS.182..389Z |s2cid=233888471 |url-access=subscription }}</ref>
====Subsection ''Strobus''==== [[File:Pinus cembra cones in Gröden.JPG|thumb|right|''Pinus cembra'']] [[File:Pinus strobus Awenda.jpg|thumb|right|''Pinus strobus'']] '''Subsection ''[[Pinus subsect. Strobus|Strobus]]''''' has five needles per fascicle and thin cone scales with no prickles. Needles tend to be flexible and soft with slightly lighter side underneath.<ref>{{Cite web |title=Eastern white pine (Pinus strobus) |url=https://www.dnr.state.mn.us/trees/eastern-white-pine.html |website=Minnesota Department of Natural Resources |location=St. Paul, Minnesota}}</ref> It is native to North and Central America, Europe, and Asia.{{sfn|Gernandt|López|García|Liston|2005|p=35}} *''[[Pinus albicaulis|P. albicaulis]]'' – whitebark pine *''[[Pinus amamiana|P. amamiana]]'' – Yakushima white pine *''[[Pinus armandii|P. armandii]]'' – Chinese white pine *''[[Pinus arunachalensis|P. arunachalensis]]'' *''[[Pinus ayacahuite|P. ayacahuite]]'' – Mexican white pine *''[[Pinus bhutanica|P. bhutanica]]'' – Bhutan white pine *''[[Pinus cembra|P. cembra]]'' – Swiss pine *''[[Pinus chiapensis|P. chiapensis]]'' – Chiapas pine *''[[Pinus dabeshanensis|P. dabeshanensis]]'' – Dabieshan pine *''[[Pinus dalatensis|P. dalatensis]]'' – Vietnamese white pine *''[[Pinus fenzeliana|P. fenzeliana]]'' – Hainan white pine *''[[Pinus flexilis|P. flexilis]]'' – limber pine *''[[Pinus koraiensis|P. koraiensis]]'' – Korean pine *''[[Pinus lambertiana|P. lambertiana]]'' – sugar pine *''[[Pinus monticola|P. monticola]]'' – western white pine *''[[Pinus morrisonicola|P. morrisonicola]]'' – Taiwan white pine *''[[Pinus parviflora|P. parviflora]]'' – Japanese white pine *''[[Pinus peuce|P. peuce]]'' – Macedonian pine *''[[Pinus pumila|P. pumila]]'' – Siberian dwarf pine *''[[Pinus ravii|P. ravii]]'' *''[[Pinus reflexa|P. reflexa]]'' – Southwestern white pine *''[[Pinus sibirica|P. sibirica]]'' – Siberian pine *''[[Pinus strobus|P. strobus]]'' – eastern white pine *''[[Pinus strobiformis|P. strobiformis]]'' – Chihuahuan white pine *''[[Pinus stylesii|P. stylesii]]'' – Styles's white pine *''[[Pinus veitchii|P. veitchii]]'' – Veitch's pine *''[[Pinus wallichiana|P. wallichiana]]'' – blue pine *''[[Pinus wangii|P. wangii]]'' – Guangdong white pine *†''[[Pinus longlingensis|P. longlingensis]]'' – [[Late Pliocene]], [[Mangbang Formation]] – Yunnan, China
==''Incertae sedis''== [[File:Pinus latahensis needles SR 87-61-08 img2.jpg|thumb|right|upright|''[[Pinus latahensis]]'']] Species which are not placed in a subgenus at this time. *†''[[Pinus latahensis]]'' – Early Eocene, [[Klondike Mountain Formation]], [[Allenby Formation]] – [[Eocene Okanagan Highlands|Okanagan Highlands Floras]] *†''[[Pinus macrophylla (Berry)|Pinus macrophylla]]'' – Early Eocene, [[Klondike Mountain Formation]], [[Allenby Formation]] – [[Eocene Okanagan Highlands|Okanagan Highlands Floras]] *†''[[Pinus peregrinus]]'' – Middle [[Eocene]], [[Golden Valley Formation]], [[North Dakota]], US *†''[[Pinus tetrafolia]]'' – Early Eocene, [[Klondike Mountain Formation]] – [[Eocene Okanagan Highlands|Okanagan Highlands Floras]]
== See also ==
*[[Hybridization in pines]] (list of pine hybrids)
==References== {{reflist}}
===Bibliography=== * {{Cite web |last1=Earle |first1=Christopher J. |date=17 October 2025 |title=''Pinus'' (Pine) description |url=https://www.conifers.org/pi/Pinus.php |website=The Gymnosperm Database |url-status=live |archive-url=https://web.archive.org/web/20250819125808/https://www.conifers.org/pi/Pinus.php |archive-date=19 August 2025 |access-date=21 August 2025 |ref={{sfnref|Earle 2025}}}} *{{cite journal |last1=Gernandt |first1=D. S. |last2=López |first2=G. G. |last3=García |first3=S. O. |last4=Liston |first4=A. |year=2005 |title=Phylogeny and classification of ''Pinus'' |journal=Taxon |volume=54 |issue=1 |pages=29–42 |doi=10.2307/25065300 |jstor=25065300|bibcode=2005Taxon..54...29G }} *{{cite journal |last1=McKown |first1=A.D. |last2=Stockey |first2=R.A. |last3=Schweger |first3=C.E. |year=2002 |title=A New Species of ''Pinus'' Subgenus ''Pinus'' Subsection ''Contortae'' From Pliocene Sediments of Ch'Ijee's Bluff, Yukon Territory, Canada |journal=International Journal of Plant Sciences |volume=163 |issue=4 |pages=687–697 |url=http://www.mckown.ca/PDF/McKownetal2002.pdf |archive-url=https://wayback.archive-it.org/all/20080221182400/http://www.mckown.ca/PDF/McKownetal2002.pdf |archive-date=2008-02-21 |doi=10.1086/340425 |bibcode=2002IJPlS.163..687M |s2cid=86234947 |access-date=2011-05-16}} *{{Cite journal |last1=Stockey |first1=R.S. |year=1983 |title=''Pinus driftwoodensis'' sp.n. from the early Tertiary of British Columbia |journal=Botanical Gazette |volume=144 |issue=1 |pages=148–156 |jstor=2474678 |doi=10.1086/337355 |s2cid=84907438}} *{{Cite journal |last1=Zeb |first1=Umar |last2=Dong |first2=Wan-Lin |last3=Zhang |first3=Ting-Ting |last4=Wang |first4=Ruo-Nan |last5=Shahzad |first5=Khurram |last6=Ma |first6=Xiong-Feng |last7=Li |first7=Zhong-Hu |title=Comparative plastid genomics of ''Pinus'' species: Insights into sequence variations and phylogenetic relationships |journal=Journal of Systematics and Evolution |volume=58 |issue=2 |doi=10.1111/jse.12492 |issn=1759-6831 |date=25 March 2019 |pages=118–132 |s2cid=109171980 |publication-date=March 2020}}
==External links== {{commons and category|Pinus}} *[http://tolweb.org/Pinus Tree of Life Web] – favors classification of ''Ducampopinus'' species in ''Strobus''. *[https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?id=3337 NCBI Taxonomy server] – files ''Ducampopinus'' species above as ''Strobus''.
{{DEFAULTSORT:Pinus Classification}} [[Category:Pinus|*]] [[Category:Lists of plant species|Pinus]] [[Category:Lists of plants|Pinus]] [[Category:Lists of trees|Pinus]]