Pinus, thepines, is a genus of approximately 110–120extant tree and shrub species. The genus is currently split into twosubgenera, subgenusPinus (hard pines), and subgenusStrobus (soft pines). Each of the subgenera have been further divided intosections and subsections based in the past on morphology, ecology and biogeography, and more recently increasingly fromchloroplast DNA sequencing^[1] and wholeplastid genomic analysis.^[2] 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.^[3] 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 constantPinus lambertiana were placed in three differentclades of the subsectionStrobus, and similar problems with many other species with widespread nonmonophyly.^[4]

Several features are used to distinguish the subgenera, sections, and subsections of pines; the number of leaves (needles) perfascicle, whether thefascicle sheaths aredeciduous orpersistent, the number offibrovascular bundles per needle (two inPinus, one inStrobus), the position of theresin ducts in the needles (internal or external), the presence or shape of the seed wings (rudimentary or effective, articulate or adnate), and the position of theumbo (dorsal or terminal) and presence of a prickle on the scales of theseed cones.^[5]

The two subgenera are thought to have ancient divergence from each other, having diverged at some point between the lateJurassic (around 160 million years ago),^[6] the midCretaceous (around 125 million years ago),^[7] to the late Cretaceous (around 100 million years ago).^[8]

SubgenusPinus includes the yellow and hard pines. Pines in this subgenus have two to five needles per fascicle (rarely as many as eight, inP. durangensis).^[9] They have twofibrovascular bundles per needle, and the fascicle sheaths arepersistent, except inP. leiophylla andP. lumholtzii. The cone scales are thicker and more rigid than those of subgenusStrobus, and have a resin sealing band before opening (resulting in the cones opening with an audible crack); thecones either open soon after they mature or areserotinous.^[10]

SectionPinus 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 subsectionPinaster. Species in this section are native toEurope,Asia, and theMediterranean, except forP. resinosa in northeasternNorth America andP. tropicalis in westernCuba.^[10]

SubsectionIncertae sedis

• †P. driftwoodensis – EarlyEocene,British Columbia,Canada^[11]

All but two species (P. resinosa andP. tropicalis) inSubsectionPinus are native to Eurasia.^[10] The cones have moderate thickness scales, and are characterised by a slighty off-centre prickle ('excentromucronate') on the umbo.^[12]

• P. densata – Sikang pine
• P. densiflora – Korean red pine
• P. henryi – Henry's pine
• P. hwangshanensis – Huangshan pine
• P. kesiya – Khasi pine
• P. luchuensis – Luchu pine
• P. massoniana – Masson's pine
• P. mugo – mountain pine
  • P. m. subsp.mugo
  • P. m. subsp.uncinata (treated as a separate speciesP. uncinata by POWO,^[13] but with minimal genetic distinction fromP. mugo^[14])
• P. nigra – Austrian pine
• P. resinosa – red pine
• P. sylvestris – Scots pine
• P. tabuliformis – Chinese red pine
• P. taiwanensis – Taiwan red pine
• P. thunbergii – Japanese black pine
• P. tropicalis – tropical pine
• P. yunnanensis – Yunnan pine
• †P. prehwangshanensis

This pine is commonly resolved into subsectionPinaster by genetic studies, but this is in strong conflict with its morphology and ecology, which is much closer to subsectionPinus, or alternatively placed in its own subsectionLeucodermes close to subsectionPinus.^[15]

• P. heldreichii – Bosnian pine

These pines are placed in subsectionPinus by some genetic studies,^[8] but in subsectionPinaster by others.^[2] In morphology and ecology, they belong in the latter group.^[16] The subsectionMerkusia has also been proposed for them.^[15]

• P. latteri – Tenasserim pine
• P. merkusii – Sumatran pine
• P. ustulata – Philippine pine (recently split fromP. merkusii)^[17]

SubsectionPinaster contains species native to warm climates in the Mediterranean region at low altitudes, as well asP. roxburghii from theHimalayas. The cones are thick-scaled and orange-brown to red-brown, and the cone scales are glossy and lack umbo spines.^[5] It is named afterP. pinaster.

• P. brutia – Turkish pine
• P. canariensis – Canary Islands pine
• P. halepensis – Aleppo pine
• P. pinaster – maritime pine
• P. pinea – stone pine
• P. roxburghii – chir pine

SectionTrifoliae (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 areserotinous. All but two American hard pines belong to this section.^[10]

The timing of divergences within this section is disputed, with subsectionsAustrales andPonderosae having diverged during the midCretaceous (about 110 million years ago) according to one study,^[6] but not until the midOligocene (about 30–35 million years ago) in others.^[7][8]

The three closed-cone (serotinous) and fire adapted species of California and Baja California form a small subsection;^[18] closely related to, and often included within subsectionAustrales. Some studies suggestPinus glabra may also belong here (despite its morphological dissimilarity)^[2] though others include it in subsectionAustrales.^[8]

• P. attenuata – knobcone pine
• P. muricata – Bishop pine
• P. radiata – Monterey pine

This subsection is native toNorth andCentral America and islands in theCaribbean.^[5][19][20] It has 26 living species.^[18]

• P. caribaea – Caribbean pine
• P. cubensis – Cuban pine
• P. echinata – shortleaf pine
• P. elliottii – slash pine
• P. glabra – spruce pine
• P. georginae
• P. greggii – Gregg's pine
• P. herrerae – Herrera's pine
• P. hondurensis – Honduras pine (treated as a variety ofP. caribaea by POWO,^[21] but considered distinct by Businský^[15])
• P. jaliscana – Jalisco pine
• P. lawsonii – Lawson's pine
• P. leiophylla – Chihuahua pine
• P. lumholtzii – Lumholtz's pine
• P. luzmariae^[22]
• P. occidentalis – Hispaniolan pine
• P. oocarpa – egg-cone pine
• P. palustris – longleaf pine
• P. patula – patula pine
• P. praetermissa – McVaugh's pine
• P. pringlei – Pringle's pine
• P. pungens – table mountain pine
• P. rigida – pitch pine
• P. serotina – pond pine
• P. taeda – loblolly pine
• P. tecunumanii – Tecun Uman's pine
• P. teocote – ocote pine
• P. vallartensis
• †P. foisyi – extinct
• †P. matthewsii – extinctPliocene species fromYukon Territory, Canada^[23]

TheContortae are native to North America and Mexico.^[5] It contains four accepted species.^[18]

• P. banksiana – jack pine
• P. clausa – sand pine
• P. contorta (infraspecific taxa traditionally cited at varietal rank,^[24] but with marked divergence in morphology and ecology, better treated at subspecific rank^[25])
  • P. c. subsp.contorta – shore pine
  • P. c. subsp.latifolia – lodgepole pine
  • P. c. subsp.murrayana – tamarack pine
• P. virginiana – Virginia pine

This subsection is native toCentral America,Mexico, the westernUnited States, and southwesternCanada,^[5][26] although its former range was possibly much wider as evidenced by upper Miocene fossils belonging to this subsection found in Japan^[27] It contains at least 13 living species and may contain five more if the disputed species become widely accepted.^[18]

• P. apulcensis (syn.P. oaxacana; currently treated as a variety ofP. pseudostrobus by POWO^[28])
• P. arizonica – Arizona pine
• P. cooperi – Cooper's pine
• P. devoniana – Michoacan pine
• P. durangensis – Durango pine
• P. engelmannii – Apache pine
• P. estevezii – (not currently accepted by POWO)^[29]
• P. gordoniana – Gordon's pine
• P. hartwegii – Hartweg's pine
• P. martinezii – (not accepted)^[30]
• P. maximinoi – thinleaf pine
• P. montezumae – Montezuma pine
• P. ponderosa – ponderosa pine
  • P. p. subsp.benthamiana – Pacific ponderosa^[31]
  • P. p. subsp.brachyptera – Southwestern ponderosa (disputed,P. brachyptera inPOWO and a subspeices inGRIN)^[32]
  • P. p. subsp.ponderosa – Columbia Basin ponderosa
  • P. p. subsp.readiana – central high plains ponderosa
  • P. p. subsp.scopulorum – Rocky Mountain ponderosa (disputed,P. scopulorum in POWO and a subspecies in GRIN)^[33]
• P. pseudostrobus – smooth-bark Mexican pine
• P. stormiae – Storm's pine
• P. yecorensis – Yecora pine
• †P. fujiii^[27]
• †P. johndayensis –Oligocene

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.^[18]

• P. coulteri – Coulter pine
• P. jeffreyi – Jeffrey pine
• P. sabiniana – gray pine
• P. torreyana – Torrey pine

SubgenusStrobus includes the white and soft pines. Pines in this subgenus have one to five needles per fascicle and onefibrovascular bundle per needle, and the fascicle sheaths aredeciduous, except inP. nelsonii, where they are persistent. Cone scales are thinner and more flexible than those of subgenusPinus, except in some species likeP. maximartinezii, andcones usually open soon after they mature.^[10]

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 assister taxon to the rest of SectionParrya, while a late embryogenesis abundant (LEA)-like gene (IFG8612) resolved it as sister to SectionQuinquefoliae;^[4] another study resolved it as sister to SubsectionBalfourianae.^[8] It has probably evolved separately from the rest of the genus since the lateEocene, with itsmost recent common ancestor around 37 million years ago.^[8] These unique characters warrent recognition as a monotypic section.^[15]

SubsectionNelsonianae is native to northeastern Mexico. It consists of the single species with persistent fascicle sheaths.

• P. nelsonii – Nelson's pinyon

Section Parrya has one to five needles per fascicle. The seeds havearticulate (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 arosette before falling away. The cones have thick scales (thin in subsectionBalfourianae) and release the seeds at maturity. This section is native to the southwestern United States and Mexico.^[10]

Subsection Balfourianae (bristlecone and foxtail pines) is native to southwestUnited States.

• P. aristata – Rocky Mountains bristlecone pine
• P. balfouriana – foxtail pine
• †P. crossii^[34][35] - (Chattian;Creede Formation, Colorado)
• P. longaeva – Great Basin bristlecone pine

The 'big-cone' pinyons, with larger cones than subsectionCembroides.^[18]

• P. maximartinezii – big-cone pinyon
• P. pinceana – weeping pinyon
• P. rzedowskii – Rzedowski's pinyon

SubsectionCembroides (pinyons or piñons) is native toMexico and the southwesternUnited States.

• P. cembroides – Mexican pinyon
• P. culminicola – Potosi pinyon
• P. discolor – border pinyon
• P. edulis – Colorado pinyon
• P. johannis – Johann's pinyon
• P. monophylla – single-leaf pinyon
• P. orizabensis – Orizaba pinyon
• P. quadrifolia – Parry pinyon
• P. remota – Texas pinyon or papershell pinyon

SectionQuinquefoliae (white pines), as its name (which means "five-leaved") suggests, has five needles per fascicle except forP. krempfii, which has two, andP. gerardiana andP. 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,P. chiapensis reaches Guatemala.^[36][37]

SubsectionGerardianae is native to East Asia. It has three or five needles per fascicle.

• P. bungeana – lacebark pine
• P. gerardiana – chilgoza pine
• P. squamata – Qiaojia pine

SubsectionKrempfianae 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.

• P. krempfii –Krempf's pine
• †P. leptokrempfii – Oligocene^[38]

SubsectionStrobus has five needles per fascicle and thin cone scales with no prickles. Needles tend to be flexible and soft with slightly lighter side underneath.^[39] It is native to North and Central America, Europe, and Asia.^[5]

• P. albicaulis – whitebark pine
• P. amamiana – Yakushima white pine
• P. armandii – Chinese white pine
• P. arunachalensis
• P. ayacahuite – Mexican white pine
• P. bhutanica – Bhutan white pine
• P. cembra – Swiss pine
• P. chiapensis – Chiapas pine
• P. dabeshanensis – Dabieshan pine
• P. dalatensis – Vietnamese white pine
• P. fenzeliana – Hainan white pine
• P. flexilis – limber pine
• P. koraiensis – Korean pine
• P. lambertiana – sugar pine
• P. monticola – western white pine
• P. morrisonicola – Taiwan white pine
• P. parviflora – Japanese white pine
• P. peuce – Macedonian pine
• P. pumila – Siberian dwarf pine
• P. ravii
• P. reflexa – Southwestern white pine
• P. sibirica – Siberian pine
• P. strobus – eastern white pine
• P. strobiformis – Chihuahuan white pine
• P. stylesii – Styles's white pine
• P. veitchii – Veitch's pine
• P. wallichiana – blue pine
• P. wangii – Guangdong white pine
• †P. longlingensis –Late Pliocene,Mangbang Formation – Yunnan, China

Species which are not placed in a subgenus at this time.

• †Pinus latahensis – Early Eocene,Klondike Mountain Formation,Allenby Formation –Okanagan Highlands Floras
• †Pinus macrophylla – Early Eocene,Klondike Mountain Formation,Allenby Formation –Okanagan Highlands Floras
• †Pinus peregrinus – MiddleEocene,Golden Valley Formation,North Dakota, US
• †Pinus tetrafolia – Early Eocene,Klondike Mountain Formation –Okanagan Highlands Floras

• Hybridization in pines (list of pine hybrids)

• Earle, Christopher J. (17 October 2025)."Pinus (Pine) description".The Gymnosperm Database.Archived from the original on 19 August 2025. Retrieved21 August 2025.
• Gernandt, D. S.; López, G. G.; García, S. O.; Liston, A. (2005). "Phylogeny and classification ofPinus".Taxon.54 (1):29–42.Bibcode:2005Taxon..54...29G.doi:10.2307/25065300.JSTOR 25065300.
• McKown, A.D.; Stockey, R.A.; Schweger, C.E. (2002)."A New Species ofPinus SubgenusPinus SubsectionContortae From Pliocene Sediments of Ch'Ijee's Bluff, Yukon Territory, Canada"(PDF).International Journal of Plant Sciences.163 (4):687–697.Bibcode:2002IJPlS.163..687M.doi:10.1086/340425.S2CID 86234947. Archived fromthe original(PDF) on 2008-02-21. Retrieved2011-05-16.
• Stockey, R.S. (1983). "Pinus driftwoodensis sp.n. from the early Tertiary of British Columbia".Botanical Gazette.144 (1):148–156.doi:10.1086/337355.JSTOR 2474678.S2CID 84907438.
• Zeb, Umar; Dong, Wan-Lin; Zhang, Ting-Ting; Wang, Ruo-Nan; Shahzad, Khurram; Ma, Xiong-Feng; Li, Zhong-Hu (25 March 2019). "Comparative plastid genomics ofPinus species: Insights into sequence variations and phylogenetic relationships".Journal of Systematics and Evolution.58 (2) (published March 2020):118–132.doi:10.1111/jse.12492.ISSN 1759-6831.S2CID 109171980.

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Pinus (category)

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