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Pinus, thepines, is a genus of approximately 111extant 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 based onchloroplast DNA sequencing^[1] and wholeplastid genomic analysis.^[2] Older classifications split the genus into three subgenera – subgenusPinus, subgenusStrobus, and subgenusDucampopinus (pinyon,bristlecone and lacebark pines)^[3] – based on cone, seed and leaf characteristics. DNA phylogeny has shown that species formerly in subgenusDucampopinus are members of subgenusStrobus, soDucampopinus is no longer used.^[1]

The species of subgenusDucampopinus were regarded as intermediate between the other two subgenera. In the modern classification, they are placed into subgenusStrobus, yet they did not fit entirely well in either so they were classified in a third subgenus. In 1888 theCalifornian botanistJohn Gill Lemmon placed them in subgenusPinus. In general, this classification emphasized cone, cone scale, seed, and leaf fascicle and sheath morphology, and species in each subsection were usually recognizable by their general appearance. Pines with one fibrovascular bundle per leaf, (the former subgeneraStrobus andDucampopinus) were known ashaploxylon pines, while pines with two fibrovascular bundles per leaf, (subgenusPinus) were calleddiploxylon pines. Diploxylon pines tend to have harder timber and a larger amount of resin than the haploxylon pines. The current division into two subgenera (Pinus andStrobus) is supported with rigorous genetic evidence.

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 (2 inPinus or 1 inStrobus), the position of theresin ducts in the needles (internal or external), the presence or shape of the seed wings (absent, rudimentary, articulate, and adnate), and the position of theumbo (dorsal or terminal) and presence of a prickle on the scales of theseed cones.^[4]

Both subgenera are thought to have a very ancient divergence from one another, having diverged during the lateJurassic.^[5]

SubgenusPinus includes the yellow and hard pines. Pines in this subgenus have one to five needles per fascicle and twofibrovascular bundles per needle, and the fascicle sheaths arepersistent, except inP. leiophylla andP. lumholtzii. Cone scales are thicker and more rigid than those of subgenusStrobus, andcones either open soon after they mature or areserotinous.^[6]

SectionPinus has two or three needles per fascicle. Cones of all species have thick scales, and all except those ofP. pinea open at maturity. Species in this section are native toEurope,Asia, and theMediterranean, except forP. resinosa in northeasternNorth America andP. tropicalis in westernCuba.^[6]

SubsectionIncertae sedis

• †P. driftwoodensis – EarlyEocene,British Columbia,Canada^[7]

All but two species (P. resinosa andP. tropicalis) inSubsectionPinus are native to Eurasia.^[6]

• P. densata – Sikang pine
• P. densiflora – Korean red pine
• P. henryi – Henry's pine
• P. hwangshanensis – Huangshan pine
• P. kesiya – Khasi pine
• P. latteri? – Tenasserim pine
• P. luchuensis – Luchu pine
• P. massoniana – Masson's pine
• P. merkusii – Sumatran pine
• P. mugo – mountain pine
• P. nigra – Austrian pine
• †P. prehwangshanensis
• 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. uncinata
• P. yunnanensis – Yunnan pine

SubsectionPinaster contains species native to the Mediterranean, as well asP. roxburghii from theHimalayas. The scales of its cones lack spines.^[4] It is named afterP. pinaster.

• P. brutia – Turkish pine
• P. canariensis – Canary Island pine
• P. halepensis – Aleppo pine
• P. heldreichii – Bosnian 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.^[6]

Phylogenetic analysis supports ancient divergences within this section, with subsectionsAustrales andPonderosae having diverged during the mid-Cretaceous.^[5]

SubsectionAustrales is native toNorth andCentral America and islands in theCaribbean.^[4][8][9]

The closed-cone (serotinous) species of California and Baja California,P. attenuata,P. muricata, andP. radiata, are sometimes placed in a separate subsection,Attenuatae.^[10]

• P. attenuata – knobcone pine
• P. caribaea – Caribbean pine
• P. cubensis – Cuban pine
• P. echinata – shortleaf pine
• P. elliottii – slash pine
• †P. foisyi – extinct
• P. glabra – spruce pine
• P. georginae
• P. greggii – Gregg's pine
• P. herrerae – Herrera's pine
• P. jaliscana – Jalisco pine
• P. lawsonii – Lawson's pine
• P. leiophylla – Chihuahua pine
• P. lumholtzii – Lumholtz's pine
• P. luzmariae^[11]
• †P. matthewsii –Pliocene,Yukon Territory, Canada^[12]
• P. muricata – bishop pine
• 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. radiata – Monterey pine
• P. rigida – pitch pine
• P. serotina – pond pine
• P. taeda – loblolly pine
• P. tecunumanii – Tecun Uman pine
• P. teocote – ocote pine
• P. vallartensis

SubsectionContortae is native to North America and Mexico.^[4]

• P. banksiana – jack pine
• P. clausa – sand pine
• P. contorta
  • P. c. bolanderi – Bolander pine
  • P. c. contorta – shore pine
  • P. c. latifolia – lodgepole pine
  • P. c. murrayana – tamarack pine
• P. virginiana – Virginia pine

SubsectionPonderosae is native toCentral America,Mexico, the westernUnited States, and southwesternCanada,^[4][13] although its former range was possibly much wider as evidenced by upper Miocene fossils belonging to this subsection found in Japan^[14]

• P. arizonica – Arizona pine
• P. cooperi – Cooper's pine
• P. coulteri – Coulter pine
• P. devoniana – Michoacan pine
• P. gordoniana
• P. durangensis – Durango pine
• P. engelmannii – Apache pine
• †P. fujiii^[14]
• P. hartwegii – Hartweg's pine
• P. jeffreyi – Jeffrey pine
• †P. johndayensis –Oligocene
• P. maximinoi – thinleaf pine
• P. montezumae – Montezuma pine
• P. ponderosa – ponderosa pine
• Pinus ponderosa var. willamettensis - Willamette Valley Ponderosa Pine
• P. pseudostrobus – smooth-bark Mexican 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.^[6]

Section Parrya has one to five needles per fascicle. The seeds either havearticulate (jointed) wings or no wings at all. In all species except forP. nelsonii, the fascicle sheaths curl back to form arosette before falling away. The cones have thick scales and release the seeds at maturity. This section is native to the southwestern United States and Mexico.^[6]

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

• P. aristata – Rocky Mountains bristlecone pine
• P. balfouriana – foxtail pine
• †P. crossii^[15][16] - (Chattian;Creede Formation, Colorado)
• P. longaeva – Great Basin bristlecone pine

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. maximartinezii – big-cone pinyon
• P. monophylla – single-leaf pinyon
• P. orizabensis – Orizaba pinyon
• P. pinceana – weeping pinyon
• P. quadrifolia – Parry pinyon
• P. remota – Texas pinyon or papershell pinyon
• P. rzedowskii – Rzedowski's pinyon

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

• P. nelsonii – Nelson's 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.^[17][18]

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^[19]

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.^[20] It is native to North and Central America, Europe, and Asia.^[4]

• 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. longlingensis –Late Pliocene,Mangbang Formation – Yunnan, China
• P. monticola – western white pine
• P. morrisonicola – Taiwan white pine
• P. parviflora – Japanese white pine
• P. hakkodensis – Hakkoda pine
• P. peuce – Macedonian pine
• P. pumila – Siberian dwarf pine
• P. ravii
• P. sibirica – Siberian pine
• P. strobus – eastern white pine
• P. strobiformis – Southwestern white pine (also Chihuahuan)
• Pinus stylesii
• P. wallichiana – blue pine
• P. wangii – Guangdong white pine

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)

• Gernandt, D. S.; López, G. G.; García, S. O.; Liston, A. (2005). "Phylogeny and classification ofPinus".Taxon.54 (1):29–42.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.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 of Pinus 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.

Wikimedia Commons has media related to:
Pinus (category)

• Tree of Life Web – favors classification ofDucampopinus species inStrobus.
• NCBI Taxonomy server – filesDucampopinus species above asStrobus.

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