List of Pinus species
Pinus, the pines, is a genus of approximately 110–120 extant tree and shrub species. The genus is currently split into two subgenera, subgenus Pinus (hard pines), and subgenus Strobus (soft pines). Each of the subgenera have been further divided into sections and subsections based in the past on morphology, ecology and biogeography, and more recently increasingly from chloroplast DNA sequencing[1] and whole plastid 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 constant Pinus lambertiana were placed in three different clades of the subsection Strobus, and similar problems with many other species with widespread nonmonophyly.[4]
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Several features are used to distinguish the subgenera, sections, and subsections of pines; the number of leaves (needles) per fascicle, whether the fascicle sheaths are deciduous or persistent, the number of fibrovascular bundles per needle (two in Pinus, one in Strobus), the position of the resin 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 the umbo (dorsal or terminal) and presence of a prickle on the scales of the seed cones.[5]
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),[6] the mid Cretaceous (around 125 million years ago),[7] to the late Cretaceous (around 100 million years ago).[8]
Subgenus Pinus
[edit]Subgenus Pinus includes the yellow and hard pines. Pines in this subgenus have one to five needles per fascicle and two fibrovascular bundles per needle, and the fascicle sheaths are persistent, except in P. leiophylla and 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 cones either open soon after they mature or are serotinous.[9]
Section Pinus
[edit]Section 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, except for P. resinosa in northeastern North America and P. tropicalis in western Cuba.[9]
Subsection Incertae sedis
- †P. driftwoodensis – Early Eocene, British Columbia, Canada[10]
Subsection Pinus
[edit]
All but two species (P. resinosa and P. tropicalis) in Subsection Pinus are native to Eurasia.[9] The cones have moderate thickness scales, and are characterised by a slighty off-centre prickle ('excentromucronate') on the umbo.[11]
- 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. 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
Subsection Incertae sedis
[edit]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.[14]
- P. heldreichii – Bosnian pine
Subsection Incertae sedis
[edit]These pines are placed in subsection Pinus by some genetic studies,[8] but in subsection Pinaster by others.[2] In morphology and ecology, they belong in the latter group.[15] The subsection Merkusia has also been proposed for them.[14]
- P. latteri – Tenasserim pine
- P. merkusii – Sumatran pine
- P. ustulata – Philippine pine (recently split from P. merkusii)[16]
Subsection Pinaster
[edit]
Subsection Pinaster contains species native to warm climates in the Mediterranean region at low altitudes, as well as 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.[5] It is named after P. 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
Section Trifoliae
[edit]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.[9]
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,[6] but not until the mid Oligocene (about 30–35 million years ago) in others.[7][8]
Subsection Attenuatae
[edit]
The three closed-cone (serotinous) and fire adapted species of California and Baja California form a small subsection;[17] closely related to, and often included within subsection Australes. Some studies suggest Pinus glabra may also belong here (despite its morphological dissimilarity)[2] though others include it in subsection Australes.[8]
- P. attenuata – knobcone pine
- P. muricata – Bishop pine
- P. radiata – Monterey pine
Subsection Australes
[edit]
This subsection is native to North and Central America and islands in the Caribbean.[5][18][19] It has 26 living species.[17]
- 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 of P. caribaea by POWO,[20] but considered distinct by Businský[14])
- P. jaliscana – Jalisco pine
- P. lawsonii – Lawson's pine
- P. leiophylla – Chihuahua pine
- P. lumholtzii – Lumholtz's pine
- P. luzmariae[21]
- 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 – extinct Pliocene species from Yukon Territory, Canada[22]
Subsection Contortae
[edit]The Contortae are native to North America and Mexico.[5] It contains four accepted species.[17]
- P. banksiana – jack pine
- P. clausa – sand pine
- P. contorta (infraspecific taxa traditionally cited at varietal rank,[23] but with marked divergence in morphology and ecology, better treated at subspecific rank[24])
- P. c. subsp. contorta – shore pine
- P. c. subsp. latifolia – lodgepole pine
- P. c. subsp. murrayana – tamarack pine
- P. virginiana – Virginia pine
Subsection Ponderosae
[edit]
This subsection is native to Central America, Mexico, the western United States, and southwestern Canada,[5][25] although its former range was possibly much wider as evidenced by upper Miocene fossils belonging to this subsection found in Japan [26] It contains at least 13 living species and may contain five more if the disputed species become widely accepted.[17]
- P. apulcensis (syn. P. oaxacana; currently treated as a variety of P. pseudostrobus by POWO[27])
- 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)[28]
- P. gordoniana – Gordon's pine
- P. hartwegii – Hartweg's pine
- P. martinezii – (not accepted)[29]
- P. maximinoi – thinleaf pine
- P. montezumae – Montezuma pine
- P. ponderosa – ponderosa pine
- P. p. subsp. benthamiana – Pacific ponderosa[30]
- P. p. subsp. brachyptera – Southwestern ponderosa (disputed, P. brachyptera in POWO and a subspeices in GRIN)[31]
- 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)[32]
- P. pseudostrobus – smooth-bark Mexican pine
- P. stormiae – Storm's pine
- P. yecorensis – Yecora pine
- †P. fujiii[26]
- †P. johndayensis – Oligocene
Subsection Sabinianae
[edit]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.[17]
- P. coulteri – Coulter pine
- P. jeffreyi – Jeffrey pine
- P. sabiniana – gray pine
- P. torreyana – Torrey pine
Subgenus Strobus
[edit]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 P. nelsonii, where they are persistent. Cone scales are thinner and more flexible than those of subgenus Pinus, except in some species like P. maximartinezii, and cones usually open soon after they mature.[9]
Section Nelsonia
[edit]
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 taxon to the rest of Section Parrya, while a late embryogenesis abundant (LEA)-like gene (IFG8612) resolved it as sister to Section Quinquefoliae;[4] another study resolved it as sister to Subsection Balfourianae.[8] 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.[8] These unique characters warrent recognition as a monotypic section.[14]
Subsection Nelsonianae
[edit]Subsection Nelsonianae is native to northeastern Mexico. It consists of the single species with persistent fascicle sheaths.
- P. nelsonii – Nelson's pinyon
Section Parrya
[edit]Section Parrya has one to five needles per fascicle. The seeds have 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 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.[9]
Subsection Balfourianae
[edit]Subsection Balfourianae (bristlecone and foxtail pines) is native to southwest United States.
- P. aristata – Rocky Mountains bristlecone pine
- P. balfouriana – foxtail pine
- †P. crossii[33][34] - (Chattian; Creede Formation, Colorado)
- P. longaeva – Great Basin bristlecone pine
Subsection Rzedowskiae
[edit]The 'big-cone' pinyons, with larger cones than subsection Cembroides.[17]
- P. maximartinezii – big-cone pinyon
- P. pinceana – weeping pinyon
- P. rzedowskii – Rzedowski's pinyon
Subsection Cembroides
[edit]
Subsection Cembroides (pinyons or piñons) is native to Mexico and the southwestern United 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
Section Quinquefoliae
[edit]Section Quinquefoliae (white pines), as its name (which means "five-leaved") suggests, has five needles per fascicle except for P. krempfii, which has two, and P. gerardiana and 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, P. chiapensis reaches Guatemala.[35][36]
Subsection Gerardianae
[edit]Subsection Gerardianae 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
Subsection Krempfianae
[edit]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.
- P. krempfii – Krempf's pine
- †P. leptokrempfii – Oligocene[37]
Subsection Strobus
[edit]
Subsection 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.[38] 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
Incertae sedis
[edit]
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 – Middle Eocene, Golden Valley Formation, North Dakota, US
- †Pinus tetrafolia – Early Eocene, Klondike Mountain Formation – Okanagan Highlands Floras
See also
[edit]- Hybridization in pines (list of pine hybrids)
References
[edit]- ^ Gernandt et al. 2005.
- ^ a b c Zeb et al. 2019.
- ^ Parks, Matthew; Cronn, Richard; Liston, Aaron (2012). "Separating the wheat from the chaff: mitigating the effects of noise in a plastome phylogenomic data set from Pinus L. (Pinaceae)". BMC Evolutionary Biology. 12 (1): 100. doi:10.1186/1471-2148-12-100. ISSN 1471-2148. PMC 3475122. PMID 22731878.
- ^ a b Syring, John; Farrell, Kathleen; Businský, Roman; Cronn, Richard; Liston, Aaron (2007-04-01). "Widespread Genealogical Nonmonophyly in Species of Pinus Subgenus Strobus" (PDF). Systematic Biology. 56 (2): 163–181. doi:10.1080/10635150701258787. ISSN 1076-836X. Retrieved 2025-11-01.
- ^ a b c d e f Gernandt et al. 2005, p. 35.
- ^ a b Stull, Gregory W.; Qu, Xiao-Jian; Parins-Fukuchi, Caroline; Yang, Ying-Ying; Yang, Jun-Bo; Yang, Zhi-Yun; Hu, Yi; Ma, Hong; Soltis, Pamela S.; Soltis, Douglas E.; Li, De-Zhu (19 July 2021). "Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms". Nature Plants. 7 (8): 1015–1025. Bibcode:2021NatPl...7.1015S. doi:10.1038/s41477-021-00964-4. ISSN 2055-0278. PMID 34282286. S2CID 236141481.
- ^ a b Saladin, Bianca; Leslie, Andrew B.; Wüest, Rafael O.; Litsios, Glenn; Conti, Elena; Salamin, Nicolas; Zimmermann, Niklaus E. (2017). "Fossils matter: improved estimates of divergence times in Pinus reveal older diversification". BMC Evolutionary Biology. 17 (1). doi:10.1186/s12862-017-0941-z. ISSN 1471-2148. PMC 5381128. PMID 28376717.
- ^ a b c d e f Jin, Wei-Tao; Gernandt, David S.; Wehenkel, Christian; Xia, Xiao-Mei; Wei, Xiao-Xin; Wang, Xiao-Quan (2021-05-18). "Phylogenomic and ecological analyses reveal the spatiotemporal evolution of global pines". Proceedings of the National Academy of Sciences. 118 (20). doi:10.1073/pnas.2022302118. ISSN 0027-8424. PMC 8157994. PMID 33941644.
- ^ a b c d e f Gernandt et al. 2005, p. 38.
- ^ Stockey 1983.
- ^ Klaus, Wilhelm (1980). "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". Plant Systematics and Evolution. 134 (3/4). Springer: 137–171. ISSN 0378-2697. JSTOR 23642506. Retrieved 2025-11-01.
- ^ "Pinus uncinata Ramond ex DC". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 1 November 2025.
- ^ Sokołowska, Joanna; Fuchs, Hanna; Celiński, Konrad (2021-06-29). "New Insight into Taxonomy of European Mountain Pines, Pinus mugo Complex, Based on Complete Chloroplast Genomes Sequencing". Plants. 10 (7): 1331. doi:10.3390/plants10071331. ISSN 2223-7747. PMC 8309040. PMID 34209970.
- ^ a b c d Businský, Roman (2008). The genus Pinus L., pines : contribution to knowledge (in Czech). ISBN 978-80-85116-60-1.
- ^ Frankis, Michael (1999). "Plate 367. Pinus brutia". Curtis's Botanical Magazine. 16 (3): 173–184. doi:10.1111/1467-8748.00216. ISSN 1355-4905. Retrieved 2025-11-01.
- ^ Businský, Roman (2014). "The Pinus merkusii agg. (Pinaceae): Literature Review, Taxa Delimitation and Typifications". Phyton. 54 (1): 1–26. doi:10.12905/0380.phython54(1)2014-0001.
- ^ a b c d e f Earle 2025.
- ^ "Pinus cubensis". Germplasm Resources Information Network. Agricultural Research Service, United States Department of Agriculture.
- ^ "Pinus occidentalis". Germplasm Resources Information Network. Agricultural Research Service, United States Department of Agriculture.
- ^ "Pinus hondurensis Sénécl". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 1 November 2025.
- ^ Wehenkel, C.; Mariscal-Lucero, S.; González-Elizondo, M.S.; Aguirre-Galindo, V.A.; Fladung, M. & López-Sánchez, C.A. (2020). "Tall Pinus luzmariae trees with genes from P. herrerae". PeerJ. 8 e8648. doi:10.7717/peerj.8648. PMC 7049253. PMID 32149029.
- ^ McKown, Stockey & Schweger 2002.
- ^ "Pinus contorta Douglas ex Loudon". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 22 October 2025.
- ^ "Pinus contorta (lodgepole pine) description". The Gymnosperm Database. 2025-02-10. Retrieved 2025-11-01.
- ^ "Pinus hartwegii". Germplasm Resources Information Network. Agricultural Research Service, United States Department of Agriculture.
- ^ a b Yamada, Toshihiro; Yamada, Mariko; Tsukagoshi, Minoru (2015-12-16). "Taxonomic revision of Pinus fujiii (Yasui) Miki (Pinaceae) and its implications for the phytogeography of the Section Trifoliae in east Asia". PLOS ONE. 10 (12) e0143512. Bibcode:2015PLoSO..1043512Y. doi:10.1371/journal.pone.0143512. ISSN 1932-6203. PMC 4689564. PMID 26673795.
- ^ "Pinus apulcensis Lindl". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 2 November 2025.
- ^ "Pinus estevezii (Martínez) J.P.Perry". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 22 October 2025.
- ^ "Pinus martinezii E.Larsen". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 22 October 2025.
- ^ "Pinus ponderosa". Germplasm Resources Information Network. Agricultural Research Service, United States Department of Agriculture.
- ^ "Pinus brachyptera Engelm". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 22 October 2025.
- ^ "Pinus scopulorum (Engelm.) Lemmon". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 22 October 2025.
- ^ Knowlton, F.H. (1923). Fossil plants from the Tertiary lake beds of South-Сentral Colorado (Report). Professional Paper. Vol. 131-G. United States Geological Survey. pp. 183–197. doi:10.3133/pp131G.
- ^ Wolfe, J.A.; Schorn, H.E. (1990). Taxonomic revision of the Spermatopsida of the Oligocene Creede flora, southern Colorado (Report). Bulletin. Vol. 1923. United States Geological Survey. pp. 1–40. doi:10.3133/b1923.
- ^ Gernandt et al. 2005, pp. 38–39.
- ^ "Pinus chiapensis". Germplasm Resources Information Network. Agricultural Research Service, United States Department of Agriculture.
- ^ Zhang, Jian-Wei; Wang, Li; D'Rozario, Ashalata; Liang, Xiao-Qing; Huang, Jian; Zhou, Zhe-Kun (2021). "Pinus leptokrempfii, an Oligocene Relative of the Flat-Needled Pine PINUS krempfii (Pinaceae) from China: Implications for Paleogeographic Origin". International Journal of Plant Sciences. 182 (5): 389–400. Bibcode:2021IJPlS.182..389Z. doi:10.1086/713957. S2CID 233888471.
- ^ "Eastern white pine (Pinus strobus)". Minnesota Department of Natural Resources. St. Paul, Minnesota.
Bibliography
[edit]- Earle, Christopher J. (17 October 2025). "Pinus (Pine) description". The Gymnosperm Database. Archived from the original on 19 August 2025. Retrieved 21 August 2025.
- Gernandt, D. S.; López, G. G.; García, S. O.; Liston, A. (2005). "Phylogeny and classification of Pinus". 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 of Pinus Subgenus Pinus Subsection Contortae 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 from the original (PDF) on 2008-02-21. Retrieved 2011-05-16.
- 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.
External links
[edit]
- Tree of Life Web – favors classification of Ducampopinus species in Strobus.
- NCBI Taxonomy server – files Ducampopinus species above as Strobus.