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 sequencingTemplate:Sfn and whole plastid genomic analysis.Template:Sfn 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">Template:Cite journal</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 clades of the subsection Strobus, and similar problems with many other species with widespread nonmonophyly.<ref name="Syring07">Template:Cite journal</ref>
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.Template:Sfn
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">Template:Cite journal</ref> the mid Cretaceous (around 125 million years ago),<ref name="Saladin2017">Template:Cite journal</ref> to the late Cretaceous (around 100 million years ago).<ref name="Jin2021">Template:Cite journal</ref>
Subgenus Pinus Template:Anchor
Subgenus Pinus includes the yellow and hard pines. Pines in this subgenus often have two needles per fascicle but occasionally as many as eight.Template:Sfn They have 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.Template:Sfn
Section Pinus
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.Template:Sfn
Template:Anchor Subsection Incertae sedis
Subsection Pinus
All but two species (P. resinosa and P. tropicalis) in Subsection Pinus are native to Eurasia.Template:Sfn The cones have moderate thickness scales, and are characterised by a slighty off-centre prickle ('excentromucronate') on the umbo.<ref>Template:Cite journal</ref>
- 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 species P. uncinata by POWO,<ref>Template:Cite POWO</ref> but with minimal genetic distinction from P. mugo<ref>Template:Cite journal</ref>)
- 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
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">Template:Cite book</ref>
- 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.Template:Sfn In morphology and ecology, they belong in the latter group.<ref>Template:Cite journal</ref> The subsection Merkusia has also been proposed for them.<ref name="Businský2008"/>
- P. latteri – Tenasserim pine
- P. merkusii – Sumatran pine
- P. ustulata – Philippine pine (recently split from P. merkusii)<ref name="Businský2014">Template:Cite journal</ref>
Subsection Pinaster
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.Template:Sfn 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
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.Template:Sfn
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
The three closed-cone (serotinous) and fire adapted species of California and Baja California form a small subsection;Template:Sfn closely related to, and often included within subsection Australes. Some studies suggest Pinus glabra may also belong here (despite its morphological dissimilarity)Template:Sfn though others include it in subsection Australes.<ref name="Jin2021"/>
- P. attenuata – knobcone pine
- P. muricata – Bishop pine
- P. radiata – Monterey pine
Subsection Australes
This subsection is native to North and Central America and islands in the Caribbean.Template:Sfn<ref>Template:GRIN</ref><ref>Template:GRIN</ref> It has 26 living species.Template:Sfn
- 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,<ref>Template:Cite POWO</ref> but considered distinct by Businský<ref name="Businský2008"/>)
- P. jaliscana – Jalisco pine
- P. lawsonii – Lawson's pine
- P. leiophylla – Chihuahua pine
- P. lumholtzii – Lumholtz's pine
- P. luzmariae<ref name=WeheMariGonzAgui20>Template:Cite journal</ref>
- 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, CanadaTemplate:Sfn
Subsection Contortae
The Contortae are native to North America and Mexico.Template:Sfn It contains four accepted species.Template:Sfn
- P. banksiana – jack pine
- P. clausa – sand pine
- P. contorta (infraspecific taxa traditionally cited at varietal rank,<ref>Template:Cite POWO</ref> but with marked divergence in morphology and ecology, better treated at subspecific rank<ref name="Earle contorta">Template:Cite web</ref>)
- P. c. subsp. contorta – shore pine
- P. c. subsp. latifolia – lodgepole pine
- P. c. subsp. murrayana – tamarack pine
- P. virginiana – Virginia pine
Subsection Ponderosae
This subsection is native to Central America, Mexico, the western United States, and southwestern Canada,Template:Sfn<ref>Template:GRIN</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">Template:Cite journal</ref> It contains at least 13 living species and may contain five more if the disputed species become widely accepted.Template:Sfn
- P. apulcensis (syn. P. oaxacana; currently treated as a variety of P. pseudostrobus by POWO<ref>Template:Cite POWO</ref>)
- 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)<ref>Template:Cite POWO</ref>
- P. gordoniana – Gordon's pine
- P. hartwegii – Hartweg's pine
- P. martinezii – (not accepted)<ref>Template:Cite POWO</ref>
- P. maximinoi – thinleaf pine
- P. montezumae – Montezuma pine
- P. ponderosa – ponderosa pine
- P. p. subsp. benthamiana – Pacific ponderosa<ref name="GRIN P ponderosa">Template:GRIN</ref>
- P. p. subsp. brachyptera – Southwestern ponderosa (disputed, P. brachyptera in POWO and a subspeices in GRIN)<ref>Template:Cite POWO</ref>
- 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)<ref>Template:Cite POWO</ref>
- P. pseudostrobus – smooth-bark Mexican pine
- P. stormiae – Storm's pine
- P. yecorensis – Yecora pine
- †P. fujiii<ref name="Yamada2015"/>
- †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.Template:Sfn
- P. coulteri – Coulter pine
- P. jeffreyi – Jeffrey pine
- P. sabiniana – gray pine
- P. torreyana – Torrey pine
Subgenus Strobus Template:Anchor
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.Template:Sfn
Section Nelsonia
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;<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 warrent 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.
- P. nelsonii – Nelson's pinyon
Section Parrya
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.Template:Sfn
Subsection Balfourianae
Subsection Balfourianae (bristlecone and foxtail pines) is native to southwest United States.
- P. aristata – Rocky Mountains bristlecone pine
- P. balfouriana – foxtail pine
- †P. crossii<ref name="Knowlton1923">Template:Cite report</ref><ref name="WolfeSchorn1990">Template:Cite report</ref> - (Chattian; Creede Formation, Colorado)
- P. longaeva – Great Basin bristlecone pine
Subsection Rzedowskiae
The 'big-cone' pinyons, with larger cones than subsection Cembroides.Template:Sfn
- P. maximartinezii – big-cone pinyon
- P. pinceana – weeping pinyon
- P. rzedowskii – Rzedowski's pinyon
Subsection Cembroides
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
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.Template:Sfn<ref>Template:GRIN</ref>
Subsection Gerardianae
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
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<ref>Template:Cite journal</ref>
Subsection Strobus
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.<ref>Template:Cite web</ref> It is native to North and Central America, Europe, and Asia.Template:Sfn
- 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
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
- Hybridization in pines (list of pine hybrids)
References
Bibliography
- Template:Cite web
- Template:Cite journal
- Template:Cite journal
- Template:Cite journal
- Template:Cite journal
External links
- Tree of Life Web – favors classification of Ducampopinus species in Strobus.
- NCBI Taxonomy server – files Ducampopinus species above as Strobus.