List of Pinus species

Pinus, the pines, is a genus of approximately 111 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 based on chloroplast DNA sequencing[1] and whole plastid genomic analysis.[2] Older classifications split the genus into three subgenera – subgenus Pinus, subgenus Strobus, and subgenus Ducampopinus (pinyon, bristlecone and lacebark pines)[3] – based on cone, seed and leaf characteristics. DNA phylogeny has shown that species formerly in subgenus Ducampopinus are members of subgenus Strobus, so Ducampopinus is no longer used.[1]

Pinus	Subgenus Pinus Section Trifoliae   Subsection Ponderosae       Subsection Contortae     Subsection Australes       Section Pinus   Subsection Pinus     Subsection Pinaster       Subgenus Strobus Section Quinquefoliae   Subsection Gerardianae       Subsection Krempfianae     Subsection Strobus       Section Parrya   Subsection Nelsonianae       Subsection Balfourianae     Subsection Cembroides

The species of subgenus Ducampopinus were regarded as intermediate between the other two subgenera. In the modern classification, they are placed into subgenus Strobus, yet they did not fit entirely well in either so they were classified in a third subgenus. In 1888 the Californian botanist John Gill Lemmon placed them in subgenus Pinus. 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 subgenera Strobus and Ducampopinus) were known as haploxylon pines, while pines with two fibrovascular bundles per leaf, (subgenus Pinus) were called diploxylon 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 and Strobus) is supported with rigorous genetic evidence.

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 (2 in Pinus or 1 in Strobus), the position of the resin ducts in the needles (internal or external), the presence or shape of the seed wings (absent, rudimentary, articulate, and adnate), and the position of the umbo (dorsal or terminal) and presence of a prickle on the scales of the seed cones.[4]

Subgenus Pinus

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. Cone scales are thicker and more rigid than those of subgenus Strobus, and cones either open soon after they mature or are serotinous.[5]

Section Pinus

Section Pinus has two or three needles per fascicle. Cones of all species have thick scales, and all except those of P. pinea open at maturity. 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.[5]

Subsection Incertae sedis

• †P. driftwoodensis – Early Eocene, British Columbia, Canada[6]

Subsection Pinus

All but two species in Subsection Pinus are native to Eurasia.[5]

Image	Leaves & Cone	Scientific name	Common Name	Distribution
		P. densata	Sikang pine	China.
		P. densiflora	Korean red pine	Japan, the Korean Peninsula, northeastern China (Heilongjiang, Jilin, Liaoning, Shandong) and the extreme southeast of Russia (southern Primorsky Krai).
		P. fragilissima		southeastern Taiwan.
		P. henryi	Henry's pine	China.
		P. hwangshanensis	Huangshan pine	eastern China, in the provinces of Anhui, Fujian, Guizhou, Hubei, Hunan, Jiangxi, and Zhejiang
		P. kesiya	Khasi pine	the Khasi Hills in the northeast Indian state of Meghalaya, to northern Thailand, Philippines, Burma, Cambodia, Laos, southernmost China, and Vietnam.
		P. latteri	Tenasserim pine	southeastern Burma, northern Thailand, Laos, Cambodia, Vietnam, and on the Chinese island of Hainan.
		P. luchuensis	Luchu pine	Ryukyu Islands of Japan
		P. massoniana	Masson's pine	Taiwan, and a wide area of central and southern China, including Hong Kong, and northern Vietnam,
		P. merkusii	Sumatran pine	northern Sumatra, and with two outlying populations in central Sumatra on Mount Kerinci and Mount Talang, and in the Philippines on Mindoro and in the Zambales Mountains on western Luzon.
		P. mugo	mountain pine	southwestern to Central Europe and Southeast Europe.
		P. nigra	European black pine	Iberian Pensinsula to the eastern Mediterranean, on the Anatolian peninsula of Turkey on Corsica and Cyprus as well as Crimea and in the high mountains of Northwest Africa.
		P. resinosa	red pine	Newfoundland west to Manitoba, and south to Pennsylvania, with several smaller, disjunct populations occurring in the Appalachian Mountains in Virginia and West Virginia, as well as a few small pockets in extreme northern New Jersey and northern Illinois.
		P. sylvestris	Scots pine	Eurasia, ranging from Western Europe to Eastern Siberia, south to the Caucasus Mountains and Anatolia, and north to well inside the Arctic Circle in Fennoscandia
		P. tabuliformis	Chinese red pine	northern China from Liaoning west to Inner Mongolia and Gansu, and south to Shandong, Henan and Shaanxi, and also northern Korea.
		P. taiwanensis	Taiwan red pine	Taiwan.
		P. thunbergii	Japanese black pine	Japan (Kyūshū, Shikoku and Honshū) and South Korea.
		P. tropicalis	tropical pine	Cuba.
		P. yunnanensis	Yunnan pine	Yunnan, Sichuan, Guizhou, and Guangxi

Subsection Pinaster

Subsection Pinaster contains species native to the Mediterranean, as well as P. roxburghii from the Himalayas. The scales of its cones lack spines.[4] It is named after P. pinaster.

Image	Leaves & Cone	Scientific name	Common Name	Distribution
		P. brutia	Turkish pine	Turkey, but it also extends to southeastern most Bulgaria, the East Aegean Islands of Aegean Sea, Crete, the Crimea, Iran, Georgia, Azerbaijan, northern Iraq, western Syria, Israel, north-west Jordan, Lebanon, and Cyprus.
		P. canariensis	Canary Island pine	outer Canary Islands (Gran Canaria, Tenerife, El Hierro and La Palma)
		P. halepensis	Aleppo pine	Morocco, Algeria, Tunisia and Spain north to southern France, Malta, Italy, Croatia, Montenegro, and Albania, and east to Greece.
		P. heldreichii	Bosnian pine	Balkans and southern Italy.
		P. pinaster	maritime pine	Portugal and Northern Spain (especially in Galicia) to southern and Western France, east to western Italy, Croatia and south to northern Tunisia, Algeria and northern Morocco.
		P. pinea	stone pine	Southern Europe, The Palestinian Territories, Lebanon and Syria.
		P. roxburghii	chir pine	Tibet and Afghanistan through Pakistan, across northern India in Jammu and Kashmir, Himachal Pradesh, Uttarakhand, Sikkim, Arunachal Pradesh), Nepal and Bhutan, to Myanmar.

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.[5]

Subsection Australes

Subsection Australes is native to North and Central America and islands in the Caribbean.[4][7][8] The closed-cone (serotinous) species of California and Baja California, P. attenuata, P. muricata, and P. radiata, are sometimes placed in a separate subsection, Attenuatae.[9]

Image	Leaves & Cone	Scientific name	Common Name	Distribution
		P. attenuata	knobcone pine	southern Oregon to Baja California with the greatest concentration in northern California and the Oregon-California border.
		P. caribaea	Caribbean pine	Central America, Cuba, the Bahamas, and the Turks and Caicos Islands.
		P. cubensis	Cuban pine	eastern highlands of the island of Cuba
		P. echinata	shortleaf pine	eastern United States from southernmost New York, south to northern Florida, west to eastern Oklahoma, and southwest to eastern Texas
		P. elliottii	slash pine	Southeastern United States.
		P. glabra	spruce pine	southern United States, from southern South Carolina south to northern Florida and west to southern Louisiana.
		P. greggii	Gregg's pine	eastern Mexico
		P. herrerae	Herrera's pine	Mexico.
		P. jaliscana	Jalisco pine	Mexico (Sierra Madre del Sur within western Jalisco state. )
		P. lawsonii	Lawson's pine	Mexico.
		P. leiophylla	Chihuahua pine	Mexico along the Sierra Madre Occidental and Sierra Madre del Sur from Chihuahua to Oaxaca, with a small extension into the United States in southeast Arizona and southwest New Mexico.
		P. lumholtzii	Lumholtz's pine	northwestern Mexico( states of Chihuahua, Durango, Zacatecas, Nayarit, Jalisco, Aguascalientes ).
		P. muricata	bishop pine	California, including several offshore Channel Islands, and a few locations in Baja California, Mexico.
		P. occidentalis	Hispaniolan pine	Dominican Republic and Haiti
		P. oocarpa	egg-cone pine	Mexico and Central America.
		P. palustris	longleaf pine	from East Texas to southern Maryland, extending into northern and central Florida
		P. patula	patula pine	Mexico (State of Veracruz on the Sierra Madre Oriental )
		P. praetermissa	McVaugh's pine	western Mexico.
		P. pringlei	Pringle's pine	Mexico.
		P. pungens	Table Mountain pine	Appalachian Mountains in the United States.
		P. radiata	Monterey pine	California and Mexico (Guadalupe Island and Cedros island)
		P. rigida	pitch pine	eastern North America, from central Maine south to Georgia and as far west as Kentucky, and in two pockets along the St. Lawrence River in southern Quebec and Ontario
		P. serotina	pond pine	United States, from southern New Jersey south to Florida and west to southern Alabama.
		P. taeda	loblolly pine	Southeastern United States, from East Texas to Florida, and north to southern New Jersey.
		P. tecunumanii	Tecun Uman pine	Mexico (Chiapas and Oaxaca) through Guatemala, Belize, El Salvador, Honduras to Nicaragua
		P. teocote	Teocote pine	Mexico.

Fossils

• †P. foisyi - extinct
• †P. matthewsii – Pliocene, Yukon Territory, Canada[10]

Subsection Contortae

Subsection Contortae is native to North America and Mexico.[4]

Image	Leaves & Cone	Scientific name	Common Name	Distribution
		P. banksiana	jack pine	Canada is east of the Rocky Mountains from the Mackenzie River in the Northwest Territories to Cape Breton Island in Nova Scotia, and the north-central and northeast of the United States from Minnesota to Maine, with the southernmost part of the range just into northwest Indiana and northwest Pennsylvania.
		P. clausa	sand pine	Southeastern United States.
		P. contorta	lodgepole pine	western North America
		P. virginiana	Virginia pine	Long Island in southern New York south through the Appalachian Mountains to western Tennessee and Alabama.

Subsection Ponderosae

Subsection Ponderosae is native to Central America, Mexico, the western United States, and southwestern Canada.[4][11]

Image	Leaves & Cone	Scientific name	Common Name	Distribution
		P. arizonica	Arizona pine	northern Mexico, southeast Arizona, southwest New Mexico, and western Texas in the United States.
		P. cooperi	Cooper's pine	Mexico.
		P. coulteri	Coulter pine	Southern California and northern Baja California (Mexico).
		P. devoniana	Michoacan pine	Mexico - from S. Sinaloa to Chiapas - and Guatemala
		P. douglasiana		Mexico
		P. durangensis	Durango pine	Sierra Madre Occidental mountain range of north-western Mexico.
		P. engelmannii	Apache pine	Northern Mexico, in the Sierra Madre Occidental with its range extending a short distance into the United States in southwestern New Mexico and southeastern Arizona.
		P. hartwegii	Hartweg's pine	Mexico and Central America east to Honduras.
		P. jeffreyi	Jeffrey pine	California, but also in the westernmost part of Nevada, southwestern Oregon, and northern Baja California.
		P. maximinoi	thinleaf pine	El Salvador, Guatemala, Honduras, and Mexico
		P. montezumae	Montezuma pine	Mexico and Central America
		P. ponderosa	ponderosa pine	western North America.
		P. pseudostrobus	smooth-bark Mexican pine	Sinaloa, Mexico to El Salvador and Honduras.
		P. sabiniana	gray pine	California in the United States.
		P. torreyana	Torrey pine	northern San Diego county, and on Santa Rosa Island.

Fossils

• †P. johndayensis – Oligocene

Subgenus Strobus

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.[5]

Section Parrya

Section Parrya has one to five needles per fascicle. The seeds either have articulate (jointed) wings or no wings at all. In all species except for P. nelsonii, the fascicle sheaths curl back to form a rosette 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.[5]

Subsection Balfourianae

Subsection Balfourianae (bristlecone pines) is native to southwest United States.

Image	Leaves & Cone	Scientific name	Common Name	Distribution
		P. aristata	Rocky Mountains bristlecone pine	Rocky Mountains in Colorado and northern New Mexico, with isolated populations in the San Francisco Peaks in Arizona and the Kaibab National Forest north of the Grand Canyon.
		P. balfouriana	foxtail pine	California, United States.
		P. longaeva	Great Basin bristlecone pine	California, Nevada, and Utah

Subsection Cembroides

Subsection Cembroides (pinyons or piñons) is native to Mexico and the southwestern United States.

Image	Leaves & Cone	Scientific name	Common Name	Distribution
		P. cembroides	Mexican pinyon	Arizona, Texas and New Mexico in the United States into Mexico.
		P. culminicola	Potosi pinyon	Mexico, northern Sierra Madre Oriental in Coahuila and Nuevo León
		P. edulis	Colorado pinyon	Colorado, southern Wyoming, eastern and central Utah, northern Arizona, New Mexico, western Oklahoma, southeastern California, and the Guadalupe Mountains in far western Texas
		P. johannis	Johann's pinyon	southeast Arizona and southwest New Mexico, United States, south in Mexico along the Sierra Madre Occidental and the Sierra Madre Oriental to southern Zacatecas and San Luis Potosí.
		P. maximartinezii	big-cone pinyon	Mexico, southern Sierra Madre Occidental in southern Zacatecas.
		P. monophylla	single-leaf pinyon	southernmost Idaho, western Utah, Arizona, southwest New Mexico, Nevada, eastern and southern California and northern Baja California.
		P. orizabensis	Orizaba pinyon	central Mexico.
		P. pinceana	weeping pinyon	Mexico, in the states of: Durango; northern Coahuila, Nuevo León, and Zacatecas; central San Luis Potosí; and southern Querétaro and Hidalgo.
		P. quadrifolia	Parry pinyon	southernmost California in the United States and northern Baja California in Mexico
		P. remota	Texas pinyon or papershell pinyon	southwestern Texas and northeastern Mexico.
		P. rzedowskii	Rzedowski's pinyon	western Michoacan state, in southwestern Mexico.

Subsection Nelsonianae

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

Image	Leaves & Cone	Scientific name	Common Name	Distribution
		P. nelsonii	Nelson's pinyon	northeastern Mexico, in Nuevo León, San Luis Potosí and Tamaulipas

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.[12][13]

Subsection Gerardianae

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

Image	Leaves & Cone	Scientific name	Common Name	Distribution
		P. bungeana	lacebark pine	northeastern and central China
		P. gerardiana	chilgoza pine	northwestern Himalayas in northern Pakistan, and northwest India
		P. squamata	Qiaojia pine	Qiaojia County, northeast Yunnan, China

Subsection Krempfianae

Subsection Krempfianae is native to Vietnam. It has two needles per fascicle, and they are atypically flattened. The cone scales are thick and have no prickles.

Image	Leaves & Cone	Scientific name	Common Name	Distribution
		P. krempfii	Krempf's Pine	Vietnam in the Da Lat–Nha Trang area

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

Image	Leaves & Cone	Scientific name	Common Name	Distribution
		P. albicaulis	whitebark pine	Sierra Nevada, Cascade Range, Pacific Coast Ranges, and Rocky Mountains from Wyoming northwards.
		P. amamiana	Yakushima white pine	Japan ( Yakushima and Tanegashima south of Kyūshū)
		P. armandii	Chinese white pine	China(southern Shanxi west to southern Gansu and south to Yunnan, with outlying populations in Anhui.)Taiwan, and northern Burma.
		P. ayacahuite	Mexican white pine	southern Mexico and western Central America
		P. bhutanica	Bhutan white pine	Bhutan and adjacent parts of northeast India (Arunachal Pradesh) and southwest China (Yunnan and Tibet).
		P. cembra	Swiss pine	the Alps and Carpathian Mountains of central Europe, in Poland (Tatra Mountains), Switzerland, France, Italy, Austria, Germany, Slovenia, Slovakia (Tatra Mountains), Ukraine and Romania.
		P. chiapensis	Chiapas pine	southern Mexico and Guatemala
		P. dabeshanensis	Dabieshan pine	Dabie Mountains in Anhui and Hubei provinces
		P. dalatensis	Vietnamese white pine	Indochina, Vietnam, Laos
		P. fenzeliana	Hainan white pine	Hainan
		P. flexilis	limber pine	Western United States, Mexico, and Canada.
		P. koraiensis	Korean pine	eastern Asia: Korea, northeastern China, Mongolia, the temperate rainforests of the Russian Far East, and central Japan.
		P. lambertiana	sugar pine	the Pacific coast of North America, from Oregon through California to Baja California.
		P. monticola	western white pine	western United States and Canada
		P. morrisonicola	Taiwan white pine	Taiwan
		P. parviflora	Japanese white pine	Korea and Japan.
		P. hakkodensis ( P. pumila × P. parviflora)	Hakkoda pine	Japan.
		P. peuce	Macedonian pine	Macedonia, Bulgaria, Albania, Montenegro, Kosovo, southwest of Serbia, and the extreme north of Greece
		P. pumila	Siberian dwarf pine	Far East, Eastern Siberia, north-east of Mongolia, north-east of China, northern Japan and Korea
		P. sibirica	Siberian pine	Siberia and Mongolia
		P. strobus	eastern white pine	Newfoundland, Canada west through the Great Lakes region to southeastern Manitoba and Minnesota, United States, and south along the Appalachian Mountains and upper Piedmont to northernmost Georgia
		P. strobiformis	Southwestern white pine (also Chihuahuan)	southwestern United States and Mexico.
		P. wallichiana	blue pine	eastern Afghanistan east across northern Pakistan and north west India to Yunnan in southwest China.
		P. wangii	Guangdong white pine	Yunnan Province of southern China

Incertae sedis

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

• †P. peregrinus – Pinus peregrinus – Middle Eocene, Golden Valley Formation, North Dakota, USA

References

1. Gernandt et al. 2005.
2. Zeb et al. 2019.
3. Frankis, Michael (2002). "Classification of the genus Pinus". The Lovett Pinetum Charitable Foundation.
4. Gernandt et al. 2005, p. 35.
5. Gernandt et al. 2005, p. 38.
6. Stockey 1983.
7. "Pinus cubensis". Germplasm Resources Information Network (GRIN). Agricultural Research Service (ARS), United States Department of Agriculture (USDA).
8. "Pinus occidentalis". Germplasm Resources Information Network (GRIN). Agricultural Research Service (ARS), United States Department of Agriculture (USDA).
9. Earle, Christopher J., ed. (2018). "Pinus". The Gymnosperm Database.
10. McKown, Stockey & Schweger 2002.
11. "Pinus hartwegii". Germplasm Resources Information Network (GRIN). Agricultural Research Service (ARS), United States Department of Agriculture (USDA).
12. Gernandt et al. 2005, pp. 38–39.
13. "Pinus chiapensis". Germplasm Resources Information Network (GRIN). Agricultural Research Service (ARS), United States Department of Agriculture (USDA).
14. "Eastern white pine (Pinus strobus)". Minnesota Department of Natural Resources. St. Paul, Minnesota.

Bibliography

• Gernandt, D. S.; López, G. G.; García, S. O.; Liston, A. (2005). "Phylogeny and classification of Pinus". Taxon. 54 (1): 29–42. 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. doi:10.1086/340425. 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.
• 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 (published March 2020). 58 (2): 118–132. doi:10.1111/jse.12492. ISSN 1759-6831.

External links

• Tree of Life Web favors classification of Ducampopinus species in Strobus.
• NCBI Taxonomy server files Ducampopinus species above as Strobus.