List of Pinus species

Last updated May 12, 2024

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]

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]

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

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.^[6]

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.^[6]

Subsection Incertae sedis

† P. driftwoodensis – Early Eocene, British Columbia, Canada ^[7]

Subsection Pinus

All but two species (P. resinosa and P. tropicalis) in Subsection Pinus 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

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.

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

Pinus pinea
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.^[6]

Phylogenetic analysis supports ancient divergences within this section, with subsections Australes and Ponderosae having diverged during the mid-Cretaceous.^[5]

Subsection Australes

Subsection Australes is native to North and Central America and islands in the Caribbean.^[4]^[8]^[9]

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.^[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

Subsection Contortae

Subsection Contortae 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

Subsection Ponderosae

Subsection Ponderosae is native to Central America, Mexico, the western United States, and southwestern Canada,^[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. douglasiana
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

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.^[6]

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.^[6]

Subsection Balfourianae

Subsection Balfourianae (bristlecone pines) is native to southwest United 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

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. 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

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 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.^[17]^[18]

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^[19]

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.^[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. 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

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

Related Research Articles

A pine is any conifer tree or shrub in the genus Pinus of the family Pinaceae. Pinus is the sole genus in the subfamily Pinoideae.

The pinyon or piñon pine group grows in southwestern North America, especially in New Mexico, Colorado, Arizona, and Utah. The trees yield edible nuts, which are a staple food of Native Americans, and widely eaten as a snack and as an ingredient in New Mexican cuisine. The name comes from the Spanish pino piñonero, a name used for both the American varieties and the stone pine common in Spain, which also produces edible nuts typical of Mediterranean cuisine. Harvesting techniques of the prehistoric American Indians are still used today to collect the pinyon seeds for personal use or for commercialization. The pinyon nut or seed is high in fats and calories. In the western United States, pinyon pines are often found in pinyon–juniper woodlands.

Pinus albicaulis, known by the common names whitebark pine, white bark pine, white pine, pitch pine, scrub pine, and creeping pine, is a conifer tree native to the mountains of the western United States and Canada, specifically subalpine areas of the Sierra Nevada, Cascade Range, Pacific Coast Ranges, and Rocky Mountains. It shares the common name "creeping pine" with several other plants.

Western white pine, also called silver pine and California mountain pine, is a species of pine in the family Pinaceae. It occurs in mountain ranges of northwestern North America and is the state tree of Idaho.

Pinus flexilis, the limber pine, is a species of pine tree-the family Pinaceae that occurs in the mountains of the Western United States, Mexico, and Canada. It is also called Rocky Mountain white pine.

Pinus strobiformis, commonly known as southwestern white pine, Mexican white pine or Chihuahua white pine, is a medium-sized white pine tree whose native habitat is in southwestern United States and Mexico. It is typically a high-elevation pine growing mixed with other conifers.

Pinus canariensis, the Canary Island pine, is a species of gymnosperm in the conifer family Pinaceae. It is a large, evergreen tree, native and endemic to the outer Canary Islands of the Atlantic Ocean.

Pinus virginiana, the Virginia pine, scrub pine, Jersey pine, possum pine, is a medium-sized tree, often found on poorer soils from Long Island in southern New York south through the Appalachian Mountains to western Tennessee and Alabama. The usual size range for this pine is 9–18 m, but can grow larger under optimum conditions. The trunk can be as large as 20 inches diameter. This tree prefers well-drained loam or clay, but will also grow on very poor, sandy soil, where it remains small and stunted. The typical life span is 65 to 90 years.

Pinus cembra, also known as Swiss pine, Swiss stone pine, Arolla pine, Austrian stone pine, or just stone pine, is a species of pine tree in the subgenus Strobus.

Pinus sibirica, or Siberian pine, in the family Pinaceae is a species of pine tree that occurs in Siberia from 58°E in the Ural Mountains east to 126°E in the Stanovoy Range in southern Sakha Republic, and from Igarka at 68°N in the lower Yenisei valley, south to 45°N in central Mongolia.

Pinus johannis, the Johann's pine, is a pine in the pinyon pine group, native to North America. The range extends from 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. It occurs at moderate to high altitudes, from 1,600–3,000 metres (5,200–9,800 ft), in cool, dry climate conditions.

Pinus culminicola, commonly known as Potosí pinyon or Potosí Piñón, is a pine in the pinyon pine group, native and endemic to northeast Mexico. The range is highly localised, confined to a small area of high summits in the northern Sierra Madre Oriental in Coahuila and Nuevo León, and only abundant on the highest peak, Cerro Potosí. It occurs at very high altitudes, from 3000–3700 m, in cool, moist subalpine climate conditions.

Pinus edulis, the Colorado pinyon, two-needle piñon, pinyon pine, or simply piñon, is a pine in the pinyon pine group native to the Southwestern United States, used for its edible pine nuts.

Pinus monophylla, the single-leaf pinyon, is a pine in the pinyon pine group, native to North America. The range is in southernmost Idaho, western Utah, Arizona, southwest New Mexico, Nevada, eastern and southern California and northern Baja California.

Pinus aristata, the Rocky Mountain bristlecone pine, is a long-living species of bristlecone pine tree native to the United States. It appears in the 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. It is usually found at very high altitudes, from 7,000–13,000 feet (2,100–4,000 m), in cold, dry subalpine climate conditions, often at the tree line, although it also forms extensive closed-canopy stands at somewhat lower elevations.

Pinus maximartinezii, called Martinez pinyon, big-cone pinyon or maxipiñon, is a pine in the pinyon pine group, native to west-central Mexico.

Pinus dalatensis, also known as Vietnamese white pine or Dalat pine, is a species of pine endemic to Indochina. In Vietnam it grows in the mountains of the central and south-central parts of the country at elevations of 1,400 to 2,300 metres. Only recently confirmed from Laos, the population located within the Nakai-Nam Theun Biodiversity Conservation Area is the largest, at the lowest elevation, and the northernmost of the known populations of P. dalatensis.

Pinus nelsonii, Nelson's pinyon, is a species of pine native to the mountains of northeastern Mexico, in Nuevo León, San Luis Potosí and Tamaulipas at 1,800–3,200 m altitude.

In botany, a fascicle is a bundle of leaves or flowers growing crowded together; alternatively the term might refer to the vascular tissues that supply such an organ with nutrients. However, vascular tissues may occur in fascicles even when the organs they supply are not fascicled.

Pinus driftwoodensis is an extinct species of conifer in the pine family solely known from early Eocene sediments exposed in south central British Columbia. The species was described from an isolated fossil ovulate cone associated with a series of wood, needles and pollen cones in chert. P. driftwoodensis was the eighth pine species to be described from a permineralized ovulate cone and the second from the Okanagan highlands.

References

1 2 Gernandt et al. 2005.
↑ Zeb et al. 2019.
↑ Frankis, Michael (2002). "Classification of the genus Pinus". The Lovett Pinetum Charitable Foundation.
1 2 3 4 5 6 Gernandt et al. 2005, p. 35.
1 2 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 (July 19, 2021). "Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms". Nature Plants. 7 (8): 1015–1025. doi:10.1038/s41477-021-00964-4. ISSN 2055-0278. PMID 34282286. S2CID 236141481.
1 2 3 4 5 6 Gernandt et al. 2005, p. 38.
↑ Stockey 1983.
↑ "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.
↑ Earle, Christopher J., ed. (2018). "Pinus". The Gymnosperm Database.
↑ 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 hartwegii". Germplasm Resources Information Network . Agricultural Research Service, United States Department of Agriculture.
1 2 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.
↑ 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. doi:10.1086/713957. S2CID 233888471.
↑ "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. 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

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

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[FOOTNOTEGernandtLópezGarcíaListon2005-1] 1 2 Gernandt et al. 2005.

[FOOTNOTEZebDongZhangWang2019-2] Zeb et al. 2019.

[3] Frankis, Michael (2002). "Classification of the genus Pinus". The Lovett Pinetum Charitable Foundation.

[FOOTNOTEGernandtLópezGarcíaListon200535-4] 1 2 3 4 5 6 Gernandt et al. 2005, p. 35.

[Stull-2021-5] 1 2 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 (July 19, 2021). "Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms". Nature Plants. 7 (8): 1015–1025. doi:10.1038/s41477-021-00964-4. ISSN 2055-0278. PMID 34282286. S2CID 236141481.

[FOOTNOTEGernandtLópezGarcíaListon200538-6] 1 2 3 4 5 6 Gernandt et al. 2005, p. 38.

[FOOTNOTEStockey1983-7] Stockey 1983.

[8] "Pinus cubensis". Germplasm Resources Information Network . Agricultural Research Service, United States Department of Agriculture.

[9] "Pinus occidentalis". Germplasm Resources Information Network . Agricultural Research Service, United States Department of Agriculture.

[10] Earle, Christopher J., ed. (2018). "Pinus". The Gymnosperm Database.

[WeheMariGonzAgui20-11] 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.

[FOOTNOTEMcKownStockeySchweger2002-12] McKown, Stockey & Schweger 2002.

[13] "Pinus hartwegii". Germplasm Resources Information Network . Agricultural Research Service, United States Department of Agriculture.

[Yamada2015-14] 1 2 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.

[Knowlton1923-15] 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 .

[WolfeSchorn1990-16] 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 .

[FOOTNOTEGernandtLópezGarcíaListon200538–39-17] Gernandt et al. 2005, pp. 38–39.

[18] "Pinus chiapensis". Germplasm Resources Information Network . Agricultural Research Service, United States Department of Agriculture.

[19] 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. doi:10.1086/713957. S2CID 233888471.

[20] "Eastern white pine (Pinus strobus)". Minnesota Department of Natural Resources. St. Paul, Minnesota.

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