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Pinus 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. ''World Flora Online'' accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as c ...
'', the pines, is a genus of approximately 111
extant Extant or Least-concern species, least concern is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Exta ...
tree and shrub species. The genus is currently split into two
subgenera In biology, a subgenus ( subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between the ge ...
: subgenus ''Pinus'' (hard pines), and subgenus ''Strobus'' (soft pines). Each of the subgenera have been further divided into sections based on
chloroplast A chloroplast () is a type of membrane-bound organelle, organelle known as a plastid that conducts photosynthesis mostly in plant cell, plant and algae, algal cells. Chloroplasts have a high concentration of chlorophyll pigments which captur ...
DNA sequencing and whole
plastid A plastid is a membrane-bound organelle found in the Cell (biology), cells of plants, algae, and some other eukaryotic organisms. Plastids are considered to be intracellular endosymbiotic cyanobacteria. Examples of plastids include chloroplasts ...
genomic analysis. Older classifications split the genus into three subgenera – subgenus ''Pinus'', subgenus ''Strobus'', and subgenus ''Ducampopinus'' ( pinyon, bristlecone and lacebark pines) – 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. 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
California California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...
n 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 In the fields of horticulture and botany, the term deciduous () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed Leaf, leaves, usually in the autumn; to the shedding of petals, aft ...
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 cone A conifer cone, or in formal botanical usage a strobilus, : strobili, is a seed-bearing organ on gymnosperm plants, especially in conifers and cycads. They are usually woody and variously conic, cylindrical, ovoid, to globular, and have scale ...
s. Both subgenera are thought to have a very ancient divergence from one another, having diverged during the late
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
.


Subgenus ''Pinus''

Subgenus ''
Pinus 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. ''World Flora Online'' accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as c ...
'' 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.


Section ''Pinus''

Section ''
Pinus 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. ''World Flora Online'' accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as c ...
'' 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 Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
,
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
, and the
Mediterranean The Mediterranean Sea ( ) is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean basin and almost completely enclosed by land: on the east by the Levant in West Asia, on the north by Anatolia in West Asia and Southern ...
, except for ''P. resinosa'' in northeastern
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
and ''P. tropicalis'' in western
Cuba Cuba, officially the Republic of Cuba, is an island country, comprising the island of Cuba (largest island), Isla de la Juventud, and List of islands of Cuba, 4,195 islands, islets and cays surrounding the main island. It is located where the ...
. Subsection ''
Incertae sedis or is a term used for a taxonomy (biology), taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
'' *†'' P. driftwoodensis'' – Early
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
,
British Columbia British Columbia is the westernmost Provinces and territories of Canada, province of Canada. Situated in the Pacific Northwest between the Pacific Ocean and the Rocky Mountains, the province has a diverse geography, with rugged landscapes that ...
,
Canada Canada is a country in North America. Its Provinces and territories of Canada, ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, making it the world's List of coun ...


Subsection ''Pinus''

All but two species (''P. resinosa'' and ''P. tropicalis'') in Subsection ''Pinus'' are native to Eurasia. *'' 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 Himalayas, or Himalaya ( ), is a mountain range in Asia, separating the plains of the Indian subcontinent from the Tibetan Plateau. The range has some of the Earth's highest peaks, including the highest, Mount Everest. More than list of h ...
. The scales of its cones lack spines. 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 *'' 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. Phylogenetic analysis supports ancient divergences within this section, with subsections ''Australes'' and ''Ponderosae'' having diverged during the mid-
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
.


Subsection ''Australes''

Subsection ''Australes'' is native to
North North is one of the four compass points or cardinal directions. It is the opposite of south and is perpendicular to east and west. ''North'' is a noun, adjective, or adverb indicating Direction (geometry), direction or geography. Etymology T ...
and
Central America Central America is a subregion of North America. Its political boundaries are defined as bordering Mexico to the north, Colombia to the southeast, the Caribbean to the east, and the Pacific Ocean to the southwest. Central America is usually ...
and islands in the
Caribbean The Caribbean ( , ; ; ; ) is a region in the middle of the Americas centered around the Caribbean Sea in the Atlantic Ocean, North Atlantic Ocean, mostly overlapping with the West Indies. Bordered by North America to the north, Central America ...
. 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''. *'' 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'' *†'' P. matthewsii'' –
Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58Yukon Territory Yukon () is a territory of Canada, bordering British Columbia to the south, the Northwest Territories to the east, the Beaufort Sea to the north, and the U.S. state of Alaska to the west. It is Canada’s westernmost territory and the smallest ...
, Canada *'' 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. *'' 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 Central America is a subregion of North America. Its political boundaries are defined as bordering Mexico to the north, Colombia to the southeast, the Caribbean to the east, and the Pacific Ocean to the southwest. Central America is usually ...
,
Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...
, the western
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
, and southwestern
Canada Canada is a country in North America. Its Provinces and territories of Canada, ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, making it the world's List of coun ...
, although its former range was possibly much wider as evidenced by upper Miocene fossils belonging to this subsection found in Japan *'' 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'' *'' P. hartwegii'' – Hartweg's pine *'' P. jeffreyi'' – Jeffrey pine *†'' P. johndayensis'' –
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
*'' 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 In the fields of horticulture and botany, the term deciduous () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed Leaf, leaves, usually in the autumn; to the shedding of petals, aft ...
, 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.


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.


Subsection ''Balfourianae''

Subsection Balfourianae (bristlecone pines) is native to southwest
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
. *'' P. aristata'' – Rocky Mountains bristlecone pine *'' P. balfouriana'' – foxtail pine *†'' P. crossii'' - (
Chattian The Chattian is, in the geologic timescale, the younger of two ages or upper of two stages of the Oligocene Epoch/Series. It spans the time between . The Chattian is preceded by the Rupelian and is followed by the Aquitanian (the lowest stage ...
; Creede Formation, Colorado) *'' P. longaeva'' – Great Basin bristlecone pine


Subsection ''Cembroides''

Subsection ''Cembroides'' (pinyons or piñons) is native to
Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...
and the southwestern
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
. *'' 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.


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


Subsection ''Strobus''

Subsection ''
Strobus ''Pine, Pinus'', the pines, is a genus of approximately 111 Neontology, extant tree and shrub species. The genus is currently split into two subgenus, subgenera: subgenus ''Pinus'' (hard pines), and subgenus ''Strobus'' (soft pines). Each of the ...
'' has five needles per fascicle and thin cone scales with no prickles. Needles tend to be flexible and soft with slightly lighter side underneath. It is native to North and Central America, Europe, and Asia. *'' 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 Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * Late (The 77s album), ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudo ...
, 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


''Incertae sedis''

Species which are not placed in a subgenus at this time. *†'' Pinus latahensis'' – Early Eocene,
Klondike Mountain Formation The Klondike Mountain Formation is an Early Eocene (Ypresian) geological formation located in the northeast central area of Washington (state), Washington state. The formation is composed of volcanic rocks in the upper unit and volcanic plus lacu ...
,
Allenby Formation The Allenby formation is a sedimentary rock formation in British Columbia which was deposited during the Ypresian stage of the Early Eocene. It consists of conglomerates, sandstones with interbedded shales and coal. The shales contain an ...
Okanagan Highlands Floras *†'' Pinus macrophylla'' – Early Eocene,
Klondike Mountain Formation The Klondike Mountain Formation is an Early Eocene (Ypresian) geological formation located in the northeast central area of Washington (state), Washington state. The formation is composed of volcanic rocks in the upper unit and volcanic plus lacu ...
,
Allenby Formation The Allenby formation is a sedimentary rock formation in British Columbia which was deposited during the Ypresian stage of the Early Eocene. It consists of conglomerates, sandstones with interbedded shales and coal. The shales contain an ...
Okanagan Highlands Floras *†'' Pinus peregrinus'' – Middle
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
, Golden Valley Formation,
North Dakota North Dakota ( ) is a U.S. state in the Upper Midwest, named after the indigenous Dakota people, Dakota and Sioux peoples. It is bordered by the Canadian provinces of Saskatchewan and Manitoba to the north and by the U.S. states of Minneso ...
, US *†''
Pinus tetrafolia 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. ''World Flora Online'' accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as cu ...
'' – Early Eocene,
Klondike Mountain Formation The Klondike Mountain Formation is an Early Eocene (Ypresian) geological formation located in the northeast central area of Washington (state), Washington state. The formation is composed of volcanic rocks in the upper unit and volcanic plus lacu ...
Okanagan Highlands Floras


See also

* Hybridization in pines (list of pine hybrids)


References


Bibliography

* * * *


External links


Tree of Life Web
– favors classification of ''Ducampopinus'' species in ''Strobus''.
NCBI Taxonomy server
– files ''Ducampopinus'' species above as ''Strobus''. {{DEFAULTSORT:Pinus Classification *
Pinus 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. ''World Flora Online'' accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as c ...
Pinus 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. ''World Flora Online'' accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as c ...
Pinus 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. ''World Flora Online'' accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as c ...