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Pinus A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family (biology), family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanic ...
'', the pines, is a genus of approximately 111
extant Extant 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 * Extant taxon, a taxon which is not extin ...
tree and shrub species. The genus is currently split into two
subgenera In biology, a subgenus (plural: 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 t ...
: 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 known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
DNA sequencing and whole
plastid The plastid (Greek: πλαστός; plastós: formed, molded – plural plastids) is a membrane-bound organelle found in the cells of plants, algae, and some other eukaryotic organisms. They are considered to be intracellular endosymbiotic cyanoba ...
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 Californian botanist
John Gill Lemmon John Gill ("J.G.") Lemmon (January 2, 1831 or 1832, Lima, Michigan – November 24, 1908 Oakland, California) was an American botanist and Civil War veteran and former prisoner of Andersonville. He was married to fellow botanist, Sara Plummer Lem ...
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 sheath 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.Shashtri, Varun. Dictionary of Botany. Publisher: Isha Books 2005. ...
s 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 leaves, usually in the autumn; to the shedding of petals, afte ...
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 (in formal botanical usage: strobilus, plural strobili) is a seed-bearing organ on gymnosperm plants. It is usually woody, ovoid to globular, including scales and bracts arranged around a central axis, especially in conifers an ...
s. Both subgenera are thought to have a very ancient divergence from one another, having diverged during the late Jurassic.


Subgenus ''Pinus''

Subgenus ''
Pinus A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family (biology), family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanic ...
'' 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 A cone is a three-dimensional geometric shape that tapers smoothly from a flat base (frequently, though not necessarily, circular) to a point called the apex or vertex. A cone is formed by a set of line segments, half-lines, or lines con ...
either open soon after they mature or are
serotinous Serotiny in botany simply means 'following' or 'later'. In the case of serotinous flowers, it means flowers which grow following the growth of leaves, or even more simply, flowering later in the season than is customary with allied species. Havi ...
.


Section ''Pinus''

Section ''
Pinus A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family (biology), family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanic ...
'' 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 Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
, 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 north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the ea ...
, except for ''P. resinosa'' in northeastern
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the C ...
and ''P. tropicalis'' in western Cuba. Subsection ''
Incertae sedis ' () or ''problematica'' is a term used for a 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, uncertaint ...
'' *†'' P. driftwoodensis'' – Early
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', "daw ...
, British Columbia, Canada


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'' – European black 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 planet's highest peaks, including the very highest, Mount Everest. Over 100 ...
. 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 Serotiny in botany simply means 'following' or 'later'. In the case of serotinous flowers, it means flowers which grow following the growth of leaves, or even more simply, flowering later in the season than is customary with allied species. Havi ...
. 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.


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 or geography. Etymology The word ''north'' is ...
and
Central America Central America ( es, América Central or ) is a subregion of the Americas. Its boundaries are defined as bordering the United States to the north, Colombia to the south, the Caribbean Sea to the east, and the Pacific Ocean to the west. C ...
and islands in the
Caribbean The Caribbean (, ) ( es, El Caribe; french: la Caraïbe; ht, Karayib; nl, De Caraïben) is a region of the Americas that consists of the Caribbean Sea, its islands (some surrounded by the Caribbean Sea and some bordering both the Caribbean S ...
. 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. 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 in the geologic time scale that extends from 5.333 million to 2.58Yukon Territory Yukon (; ; formerly called Yukon Territory and also referred to as the Yukon) is the smallest and westernmost of Canada's three territories. It also is the second-least populated province or territory in Canada, with a population of 43,964 as ...
, 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


Subsection ''Contortae''

Subsection '' Contortae'' is native to North America and Mexico. *'' P. banksiana'' – jack pine *'' P. clausa'' – sand pine *'' P. contorta contorta'' – shore pine *'' P. contorta latifolia'' – lodgepole pine *'' P. virginiana'' – Virginia pine


Subsection ''Ponderosae''

Subsection '' Ponderosae'' is native to
Central America Central America ( es, América Central or ) is a subregion of the Americas. Its boundaries are defined as bordering the United States to the north, Colombia to the south, the Caribbean Sea to the east, and the Pacific Ocean to the west. C ...
,
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
, the western
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territo ...
, and southwestern Canada, 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. douglasiana'' *'' 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 of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
*'' 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 leaves, usually in the autumn; to the shedding of petals, afte ...
, 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 A cone is a three-dimensional geometric shape that tapers smoothly from a flat base (frequently, though not necessarily, circular) to a point called the apex or vertex. A cone is formed by a set of line segments, half-lines, or lines con ...
usually open soon after they mature.


Section ''Parrya''

Section '' Parrya'' has one to five needles per fascicle. The seeds either have
articulate Articulate may refer to: * '' Articulate!'', a board game in which players describe words from different categories * Articulate brachiopods, brachiopods with toothed hinges and simple opening and closing muscles * Articulate sound, to move the ...
(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 (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territo ...
. *'' P. aristata'' – Rocky Mountains bristlecone pine *'' P. balfouriana'' – foxtail pine *'' P. longaeva'' – Great Basin bristlecone pine


Subsection ''Cembroides''

Subsection ''Cembroides (pinyons or piñons) is native to
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
and the southwestern
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territo ...
. *'' 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'' 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 ''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 ...
'' 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. 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. 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. The formation, named for the type location designated in 1962, Klondike Mountain north of Republic, Washi ...
,
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 abun ...
- 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. The formation, named for the type location designated in 1962, Klondike Mountain north of Republic, Washi ...
,
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 abun ...
- Okanagan Highlands Floras *†'' Pinus peregrinus'' – Middle
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', "daw ...
,
Golden Valley Formation The Golden Valley Formation is a stratigraphic unit of Late Paleocene to Early Eocene age in the Williston Basin of North Dakota.Hickey, 1977 It is present in western North Dakota and was named for the city of Golden Valley by W.E. Benson and W.M. ...
,
North Dakota North Dakota () is a U.S. state in the Upper Midwest, named after the indigenous Dakota Sioux. North Dakota is bordered by the Canadian provinces of Saskatchewan and Manitoba to the north and by the U.S. states of Minnesota to the east, South ...
, USA *†'' Pinus tetrafolia'' - 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. The formation, named for the type location designated in 1962, Klondike Mountain north of Republic, Washi ...
- 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 (biology), family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanic ...
Pinus A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family (biology), family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanic ...
Pinus A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family (biology), family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanic ...