Pinus subsection Ponderosae
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'' Pinus'', 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 extinct, ...
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 Section, Sectioning or Sectioned may refer to: Arts, entertainment and media * Section (music), a complete, but not independent, musical idea * Section (typography), a subdivision, especially of a chapter, in books and documents ** Section sig ...
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 Cell (biology), cells of plants, algae, and some other eukaryotic organisms. They are considered to be intracellular endosy ...
genomic analysis. Older classifications split the genus into three subgenera – subgenus ''Pinus'', subgenus ''Strobus'', and subgenus ''Ducampopinus'' ( pinyon,
bristlecone The term bristlecone pine covers three species of pine tree (family Pinaceae, genus '' Pinus'', subsection ''Balfourianae''). All three species are long-lived and highly resilient to harsh weather and bad soils. One of the three species, ''P ...
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 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 Fascicle or ''fasciculus'' may refer to: Anatomy and histology * Muscle fascicle, a bundle of skeletal muscle fibers * Nerve fascicle, a bundle of axons (nerve fibers) ** Superior longitudinal fasciculus *** Arcuate fasciculus ** Gracile fas ...
, 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 or
persistent Persistent may refer to: * Persistent data * Persistent data structure * Persistent identifier * Persistent memory * Persistent organic pollutant * Persistent Systems, a technology company * USS ''Persistent'', three United States Navy ships See ...
, the number of
fibrovascular bundle A vascular bundle is a part of the transport system in vascular plants. The transport itself happens in the stem, which exists in two forms: xylem and phloem. Both these tissues are present in a vascular bundle, which in addition will includ ...
s per needle (2 in ''Pinus'' or 1 in ''Strobus''), the position of the
resin duct Resin canals or resin ducts are elongated, tube-shaped intercellular spaces surrounded by epithelial cells which secrete resin into the canal. These canals are orientated longitudinally and radially in between fusiform rays. They are usually fou ...
s 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'' includes the yellow and hard pines. Pines in this subgenus have one to five needles per fascicle and two
fibrovascular bundle A vascular bundle is a part of the transport system in vascular plants. The transport itself happens in the stem, which exists in two forms: xylem and phloem. Both these tissues are present in a vascular bundle, which in addition will includ ...
s per needle, and the fascicle sheaths are
persistent Persistent may refer to: * Persistent data * Persistent data structure * Persistent identifier * Persistent memory * Persistent organic pollutant * Persistent Systems, a technology company * USS ''Persistent'', three United States Navy ships See ...
, 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'' 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 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 Car ...
and ''P. tropicalis'' in western Cuba. Subsection '' Incertae sedis'' *†'' P. driftwoodensis'' – Early Eocene, 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 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.


Subsection ''Australes''

Subsection ''Australes'' is native to North and Central America 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 Se ...
. 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, Yukon Territory, 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 Contortae as a term has appeared in several senses in botanical taxonomy, most conspicuously as follows: Order Contortae Order Contortae is no longer recognised in botanical taxonomy. After a varied history in the nineteenth century, predominant o ...
'' 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, Mexico, the western United States, 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 The Willamette Valley ponderosa pine is a population of the ponderosa pine (''Pinus ponderosa'') native to the Willamette Valley in Oregon. It is adapted for Western Oregon's wet winter and dry summer. History The Willamette Valley ponderosa va ...
'' - 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 A vascular bundle is a part of the transport system in vascular plants. The transport itself happens in the stem, which exists in two forms: xylem and phloem. Both these tissues are present in a vascular bundle, which in addition will includ ...
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.


Section ''Parrya''

Section ''
Parrya ''Parrya'' is a genus of flowering plants in the family Brassicaceae, found in the arctic and subarctic biomes of the Northern Hemisphere, and in Central Asia. The center of diversity is the Tian Shan and Pamir-Alay The Pamir-Alay (also ''Pami ...
'' 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. *'' 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 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.


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 ''Pinus krempfii'', or Krempf's pine, is a rare species of pine, endemic to the central highlands of Vietnam in the Da Lat–Nha Trang Nha Trang ( or ; ) is a coastal city and capital of Khánh Hòa Province, on the South Central Coast of ...
*†'' P. leptokrempfii'' - Oligocene


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. 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 ''Pinus latahensis'' is an extinct species of conifer in the pine family Pinaceae. The species is known from fossil leaves found in the early Eocene deposits of northern Washington state, United States, and southern British Columbia, Canada. A ...
'' - 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, Wash ...
, Allenby Formation -
Okanagan Highlands Floras The Okanagan ( ), also known as the Okanagan Valley and sometimes as the Okanagan Country, is a region in the Canadian province of British Columbia defined by the basin of Okanagan Lake and the Canadian portion of the Okanagan River. It is part ...
*†'' 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, Wash ...
, Allenby Formation -
Okanagan Highlands Floras The Okanagan ( ), also known as the Okanagan Valley and sometimes as the Okanagan Country, is a region in the Canadian province of British Columbia defined by the basin of Okanagan Lake and the Canadian portion of the Okanagan River. It is part ...
*†''
Pinus peregrinus ''Pinus peregrinus'' is an extinction, extinct species of pine in the family Pinaceae known from Clarkforkian age Paleocene fossils found in western North Dakota, USA. The species was described from three wing seed specimens found at localities ...
'' – Middle Eocene,
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, USA *†''
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. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden accep ...
'' - 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, Wash ...
-
Okanagan Highlands Floras The Okanagan ( ), also known as the Okanagan Valley and sometimes as the Okanagan Country, is a region in the Canadian province of British Columbia defined by the basin of Okanagan Lake and the Canadian portion of the Okanagan River. It is part ...


See also

*
Hybridization in pines Both naturally and artificially occurring pine species ''Pine, (Pinus)'' can Hybrid (biology), hybridize, combining their genetic material and sometimes creating hybrids that can be more or less Heterosis, vigorous than their parent species. An exam ...
(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 Pinus Pinus