List of Pinus species


Pinus, the pines, is a genus of approximately 110–120 extant tree and shrub species. The genus is currently split into two subgenera, subgenus Pinus, and subgenus Strobus. Each of the subgenera have been further divided into sections and subsections based in the past on morphology, ecology and biogeography, and more recently increasingly from chloroplast DNA sequencing and whole plastid genomic analysis. While the genetic analysis has given robust results at the higher levels, they often give conflicting results lower in the phylogenetic trees, with species allocated to different subsections by different studies or even within a study. Within subsections, the genetic relationships between species can be even more complex and conflicting; in one study, three samples of the very distinctive and morphologically constant Pinus lambertiana were placed in three different clades of the subsection Strobus, and similar problems with many other species with widespread nonmonophyly.
Several features are used to distinguish the subgenera, sections, and subsections of pines; the number of leaves per fascicle, whether the fascicle sheaths are deciduous or persistent, the number of fibrovascular bundles per needle, the position of the resin ducts in the needles, the presence or shape of the seed wings, and the position of the umbo and presence of a prickle on the scales of the seed cones.
The two subgenera are thought to have ancient divergence from each other, having diverged at some point between the late Jurassic, the mid Cretaceous, to the late Cretaceous.

Subgenus ''Pinus''

Subgenus Pinus includes the yellow and hard pines. Pines in this subgenus have two to five needles per fascicle. They have two fibrovascular bundles per needle, and the fascicle sheaths are persistent, except in P. leiophylla and P. lumholtzii. The cone scales are thicker and more rigid than those of subgenus Strobus, and have a resin sealing band before opening ; the cones either open soon after they mature or are serotinous.

Section ''Pinus''

Section Pinus has two or three needles per fascicle. The cones have moderately thin to thick scales; in most they open at maturity, but are weakly serotinous in some species in subsection Pinaster. 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.
Subsection Incertae sedis
All but two species in Subsection Pinus are native to Eurasia. The cones have moderate thickness scales, and are characterised by a slighty off-centre prickle on the umbo.
This pine is commonly resolved into subsection Pinaster by genetic studies, but this is in strong conflict with its morphology and ecology, which is much closer to subsection Pinus, or alternatively placed in its own subsection Leucodermes close to subsection Pinus.
These pines are placed in subsection Pinus by some genetic studies, but in subsection Pinaster by others. In morphology and ecology, they belong in the latter group. The subsection Merkusia has also been proposed for them.
Subsection Pinaster contains species native to warm climates in the Mediterranean region at low altitudes, as well as P. roxburghii from the Himalayas. The cones are thick-scaled and orange-brown to red-brown, and the cone scales are glossy and lack umbo spines. It is named after P. pinaster.
Section Trifoliae, despite its name, 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.
The timing of divergences within this section is disputed, with subsections Australes and Ponderosae having diverged during the mid Cretaceous according to one study, but not until the mid Oligocene in others.

Subsection ''Attenuatae''

The three closed-cone and fire adapted species of California and Baja California form a small subsection; closely related to, and often included within subsection Australes. Some studies suggest Pinus glabra may also belong here though others include it in subsection Australes.
This subsection is native to North and Central America and islands in the Caribbean. It has 26 living species.
The Contortae are native to North America and Mexico. It contains four accepted species.
This subsection 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 It contains at least 13 living species and may contain five more if the disputed species become widely accepted.
These are pines of the western United States and Mexico with four existing species. Within the subsection the Coulter pine is closely related with the Jeffery pine and the gray pine is likewise paired with the Torrey pine.
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.

Section ''Nelsonia''

This unique pine is remarkable in differing from all other pines in multiple cone and foliage characters. Genetically, its position has varied between and within studies; three nuclear genes and chloroplast DNA placed it as sister taxon to the rest of Section Parrya, while a late embryogenesis abundant -like gene resolved it as sister to Section Quinquefoliae; another study resolved it as sister to Subsection Balfourianae. It has probably evolved separately from the rest of the genus since the late Eocene, with its most recent common ancestor around 37 million years ago. These unique characters warrant recognition as a monotypic section.

Subsection ''Nelsonianae''

Subsection Nelsonianae is native to northeastern Mexico. It consists of the single species with persistent fascicle sheaths.
Section Parrya has one to five needles per fascicle. The seeds have articulate wings; in most, the wing is vestigial, and remains attached to the cone scale when the seed is released apparently wingless. In all species, 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 is native to southwest United States.
The 'big-cone' pinyons, with larger cones than subsection Cembroides.
Subsection Cembroides is native to Mexico and the southwestern United States.
Section Quinquefoliae, as its name 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.
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.
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.
Species which are not placed in a subgenus at this time.