Pinaceae
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The Pinaceae (), or pine family, are
conifer Conifers () are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a sin ...
trees or shrubs, including many of the well-known conifers of commercial importance such as cedars,
fir Firs are evergreen coniferous trees belonging to the genus ''Abies'' () in the family Pinaceae. There are approximately 48–65 extant species, found on mountains throughout much of North and Central America, Eurasia, and North Africa. The genu ...
s, hemlocks, piñons,
larch Larches are deciduous conifers in the genus ''Larix'', of the family Pinaceae (subfamily Laricoideae). Growing from tall, they are native to the cooler regions of the northern hemisphere, where they are found in lowland forests in the high la ...
es,
pine 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 ...
s and
spruce A spruce is a tree of the genus ''Picea'' ( ), a genus of about 40 species of coniferous evergreen trees in the family Pinaceae, found in the northern temperate and boreal ecosystem, boreal (taiga) regions of the Northern hemisphere. ''Picea'' ...
s. The family is included in the order
Pinales The order Pinales in the division Pinophyta, class Pinopsida, comprises all the extant conifers. The distinguishing characteristic is the reproductive structure known as a cone produced by all Pinales. All of the extant conifers, such as Arauca ...
, formerly known as Coniferales. Pinaceae have distinctive cones with woody scales bearing typically two ovules, and are supported as
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
by both morphological trait and genetic analysis. They are the largest extant conifer family in species diversity, with between 220 and 250 species (depending on taxonomic opinion) in 11 genera, and the second-largest (after
Cupressaceae Cupressaceae or the cypress family is a family of conifers. The family includes 27–30 genera (17 monotypic), which include the junipers and redwoods, with about 130–140 species in total. They are monoecious, subdioecious or (rarely) dioecio ...
) in geographical range, found in most of the
Northern Hemisphere The Northern Hemisphere is the half of Earth that is north of the equator. For other planets in the Solar System, north is defined by humans as being in the same celestial sphere, celestial hemisphere relative to the invariable plane of the Solar ...
, with the majority of the species in temperate climates, but ranging from subarctic to tropical. The family often forms the dominant component of boreal, coastal, and
montane forest Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures lapse rate, fall as elevation increases, causing the ecosystem to stratify. This stratification is ...
s. One species, '' Pinus merkusii'', grows just south of the
equator The equator is the circle of latitude that divides Earth into the Northern Hemisphere, Northern and Southern Hemisphere, Southern Hemispheres of Earth, hemispheres. It is an imaginary line located at 0 degrees latitude, about in circumferen ...
in Southeast Asia. Major centres of diversity are found in the mountains of
southwest China Southwestern China () is a region in the People's Republic of China. It consists of five provincial administrative regions, namely Chongqing, Sichuan, Guizhou, Yunnan, and Xizang. Geography Southwestern China is a rugged and mountainous region, ...
, Mexico, central Japan, and
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 ...
.


Description

Members of the family Pinaceae are
tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, e.g., including only woody plants with secondary growth, only ...
s (rarely
shrub A shrub or bush is a small to medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees by their multiple ...
s) growing from tall, mostly
evergreen In botany, an evergreen is a plant which has Leaf, foliage that remains green and functional throughout the year. This contrasts with deciduous plants, which lose their foliage completely during the winter or dry season. Consisting of many diffe ...
(except the
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 ...
'' Larix'' and '' Pseudolarix''),
resin A resin is a solid or highly viscous liquid that can be converted into a polymer. Resins may be biological or synthetic in origin, but are typically harvested from plants. Resins are mixtures of organic compounds, predominantly terpenes. Commo ...
ous,
monoecious Monoecy (; adj. monoecious ) is a sexual system in seed plants where separate male and female cones or flowers are present on the same plant. It is a monomorphic sexual system comparable with gynomonoecy, andromonoecy and trimonoecy, and contras ...
, with subopposite or whorled branches, and spirally arranged, linear (needle-like) leaves. The embryos of Pinaceae have three to 24
cotyledon A cotyledon ( ; ; "a cavity, small cup, any cup-shaped hollow", gen. (), ) is a "seed leaf" – a significant part of the embryo within the seed of a plant – and is formally defined as "the embryonic leaf in seed-bearing plants, one or mor ...
s. The female cones are large and usually woody, long, with numerous spirally arranged scales, and two winged
seed In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
s on each scale. The male cones are small, long, and fall soon after pollination; pollen dispersal is by wind. Seed dispersal is mostly by wind, but some species have large seeds with reduced wings, and are dispersed by birds. Analysis of Pinaceae cones reveals how selective pressure has shaped the evolution of variable cone size and function throughout the family. Variation in cone size in the family has likely resulted from the variation of seed dispersal mechanisms available in their environments over time. All Pinaceae with seeds weighing less than 90 milligrams are seemingly adapted for wind dispersal. Pines having seeds larger than 100 mg are more likely to have benefited from adaptations that promote animal dispersal, particularly by birds. Pinaceae that persist in areas where
tree squirrel Tree squirrels are the members of the squirrel Family (biology), family (Sciuridae) commonly just referred to as "squirrels". They include more than 100 arboreal species native to all continents except Antarctica and Oceania. They do not form a ...
s are abundant do not seem to have evolved adaptations for bird dispersal. Boreal conifers have many adaptions for winter. The narrow conical shape of northern conifers, and their downward-drooping limbs help them shed snow, and many of them seasonally alter their biochemistry to make them more resistant to freezing, called "hardening". File:Floral Morphology and Anatomy of Picea sp. (Spruce).jpg, Floral Morphology and Anatomy of ''Picea'' sp. (Spruce) File:Vagamon Pine Forest.jpg, Cultivated pine forest in Western Ghats, India


Classification

Classification of the subfamilies and genera of Pinaceae has been subject to debate in the past. Pinaceae ecology, morphology, and history have all been used as the basis for methods of analyses of the family. An 1891 publication divided the family into two subfamilies, using the number and position of resin canals in the primary vascular region of the young taproot as the primary consideration. In a 1910 publication, the family was divided into two tribes based on the occurrence and type of long–short shoot dimorphism. A more recent classification divided the subfamilies and genera based on the consideration of features of ovulate cone anatomy among extant and fossil members of the family. Below is an example of how the morphology has been used to classify Pinaceae. The 11 genera are grouped into four subfamilies, based on the microscopical anatomy and the morphology of the cones, pollen, wood, seeds, and leaves: * Subfamily Pinoideae (''
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 ...
''): cones are biennial, rarely triennial, with each year's scale-growth distinct, forming an umbo on each scale, the cone scale base is broad, concealing the seeds fully from
abaxial {{Short pages monitor