Pinus – pines (c. 115)
Picea – spruces (c. 35)
Cathaya (1 species)
Larix – larches (about 14)
Pseudotsuga – Douglas-firs (5)
Abies – firs (about 50)
Cedrus – cedars (2–4)
Pseudolarix – golden larch (1)
Tsuga – hemlock (9)
Pinaceae (pine family) are trees or shrubs, including many of the
well-known conifers of commercial importance such as cedars, firs,
hemlocks, larches, pines and spruces. The family is included in the
order Pinales, formerly known as Coniferales.
Pinaceae are supported
as monophyletic by their protein-type sieve cell plastids, pattern of
proembryogeny, and lack of bioflavonoids. 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) in geographical range, found in
most of the Northern Hemisphere, 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
forests. One species,
Pinus merkusii, grows just south of the equator
in Southeast Asia. Major centres of diversity are found in the
mountains of southwest China, Mexico, central Japan, and California.
4 Further reading
5 External links
Pine forest in Vagamon, southern Western Ghats, Kerala (India)
Members of the family
Pinaceae are trees (rarely shrubs) growing from
2 to 100 m (7 to 300 ft) tall, mostly evergreen (except the
Larix and Pseudolarix), resinous, monoecious, with
subopposite or whorled branches, and spirally arranged, linear
(needle-like) leaves. The embryos of
Pinaceae have three to 24
The female cones are large and usually woody, 2–60 cm
(1–24 in) long, with numerous spirally arranged scales, and two
winged seeds on each scale. The male cones are small,
0.5–6.0 cm (0.2–2 in) 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 mg 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 squirrels 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".
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 subfamiles,
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:
Pinoideae (Pinus): 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
view, the seed is without resin vesicles, the seed wing holds the seed
in a pair of claws, leaves have primary stomatal bands adaxial (above
the xylem) or equally on both surfaces.
Piceoideae (Picea): cones are annual, without a distinct
umbo, the cone scale base is broad, concealing the seeds fully from
abaxial view, seed is without resin vesicles, blackish, the seed wing
holds the seed loosely in a cup, leaves have primary stomatal bands
adaxial (above the xylem) or equally on both surfaces.
Laricoideae (Larix, Pseudotsuga, and Cathaya): cones are
annual, without a distinct umbo, the cone scale base is broad,
concealing the seeds fully from abaxial view, the seed is without
resin vesicles, whitish, the seed wing holds the seed tightly in a
cup, leaves have primary stomatal bands abaxial (below the phloem
Subfamily Abietoideae (Abies, Cedrus, Pseudolarix, Keteleeria,
Nothotsuga, and Tsuga): cones are annual, without a distinct umbo, the
cone scale base is narrow, with the seeds partly visible in abaxial
view, the seed has resin vesicles, the seed wing holds the seed
tightly in a cup, leaves have primary stomatal bands abaxial (below
the phloem vessels) only.
^ a b Aljos Farjon (1998). World Checklist and Bibliography of
Conifers. Royal Botanic Gardens, Kew. ISBN 1-900347-54-7.
^ Christopher J. Earle (ed.). "
Pinus merkusii Junghuhn et de Vriese ex
de Vriese 1845". The Gymnosperm Database. Retrieved March 17,
^ Craig W. Benkman (1995). "Wind dispersal capacity of pine seeds and
the evolution of different seed dispersal modes in pines" (PDF).
Oikos. 73 (2): 221–224. doi:10.2307/3545911.
^ Robert A. Price, Jeanine Olsen-Stojkovich & Jerold M. Lowenstein
(1987). "Relationships among the genera of Pinaceae: an immunological
comparison". Systematic Botany. 12 (1): 91–97. doi:10.2307/2419217.
Heinz-Dietmar Behnke (1974). "Sieve element plastids of Gymnospermae:
their ultrastructure and relation to systematics".
and Evolution. 123 (1): 1–12. doi:10.1007/BF00983281.
D. F. Greene & E. A. Johnson (1990). "The dispersal of winged
fruits and seeds differing in autorotative behavior". Canadian Journal
of Botany. 68 (12): 2693–2697. doi:10.1139/b90-340.
A. Liston, D. S. Gernandt, T. F. Vining, C. S. Campbell & D.
Piñero (2003). R. R. Mill, ed. Molecular phylogeny of Pinaceae
and Pinus. Proceedings of the Fourth International
Acta Horticulturae. 615. Brugge: International Society for
Horticultural Science. pp. 107–114. CS1 maint: Multiple
names: authors list (link)
Zsolt Debreczy; Istvan Racz (2012). Conifers Around the World.
DendroPress. ISBN 9632190610.
Arboretum de Villardebelle French Arboretum of conifers around the
Gymnosperm Database – Pinaceae
Pinaceae on the web page of the Tree-of-Life project
Pine Trees From Around the World by The Spruce
Pinaceae from the Jepson Manual, covers Californian species and
much of western North America
Pinaceae in Flora of North America
Pinus in USDA Plants Database