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Conifers are a group of cone-bearing
seed plants A spermatophyte (; ), also known as phanerogam (taxon Phanerogamae) or phaenogam (taxon Phaenogamae), is any plant that produces seeds, hence the alternative name seed plant. Spermatophytes are a subset of the embryophytes or land plants. They inc ...
, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant
class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used differently ...
, Pinopsida. All
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 ...
conifers are
perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...
woody plant A woody plant is a plant that produces wood as its structural tissue and thus has a hard stem. In cold climates, woody plants further survive winter or dry season above ground, as opposite to herbaceous plants that die back to the ground until ...
s with secondary growth. The great majority are trees, though a few are shrubs. Examples include cedars, Douglas-firs, cypresses, firs,
juniper Junipers are coniferous trees and shrubs in the genus ''Juniperus'' () of the cypress family Cupressaceae. Depending on the taxonomy, between 50 and 67 species of junipers are widely distributed throughout the Northern Hemisphere, from the Arcti ...
s, kauri,
larch Larches are deciduous conifers in the genus ''Larix'', of the family Pinaceae (subfamily Laricoideae). Growing from tall, they are native to much of the cooler temperate northern hemisphere, on lowlands in the north and high on mountains furt ...
es, pines, hemlocks,
redwoods Sequoioideae, popularly known as redwoods, is a subfamily of coniferous trees within the family Cupressaceae. It includes the largest and tallest trees in the world. Description The three redwood subfamily genera are '' Sequoia'' from coasta ...
, spruces, and yews.Campbell, Reece, "Phylum Coniferophyta". Biology. 7th. 2005. Print. P. 595 As of 1998, the division Pinophyta was estimated to contain eight families, 68 genera, and 629 living species. Although the total number of species is relatively small, conifers are
ecologically Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overlaps w ...
important. They are the dominant plants over large areas of land, most notably the taiga of the Northern Hemisphere, but also in similar cool climates in mountains further south. Boreal conifers have many wintertime adaptations. The narrow conical shape of northern conifers, and their downward-drooping limbs, help them shed snow. Many of them seasonally alter their biochemistry to make them more resistant to freezing. While
tropical rainforest Tropical rainforests are rainforests that occur in areas of tropical rainforest climate in which there is no dry season – all months have an average precipitation of at least 60 mm – and may also be referred to as ''lowland equatori ...
s have more
biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic (''genetic variability''), species (''species diversity''), and ecosystem (''ecosystem diversity'') le ...
and turnover, the immense conifer forests of the world represent the largest terrestrial
carbon sink A carbon sink is anything, natural or otherwise, that accumulates and stores some carbon-containing chemical compound for an indefinite period and thereby removes carbon dioxide () from the atmosphere. Globally, the two most important carbon ...
. Conifers are of great economic value for softwood lumber and paper production.


Evolutionary history

The earliest conifers appear in the fossil record during the Late
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carboniferous ...
( Pennsylvanian), over 300 million years ago. Conifers have been suggested to be most closely related to the
Cordaitales Cordaitales are an extinct order of gymnosperms, known from the early Carboniferous to the late Permian. Many Cordaitales had elongated strap-like leaves, resembling some modern-day conifers of the Araucariaceae and Podocarpaceae. They had cone ...
'','' a group of Carboniferous-Permian trees and clambering plants whose reproductive structures have some similarities to those of conifers. The most primitive conifers belong to the paraphyletic assemblage of " walchian conifers", which were small trees, and probably originated in dry upland habitats. The range of conifers expanded during the Early Permian (
Cisuralian The Cisuralian is the first series/epoch of the Permian. The Cisuralian was preceded by the Pennsylvanian and followed by the Guadalupian. The Cisuralian Epoch is named after the western slopes of the Ural Mountains in Russia and Kazakhstan and ...
) to lowlands due to increasing aridity. Walchian conifers were gradually replaced by more advanced voltzialean or "transition" conifers. Conifers were largely unaffected by the Permian–Triassic extinction event, and were dominant land plants of the Mesozoic era. Modern groups of conifers emerged from the Voltziales during the Late Permian through Jurassic. Conifers underwent a major decline in the Late Cretaceous corresponding to the explosive
adaptive radiation In evolutionary biology, adaptive radiation is a process in which organisms diversify rapidly from an ancestral species into a multitude of new forms, particularly when a change in the environment makes new resources available, alters biotic in ...
of flowering plants.


Taxonomy and naming

''Conifer'' is a Latin word, a compound of ''conus'' (cone) and ''ferre'' (to bear), meaning "the one that bears (a) cone(s)". The division name Pinophyta conforms to the rules of the ''
International Code of Nomenclature for algae, fungi, and plants The ''International Code of Nomenclature for algae, fungi, and plants'' (ICN) is the set of rules and recommendations dealing with the formal botanical names that are given to plants, fungi and a few other groups of organisms, all those "tra ...
(ICN)'', which state (Article 16.1) that the names of higher
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
in plants (above the rank of family) are either formed from the name of an included family (usually the most common and/or representative), in this case Pinaceae (the pine family), or are descriptive. A descriptive name in widespread use for the conifers (at whatever rank is chosen) is Coniferae (Art 16 Ex 2). According to the ''ICN'', it is possible to use a name formed by replacing the termination ''-aceae'' in the name of an included family, in this case preferably Pinaceae, by the appropriate termination, in the case of this division ''-ophyta''. Alternatively, "
descriptive botanical name Descriptive botanical names are scientific names of groups of plants that are irregular, not being derived systematically from the name of a type genus. They may describe some characteristics of the group in general or may be a name already in exis ...
s" may also be used at any
rank Rank is the relative position, value, worth, complexity, power, importance, authority, level, etc. of a person or object within a ranking, such as: Level or position in a hierarchical organization * Academic rank * Diplomatic rank * Hierarchy * H ...
above family. Both are allowed. This means that if conifers are considered a division, they may be called Pinophyta or Coniferae. As a class, they may be called Pinopsida or Coniferae. As an order they may be called Pinales or Coniferae or
Coniferales 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 cedar ...
. Conifers are the largest and economically most important component group of the gymnosperms, but nevertheless they comprise only one of the four groups. The division Pinophyta consists of just one class, Pinopsida, which includes both living and fossil taxa. Subdivision of the living conifers into two or more orders has been proposed from time to time. The most commonly seen in the past was a split into two orders,
Taxales The plant order Taxales was until recently treated as a distinct order in the division Pinophyta, class Pinopsida, and included only those species in the family Taxaceae, known commonly as yews. Under this interpretation, all other conifers were c ...
(Taxaceae only) and
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 cedar ...
(the rest), but recent research into DNA sequences suggests that this interpretation leaves the Pinales without Taxales as paraphyletic, and the latter order is no longer considered distinct. A more accurate subdivision would be to split the class into three orders, Pinales containing only Pinaceae, Araucariales containing Araucariaceae and Podocarpaceae, and Cupressales containing the remaining families (including Taxaceae), but there has not been any significant support for such a split, with the majority of opinion preferring retention of all the families within a single order Pinales, despite their antiquity and diverse
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
. , the conifers were accepted as composed of seven families, with a total of 65–70 genera and 600–630 species (696 accepted names). The seven most distinct families are linked in the box above right and phylogenetic diagram left. In other interpretations, the
Cephalotaxaceae Cephalotaxaceae is a small grouping of conifers, that included one to three genera closely allied to Taxaceae. However, members of Cephalotaxaceae are now included in Taxaceae by botanists, instead of as a distinct family, based on phylogenetic e ...
may be better included within the Taxaceae, and some authors additionally recognize
Phyllocladaceae ''Phyllocladus'', the celery pines, is a small genus of conifers, now usually treated in the family Podocarpaceae.James E. Eckenwalder. 2009. ''Conifers of the World''. Timber Press: Portland, OR, USA. .'' ''Species occur mainly in New Zealand, ...
as distinct from Podocarpaceae (in which it is included here). The family Taxodiaceae is here included in family Cupressaceae, but was widely recognized in the past and can still be found in many field guides. A new classification and linear sequence based on molecular data can be found in an article by Christenhusz et al.Christenhusz, M.J.M., Reveal, J., Farjon, A., Gardner, M.F., Mill, R.R. & Chase, M.W. (2011) A new classification and linear sequence of extant gymnosperms. Phytotaxa 19: 55–70. The conifers are an ancient group, with a fossil record extending back about 300 million years to the Paleozoic in the late
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carboniferous ...
period; even many of the modern genera are recognizable from fossils 60–120 million years old. Other classes and orders, now long extinct, also occur as fossils, particularly from the late Paleozoic and Mesozoic eras. Fossil conifers included many diverse forms, the most dramatically distinct from modern conifers being some herbaceous conifers with no woody stems. Major fossil orders of conifers or conifer-like plants include the
Cordaitales Cordaitales are an extinct order of gymnosperms, known from the early Carboniferous to the late Permian. Many Cordaitales had elongated strap-like leaves, resembling some modern-day conifers of the Araucariaceae and Podocarpaceae. They had cone ...
, Vojnovskyales, Voltziales and perhaps also the Czekanowskiales (possibly more closely related to the Ginkgophyta). Multiple studies also indicate that the Gnetophyta belong within the conifers despite their distinct appearances, either placing them as a sister group to
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 cedar ...
(the 'gnepine' hypothesis) or as being more derived than Pinales but sister to the rest of the group. Most recent studies favor the 'gnepine' hypothesis.


Morphology

All living conifers are woody plants, and most are trees, the majority having monopodial growth form (a single, straight trunk with side branches) with strong apical dominance. Many conifers have distinctly scented resin, secreted to protect the tree against insect infestation and fungal infection of wounds. Fossilized resin hardens into
amber Amber is fossilized tree resin that has been appreciated for its color and natural beauty since Neolithic times. Much valued from antiquity to the present as a gemstone, amber is made into a variety of decorative objects."Amber" (2004). In Ma ...
. The size of mature conifers varies from less than one metre, to over 100 metres. The world's tallest, thickest, largest, and oldest living trees are all conifers. The tallest is a
Coast Redwood ''Sequoia sempervirens'' ()''Sunset Western Garden Book,'' 1995:606–607 is the sole living species of the genus ''Sequoia (genus), Sequoia'' in the cypress family Cupressaceae (formerly treated in Taxodiaceae). Common names include coast r ...
(''Sequoia sempervirens''), with a height of 115.55 metres (although one Victorian mountain ash, '' Eucalyptus regnans'', allegedly grew to a height of 140 metres, although the exact dimensions were not confirmed). The thickest, meaning the tree with the greatest trunk diameter, is a
Montezuma Cypress ''Taxodium mucronatum'', commonly known as Montezuma bald cypress, Montezuma cypress, or ahuehuete, is a species of ''Taxodium'' that is primarily native to Mexico and Guatemala, with a few populations in the southwestern United States. Ahuehuet ...
(''Taxodium mucronatum''), 11.42 metres in diameter. The largest tree by three-dimensional volume is a Giant Sequoia ('' Sequoiadendron giganteum''), with a volume 1486.9 cubic metres. The smallest is the pygmy pine (''Lepidothamnus laxifolius'') of New Zealand, which is seldom taller than 30 cm when mature. The oldest is a Great Basin Bristlecone Pine (''
Pinus longaeva ''Pinus longaeva'' (commonly referred to as the Great Basin bristlecone pine, intermountain bristlecone pine, or western bristlecone pine) is a long-living species of bristlecone pine tree found in the higher mountains of California, Nevada, and ...
''), 4,700 years old.


Foliage

Since most conifers are evergreens, the
leaves A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...
of many conifers are long, thin and have a needle-like appearance, but others, including most of the
Cupressaceae Cupressaceae is a conifer family, the cypress family, with worldwide distribution. The family includes 27–30 genera (17 monotypic), which include the junipers and redwoods, with about 130–140 species in total. They are monoecious, subdioeci ...
and some of the
Podocarpaceae Podocarpaceae is a large family of mainly Southern Hemisphere conifers, known in English as podocarps, comprising about 156 species of evergreen trees and shrubs.James E. Eckenwalder. 2009. ''Conifers of the World''. Portland, Oregon: Timber P ...
, have flat, triangular scale-like leaves. Some, notably ''
Agathis ''Agathis'', commonly known as kauri or dammara, is a genus of 22 species of evergreen tree. The genus is part of the ancient conifer family Araucariaceae, a group once widespread during the Jurassic and Cretaceous periods, but now largely res ...
'' in Araucariaceae and ''
Nageia ''Nageia'' is a genus of conifers belonging to the podocarp family Podocarpaceae.Christopher N. Page. 1990. "Podocarpaceae" pages 332-346. In: Klaus Kubitzki (general editor); Karl U. Kramer and Peter S. Green (volume editors) ''The Families and ...
'' in Podocarpaceae, have broad, flat strap-shaped leaves. Others such as ''
Araucaria columnaris ''Araucaria columnaris'', the coral reef araucaria, Cook pine (or Cook's pine), New Caledonia pine, Cook araucaria, or columnar araucaria, is a species of conifer in the family Araucariaceae. Distribution The tree is endemic to New Caledonia in ...
'' have leaves that are awl-shaped. In the majority of conifers, the leaves are arranged spirally, exceptions being most of Cupressaceae and one genus in Podocarpaceae, where they are arranged in decussate opposite pairs or whorls of 3 (−4). In many species with spirally arranged leaves, such as ''
Abies grandis ''Abies grandis'' (grand fir, giant fir, lowland white fir, great silver fir, western white fir, Vancouver fir, or Oregon fir) is a fir native to the Pacific Northwest and Northern California of North America, occurring at altitudes of sea level ...
'' (pictured), the leaf bases are twisted to present the leaves in a very flat plane for maximum light capture. Leaf size varies from 2 mm in many scale-leaved species, up to 400 mm long in the needles of some pines (e.g. Apache Pine, ''
Pinus engelmannii ''Pinus engelmannii'', commonly known as the Apache pine, is a tree of Northern Mexico, in the Sierra Madre Occidental with its range extending a short distance into the United States in southwestern New Mexico and southeastern Arizona. This pin ...
''). The
stoma In botany, a stoma (from Greek ''στόμα'', "mouth", plural "stomata"), also called a stomate (plural "stomates"), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is bor ...
ta are in lines or patches on the leaves and can be closed when it is very dry or cold. The leaves are often dark green in colour, which may help absorb a maximum of energy from weak sunshine at high
latitude In geography, latitude is a coordinate that specifies the north–south position of a point on the surface of the Earth or another celestial body. Latitude is given as an angle that ranges from –90° at the south pole to 90° at the north pole ...
s or under forest canopy shade. Conifers from hotter areas with high sunlight levels (e.g. Turkish Pine ''
Pinus brutia ''Pinus brutia'', commonly known as the Turkish pine, is a species of pine native to the eastern Mediterranean region. The bulk of its range is in Turkey. Turkish pine is also known by several other common names: Calabrian pine (from a naturali ...
'') often have yellower-green leaves, while others (e.g.
blue spruce The blue spruce (''Picea pungens''), also commonly known as green spruce, Colorado spruce, or Colorado blue spruce, is a species of spruce tree. It is native to North America, and is found in USDA growing zones 1 through 7. It is found naturally ...
, ''Picea pungens'') may develop blue or silvery leaves to reflect ultraviolet light. In the great majority of genera the leaves are evergreen, usually remaining on the plant for several (2–40) years before falling, but five genera (''
Larix Larches are deciduous conifers in the genus ''Larix'', of the family Pinaceae (subfamily Laricoideae). Growing from tall, they are native to much of the cooler temperate northern hemisphere, on lowlands in the north and high on mountains fu ...
'', '' Pseudolarix'', ''
Glyptostrobus ''Glyptostrobus'' is a small genus of conifers in the family Cupressaceae (formerly in the family Taxodiaceae). The sole living species, '' Glyptostrobus pensilis'', is native to subtropical southeastern China, from Fujian west to southeast Yun ...
'', ''
Metasequoia ''Metasequoia'', or dawn redwoods, is a genus of fast-growing deciduous trees, one of three species of conifers known as redwoods. The living species ''Metasequoia glyptostroboides'' is native to Lichuan county in Hubei province, China. Althou ...
'' and '' Taxodium'') are deciduous, shedding their leaves in autumn. The seedlings of many conifers, including most of the Cupressaceae, and ''Pinus'' in Pinaceae, have a distinct juvenile foliage period where the leaves are different, often markedly so, from the typical adult leaves.


Tree ring structure

Tree rings are records of the
influence Influence or influencer may refer to: *Social influence, in social psychology, influence in interpersonal relationships ** Minority influence, when the minority affect the behavior or beliefs of the majority *Influencer marketing, through individ ...
of environmental conditions, their anatomical characteristics record growth rate changes produced by these changing conditions. The microscopic structure of conifer wood consists of two types of
cells Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery w ...
: parenchyma, which have an oval or polyhedral shape with approximately identical dimensions in three directions, and strongly elongated tracheids. Tracheids make up more than 90% of timber volume. The tracheids of earlywood formed at the beginning of a growing season have large radial sizes and smaller, thinner
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering me ...
s. Then, the first tracheids of the transition zone are formed, where the radial size of cells and thickness of their cell walls changes considerably. Finally, the latewood tracheids are formed, with small radial sizes and greater cell wall thickness. This is the basic
pattern A pattern is a regularity in the world, in human-made design, or in abstract ideas. As such, the elements of a pattern repeat in a predictable manner. A geometric pattern is a kind of pattern formed of geometric shapes and typically repeated l ...
of the internal
cell Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery w ...
structure of conifer tree rings.


Reproduction

Most conifers are monoecious, but some are subdioecious or dioecious; all are
wind-pollinated Anemophily or wind pollination is a form of pollination whereby pollen is distributed by wind. Almost all gymnosperms are anemophilous, as are many plants in the order Poales, including grasses, sedges, and rushes. Other common anemophilo ...
. Conifer seeds develop inside a protective cone called a strobilus. The cones take from four months to three years to reach maturity, and vary in size from 2 mm to 600 mm long. In Pinaceae,
Araucariaceae Araucariaceae – also known as araucarians – is an extremely ancient family of coniferous trees. The family achieved its maximum diversity during the Jurassic and Cretaceous periods and the early Cenozoic, when it was distributed almost world ...
,
Sciadopityaceae ''Sciadopitys verticillata'', the or Japanese umbrella-pine, is a unique conifer endemic to Japan. It is the sole member of the family Sciadopityaceae and genus ''Sciadopitys'', a living fossil with no close relatives. The oldest fossils of ...
and most
Cupressaceae Cupressaceae is a conifer family, the cypress family, with worldwide distribution. The family includes 27–30 genera (17 monotypic), which include the junipers and redwoods, with about 130–140 species in total. They are monoecious, subdioeci ...
, the cones are woody, and when mature the scales usually spread open allowing the seeds to fall out and be dispersed by the wind. In some (e.g. firs and cedars), the cones disintegrate to release the seeds, and in others (e.g. the pines that produce
pine nut Pine nuts, also called piñón (), pinoli (), pignoli or chilgoza (), are the edible seeds of pines (family Pinaceae, genus ''Pinus''). According to the Food and Agriculture Organization, only 29 species provide edible nuts, while 20 are trade ...
s) the nut-like seeds are dispersed by birds (mainly
nutcracker A nutcracker is a tool designed to open nuts by cracking their shells. There are many designs, including levers, screws, and ratchets. The lever version is also used for cracking lobster and crab shells. A decorative version portrays a person w ...
s, and
jay A jay is a member of a number of species of medium-sized, usually colorful and noisy, passerine birds in the Crow family, Corvidae. The evolutionary relationships between the jays and the magpies are rather complex. For example, the Eurasian ...
s), which break up the specially adapted softer cones. Ripe cones may remain on the plant for a varied amount of time before falling to the ground; in some fire-adapted pines, the seeds may be stored in closed cones for up to 60–80 years, being released only when a fire kills the parent tree. In the families
Podocarpaceae Podocarpaceae is a large family of mainly Southern Hemisphere conifers, known in English as podocarps, comprising about 156 species of evergreen trees and shrubs.James E. Eckenwalder. 2009. ''Conifers of the World''. Portland, Oregon: Timber P ...
,
Cephalotaxaceae Cephalotaxaceae is a small grouping of conifers, that included one to three genera closely allied to Taxaceae. However, members of Cephalotaxaceae are now included in Taxaceae by botanists, instead of as a distinct family, based on phylogenetic e ...
, Taxaceae, and one
Cupressaceae Cupressaceae is a conifer family, the cypress family, with worldwide distribution. The family includes 27–30 genera (17 monotypic), which include the junipers and redwoods, with about 130–140 species in total. They are monoecious, subdioeci ...
genus (''
Juniper Junipers are coniferous trees and shrubs in the genus ''Juniperus'' () of the cypress family Cupressaceae. Depending on the taxonomy, between 50 and 67 species of junipers are widely distributed throughout the Northern Hemisphere, from the Arcti ...
us''), the scales are soft, fleshy, sweet, and brightly colored, and are eaten by fruit-eating birds, which then pass the seeds in their droppings. These fleshy scales are (except in ''Juniperus'') known as
aril An aril (pronounced ), also called an arillus, is a specialized outgrowth from a seed that partly or completely covers the seed. An arillode or false aril is sometimes distinguished: whereas an aril grows from the attachment point of the see ...
s. In some of these conifers (e.g. most Podocarpaceae), the cone consists of several fused scales, while in others (e.g. Taxaceae), the cone is reduced to just one seed scale or (e.g. Cephalotaxaceae) the several scales of a cone develop into individual arils, giving the appearance of a cluster of berries. The male cones have structures called microsporangia that produce yellowish pollen through meiosis. Pollen is released and carried by the wind to female cones. Pollen grains from living pinophyte species produce pollen tubes, much like those of angiosperms. The gymnosperm male gametophytes (pollen grains) are carried by wind to a female cone and are drawn into a tiny opening on the ovule called the micropyle. It is within the ovule that pollen-germination occurs. From here, a pollen tube seeks out the female gametophyte, which contains archegonia each with an egg, and if successful, fertilization occurs. The resulting zygote develops into an embryo, which along with the female gametophyte (nutritional material for the growing embryo) and its surrounding integument, becomes a seed. Eventually, the seed may fall to the ground and, if conditions permit, grow into a new plant. In forestry, the terminology of flowering plants has commonly though inaccurately been applied to cone-bearing trees as well. The male cone and unfertilized female cone are called ''male flower'' and ''female flower'', respectively. After fertilization, the female cone is termed ''fruit'', which undergoes ''ripening'' (maturation). It was found recently that the pollen of conifers transfers the mitochondrial
organelle In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' the ...
s to the embryo, a sort of
meiotic Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately res ...
drive that perhaps explains why ''
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 ...
'' and other conifers are so productive, and perhaps also has bearing on observed sex-ratio bias. File:Abies lasiocarpa 6972.JPG, Pinaceae: unopened female cones of
subalpine fir ''Abies lasiocarpa'', the subalpine fir or Rocky Mountain fir, is a western North American fir tree. Description ''Abies lasiocarpa'' is a medium-sized evergreen conifer with a very narrow conic crown, growing to tall, exceptionally , with a t ...
(''Abies lasiocarpa'') Taxus baccata MHNT.jpg, Taxaceae: the fleshy aril that surrounds each seed in the
European Yew ''Taxus baccata'' is a species of evergreen tree in the family Taxaceae, native to western, central and southern Europe (including Britain and Ireland), northwest Africa, northern Iran, and southwest Asia.Rushforth, K. (1999). ''Trees of Britain ...
(''Taxus baccata'') is a highly modified seed cone scale Immature fir cone.jpg, Pinaceae: pollen cone of a
Japanese Larch ''Larix kaempferi'', the Japanese larch or karamatsu () in Japanese, is a species of larch native to Japan, in the mountains of Chūbu and Kantō regions in central Honshū.Farjon, A. (1990). ''Pinaceae. Drawings and Descriptions of the Genera' ...
(''Larix kaempferi'')


Life cycle

Conifers are heterosporous, generating two different types of spores: male microspores and female
megaspore Megaspores, also called macrospores, are a type of spore that is present in heterosporous plants. These plants have two spore types, megaspores and microspores. Generally speaking, the megaspore, or large spore, germinates into a female gametophy ...
s. These spores develop on separate male and female
sporophylls A sporophyll is a leaf that bears sporangia. Both microphylls and megaphylls can be sporophylls. In heterosporous plants, sporophylls (whether they are microphylls or megaphylls) bear either megasporangia and thus are called megasporophylls, or ...
on separate male and female cones. In the male cones, microspores are produced from microsporocytes by meiosis. The microspores develop into pollen grains, which are male gametophytes. Large amounts of pollen are released and carried by the wind. Some pollen grains will land on a female cone for pollination. The generative cell in the pollen grain divides into two haploid sperm cells by mitosis leading to the development of the pollen tube. At fertilization, one of the sperm cells unites its haploid nucleus with the haploid nucleus of an egg cell. The female cone develops two ovules, each of which contains haploid megaspores. A megasporocyte is divided by meiosis in each ovule. Each winged pollen grain is a four celled male gametophyte. Three of the four cells break down leaving only a single surviving cell which will develop into a female
multicellular A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially uni ...
gametophyte. The female gametophytes grow to produce two or more archegonia, each of which contains an egg. Upon fertilization, the diploid egg will give rise to the embryo, and a seed is produced. The female cone then opens, releasing the seeds which grow to a young seedling. # To fertilize the ovum, the male cone releases pollen that is carried on the wind to the female cone. This is pollination. (Male and female cones usually occur on the same plant.) # The pollen fertilizes the female gamete (located in the female cone). Fertilization in some species does not occur until 15 months after pollination. # A fertilized female gamete (called a zygote) develops into an embryo. # A seed develops which contains the embryo. The seed also contains the integument cells surrounding the embryo. This is an evolutionary characteristic of the Spermatophyta. # Mature seed drops out of cone onto the ground. # Seed germinates and seedling grows into a mature plant. # When the plant is mature, it produces cones and the cycle continues.


Female reproductive cycles

Conifer reproduction is synchronous with seasonal changes in temperate zones. Reproductive development slows to a halt during each winter season and then resumes each spring. The male strobilus development is completed in a single year. Conifers are classified by three reproductive cycles that refer to the completion of female strobilus development from initiation to seed maturation. All three types of reproductive cycle have a long gap between pollination and
fertilization Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Proce ...
. One year reproductive cycle:The genera include ''
Abies Firs (''Abies'') are a genus of 48–56 species of evergreen coniferous trees in the family Pinaceae. They are found on mountains throughout much of North and Central America, Europe, Asia, and North Africa. The genus is most closely re ...
'', ''
Picea A spruce is a tree of the genus ''Picea'' (), a genus of about 35 species of coniferous evergreen trees in the family Pinaceae, found in the northern temperate and boreal ( taiga) regions of the Earth. ''Picea'' is the sole genus in the subfam ...
'', ''
Cedrus ''Cedrus'', common English name cedar, is a genus of coniferous trees in the plant family Pinaceae (subfamily Abietoideae). They are native to the mountains of the western Himalayas and the Mediterranean region, occurring at altitudes of 1,500 ...
'', '' Pseudotsuga,'' '' Tsuga'', ''
Keteleeria ''Keteleeria'' is a genus of three species of coniferous trees in the family Pinaceae first described as a genus in 1866. The genus name ''Keteleeria'' honours J.B. Keteleer (1813–1903), a French nurseryman. The group is related to the genera ...
'' ''( Pinaceae)'' and ''
Cupressus ''Cupressus'' is one of several genera of evergreen conifers within the family Cupressaceae that have the common name cypress; for the others, see cypress. It is considered a polyphyletic group. Based on genetic and morphological analysis, the ...
, Thuja,
Cryptomeria ''Cryptomeria'' (literally "hidden parts") is a monotypic genus of conifer in the cypress family Cupressaceae, formerly belonging to the family Taxodiaceae. It includes only one species, ''Cryptomeria japonica'' ( syn. ''Cupressus japonica'' L ...
,
Cunninghamia ''Cunninghamia'' is a genus of one or two living species of evergreen coniferous trees in the cypress family Cupressaceae. They are native to China, northern Vietnam and Laos, and perhaps also Cambodia. They may reach in height. In vernacular u ...
'' and '' Sequoia (
Cupressaceae Cupressaceae is a conifer family, the cypress family, with worldwide distribution. The family includes 27–30 genera (17 monotypic), which include the junipers and redwoods, with about 130–140 species in total. They are monoecious, subdioeci ...
)''. Female strobili are initiated in late summer or fall in a year, then they overwinter. Female strobili emerge followed by pollination in the following spring. Fertilization takes place in summer of the following year, only 3–4 months after pollination. Cones mature and seeds are then shed by the end of that same year. Pollination and fertilization occur in a single growing season.Singh, H. 1978. Embryology of gymnosperms. Berlin, Gebruder Borntraeger. Two-year reproductive cycle:The genera includes '' Widdringtonia'', ''
Sequoiadendron ''Sequoiadendron'' is a genus of evergreen trees, with two species, only one of which survives to the present: * ''Sequoiadendron giganteum'', extant, commonly known as wellingtonia, giant redwood and giant sequoia, growing naturally in the Sierr ...
'' (''
Cupressaceae Cupressaceae is a conifer family, the cypress family, with worldwide distribution. The family includes 27–30 genera (17 monotypic), which include the junipers and redwoods, with about 130–140 species in total. They are monoecious, subdioeci ...
'') and most species of ''Pinus''. Female strobilus initials are formed in late summer or fall then overwinter. Female strobili emerge and receive pollen in the first year spring and become conelets. The conelet goes through another winter rest and, in the spring of the 2nd year archegonia form in the conelet. Fertilization of the archegonia occurs by early summer of the 2nd year, so the pollination-fertilization interval exceeds a year. After fertilization, the conelet is considered an immature cone. Maturation occurs by autumn of the 2nd year, at which time seeds are shed. In summary, the 1-year and the 2-year cycles differ mainly in the duration of the pollination- fertilization interval. Three-year reproductive cycle: Three of the conifer species are pine species ('' Pinus pinea'', ''
Pinus leiophylla ''Pinus leiophylla'', commonly known as Chihuahua pine, smooth-leaf pine,''Pinus leiophylla'' was originally described and published in ''Linnaea'' 6:354. 1831. and yellow pine (in Mexico, tlacocote and ocote chino), is a tree with a range pri ...
'', ''
Pinus torreyana The Torrey pine (''Pinus torreyana'') is a rare pine species in California, United States. It is a critically endangered species growing only in coastal San Diego County, and on Santa Rosa Island, offshore from Santa Barbara in Santa Barbara ...
'') which have pollination and fertilization events separated by a 2-year interval. Female strobili initiated during late summer or autumn in a year, then overwinter until the following spring. Female strobili emerge then pollination occurs in spring of the 2nd year then the pollinated strobili become conelets in the same year (i.e. the second year). The female
gametophytes A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the sex ...
in the conelet develop so slowly that the
megaspore Megaspores, also called macrospores, are a type of spore that is present in heterosporous plants. These plants have two spore types, megaspores and microspores. Generally speaking, the megaspore, or large spore, germinates into a female gametophy ...
does not go through free-nuclear divisions until autumn of the 3rd year. The conelet then overwinters again in the free-nuclear female gametophyte stage. Fertilization takes place by early summer of the 4th year and seeds mature in the cones by autumn of the 4th year.


Tree development

The growth and form of a forest tree are the result of activity in the primary and secondary meristems, influenced by the distribution of photosynthate from its needles and the hormonal gradients controlled by the apical meristems (Fraser et al. 1964).Fraser, D.A.; Belanger, L.; McGuire, D.; Zdrazil, Z. 1964. Total growth of the aerial parts of a white spruce tree at Chalk River, Ontario, Canada. Can. J. Bot. 42:159–179. External factors also influence growth and form. Fraser recorded the development of a single white spruce tree from 1926 to 1961. Apical growth of the stem was slow from 1926 through 1936 when the tree was competing with herbs and shrubs and probably shaded by larger trees. Lateral branches began to show reduced growth and some were no longer in evidence on the 36-year-old tree. Apical growth totaling about 340 m, 370 m, 420 m, 450 m, 500 m, 600 m, and 600 m was made by the tree in the years 1955 through 1961, respectively. The total number of needles of all ages present on the 36-year-old tree in 1961 was 5.25 million weighing 14.25 kg. In 1961, needles as old as 13 years remained on the tree. The ash weight of needles increased progressively with age from about 4% in first-year needles in 1961 to about 8% in needles 10 years old. In discussing the data obtained from the one 11 m tall white spruce, Fraser et al. (1964) speculated that if the photosynthate used in making apical growth in 1961 was manufactured the previous year, then the 4 million needles that were produced up to 1960 manufactured food for about 600,000 mm of apical growth or 730 g dry weight, over 12 million mm3 of wood for the 1961 annual ring, plus 1 million new needles, in addition to new tissue in branches, bark, and roots in 1960. Added to this would be the photosynthate to produce energy to sustain respiration over this period, an amount estimated to be about 10% of the total annual photosynthate production of a young healthy tree. On this basis, one needle produced food for about 0.19 mg dry weight of apical growth, 3 mm3 wood, one-quarter of a new needle, plus an unknown amount of branch wood, bark and roots. The order of priority of photosynthate distribution is probably: first to apical growth and new needle formation, then to buds for the next year's growth, with the cambium in the older parts of the branches receiving sustenance last. In the white spruce studied by Fraser et al. (1964), the needles constituted 17.5% of the over-day weight. Undoubtedly, the proportions change with time.


Seed-dispersal mechanism

Wind and animal dispersals are two major mechanisms involved in the dispersal of conifer seeds. Wind born seed dispersal involves two processes, namely; local neighborhood dispersal (LND) and long-distance dispersal (LDD). Long-distance dispersal distances range from from the source. Birds of the crow family,
Corvidae Corvidae is a cosmopolitan family of oscine passerine birds that contains the crows, ravens, rooks, jackdaws, jays, magpies, treepies, choughs, and nutcrackers. In colloquial English, they are known as the crow family or corvids. Currentl ...
, are the primary distributor of the conifer seeds. These birds are known to
cache Cache, caching, or caché may refer to: Places United States * Cache, Idaho, an unincorporated community * Cache, Illinois, an unincorporated community * Cache, Oklahoma, a city in Comanche County * Cache, Utah, Cache County, Utah * Cache Count ...
32,000 pine seeds and transport the seeds as far as from the source. The birds store the seeds in the soil at depths of under conditions which favor
germination Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, fer ...
.


Invasive species

A number of conifers originally introduced for forestry have become invasive species in parts of New Zealand, including radiata pine ('' Pinus radiata''), lodgepole pine ('' P. contorta''), Douglas fir (''Pseudotsuga mensiezii'') and European larch ('' Larix decidua''). In parts of South Africa, maritime pine (''
Pinus pinaster ''Pinus pinaster'', the maritime pine or cluster pine, is a pine native to the south Atlantic Europe region and parts of the western Mediterranean. It is a hard, fast growing pine bearing small seeds with large wings. Description ''Pinus pinast ...
''), patula pine ('' P. patula'') and radiata pine have been declared invasive species. These
wilding conifers Wilding conifers, also known as wilding pines, are invasive trees in the high country of New Zealand. Millions of dollars are spent on controlling their spread. In the South Island they threaten 210,000 hectares of public land administered by ...
are a serious environmental issue causing problems for pastoral farming and for conservation. Radiata pine was introduced to Australia in the 1870s. It is "the dominant tree species in the Australian plantation estate""Fauna conservation in Australian plantation forests: a review"
, May 2007, D.B. Lindenmayer and R.J. Hobbs
– so much so that many Australians are concerned by the resulting loss of native wildlife habitat. The species is widely regarded as an environmental weed across southeastern and southwestern Australia and the removal of individual plants beyond plantations is encouraged.


Predators

At least 20 species of roundheaded borers of the family
Cerambycidae The longhorn beetles (Cerambycidae), also known as long-horned or longicorns, are a large family of beetles, with over 35,000 species described. Most species are characterized by extremely long antennae, which are often as long as or longer than ...
feed on the wood of spruce, fir, and hemlock (Rose and Lindquist 1985).Rose, A.H.; Lindquist, O.H. 1985. Insects of eastern spruces, fir and, hemlock, revised edition. Gov’t Can., Can. For. Serv., Ottawa, For. Tech. Rep. 23. 159 p. (cited in Coates et al. 1994, cited orig ed 1977) Borers rarely bore tunnels in living trees, although when populations are high, adult beetles feed on tender twig bark, and may damage young living trees. One of the most common and widely distributed borer species in North America is the whitespotted sawyer (''Monochamus scutellatus''). Adults are found in summer on newly fallen or recently felled trees chewing tiny slits in the bark in which they lay eggs. The eggs hatch in about 2 weeks and the tiny larvae tunnel to the wood and score its surface with their feeding channels. With the onset of cooler weather, they bore into the wood making oval entrance holes and tunnel deeply. Feeding continues the following summer when larvae occasionally return to the surface of the wood and extend the feeding channels generally in a U-shaped configuration. During this time, small piles of frass extruded by the larvae accumulate under logs. Early in the spring of the second year following egg-laying, the larvae, about 30 mm long, pupate in the tunnel enlargement just below the wood surface. The resulting adults chew their way out in early summer, leaving round exit holes, so completing the usual 2-year life cycle.


Cultivation

Conifers – notably ''
Abies Firs (''Abies'') are a genus of 48–56 species of evergreen coniferous trees in the family Pinaceae. They are found on mountains throughout much of North and Central America, Europe, Asia, and North Africa. The genus is most closely re ...
'' (fir), ''
Cedrus ''Cedrus'', common English name cedar, is a genus of coniferous trees in the plant family Pinaceae (subfamily Abietoideae). They are native to the mountains of the western Himalayas and the Mediterranean region, occurring at altitudes of 1,500 ...
'', ''
Chamaecyparis lawsoniana ''Chamaecyparis lawsoniana'', known as Port Orford cedar or Lawson cypress, is a species of conifer in the genus '' Chamaecyparis'', family Cupressaceae. It is native to Oregon and northwestern California, and grows from sea level up to in the ...
'' (Lawson's cypress), ''
Cupressus ''Cupressus'' is one of several genera of evergreen conifers within the family Cupressaceae that have the common name cypress; for the others, see cypress. It is considered a polyphyletic group. Based on genetic and morphological analysis, the ...
'' (cypress),
juniper Junipers are coniferous trees and shrubs in the genus ''Juniperus'' () of the cypress family Cupressaceae. Depending on the taxonomy, between 50 and 67 species of junipers are widely distributed throughout the Northern Hemisphere, from the Arcti ...
, ''
Picea A spruce is a tree of the genus ''Picea'' (), a genus of about 35 species of coniferous evergreen trees in the family Pinaceae, found in the northern temperate and boreal ( taiga) regions of the Earth. ''Picea'' is the sole genus in the subfam ...
'' (spruce), ''
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 ...
'' (pine), '' Taxus'' (yew), '' Thuja'' (cedar) – have been the subject of selection for ornamental purposes (for more information see the silviculture page). Plants with unusual growth habits, sizes, and colours are propagated and planted in parks and gardens throughout the world.


Conditions for growth

Conifers can absorb nitrogen in either the ammonium (NH4+) or
nitrate Nitrate is a polyatomic ion with the chemical formula . Salts containing this ion are called nitrates. Nitrates are common components of fertilizers and explosives. Almost all inorganic nitrates are soluble in water. An example of an insolubl ...
(NO3) form, but the forms are not physiologically equivalent. Form of nitrogen affected both the total amount and relative composition of the soluble nitrogen in white spruce tissues (Durzan and Steward 1967).Durzan, D.J.; Steward, F.C. 1967. The nitrogen metabolism of ''Picea glauca'' (Moench) Voss and ''Pinus banksiana'' Lamb. as influenced by mineral nutrition. Can. J. Bot. 45:695–710. Ammonium nitrogen was shown to foster
arginine Arginine is the amino acid with the formula (H2N)(HN)CN(H)(CH2)3CH(NH2)CO2H. The molecule features a guanidino group appended to a standard amino acid framework. At physiological pH, the carboxylic acid is deprotonated (−CO2−) and both the a ...
and
amide In organic chemistry, an amide, also known as an organic amide or a carboxamide, is a compound with the general formula , where R, R', and R″ represent organic groups or hydrogen atoms. The amide group is called a peptide bond when it is ...
s and lead to a large increase of free
guanidine Guanidine is the compound with the formula HNC(NH2)2. It is a colourless solid that dissolves in polar solvents. It is a strong base that is used in the production of plastics and explosives. It is found in urine predominantly in patients experie ...
compounds, whereas in leaves nourished by nitrate as the sole source of nitrogen guanidine compounds were less prominent. Durzan and Steward noted that their results, drawn from determinations made in late summer, did not rule out the occurrence of different interim responses at other times of the year. Ammonium nitrogen produced significantly heavier (dry weight) seedlings with higher nitrogen content after 5 weeks (McFee and Stone 1968)McFee, W.W.; Stone, E.L. 1968. Ammonium and nitrate as nitrogen sources for ''Pinus radiata ''and ''Picea glauca''. Soil Sci. Soc. Amer. Proc. 32(6):879–884. than did the same amount of nitrate nitrogen. Swan (1960)Swan, H.S.D. 1960. The mineral nutrition of Canadian pulpwood species. 1. The influence of nitrogen, phosphorus, potassium, and magnesium deficiencies on the growth and development of white spruce, black spruce, jack pine, and western hemlock seedlings grown in a controlled environment. Pulp Paper Res. Instit. Can., Montreal QC, Woodlands Res. Index No. 116, Tech. Rep. 168. 66 p. found the same effect in 105-day-old white spruce. The general short-term effect of nitrogen fertilization on coniferous seedlings is to stimulate shoot growth more so than root growth (Armson and Carman 1961).Armson, K.A.; Carman, R.D. 1961. Forest tree nursery soil management. Ont. Dep. Lands & Forests, Timber Branch, Ottawa ON. 74 p. Over a longer period, root growth is also stimulated. Many nursery managers were long reluctant to apply nitrogenous fertilizers late in the growing season, for fear of increased danger of frost damage to succulent tissues. A presentation at the North American Forest Tree Nursery Soils Workshop at Syracuse in 1980 provided strong contrary evidence: Bob Eastman, President of the Western Maine Forest Nursery Co. stated that for 15 years he has been successful in avoiding winter “burn” to
Norway spruce ''Picea abies'', the Norway spruce or European spruce, is a species of spruce native to Northern, Central and Eastern Europe. It has branchlets that typically hang downwards, and the largest cones of any spruce, 9–17 cm long. It is very clo ...
and white spruce in his nursery operation by fertilizing with 50–80 lb/ac (56–90 kg/ha) nitrogen in September, whereas previously winter burn had been experienced annually, often severely. Eastman also stated that the overwintering storage capacity of stock thus treated was much improved (Eastman 1980).Eastman, B. 1980. The Western Maine Forest Nursery Company. pp. 291–295 In Proc. of the North American Forest Tree Nursery Soils Workshop, July 28 – August 1, 1980, Syracuse, New York. Environment Canada, Canadian Forestry Service, USDA For. Serv. The concentrations of nutrients in plant tissues depend on many factors, including growing conditions. Interpretation of concentrations determined by analysis is easy only when a nutrient occurs in excessively low or occasionally excessively high concentration. Values are influenced by environmental factors and interactions among the 16 nutrient elements known to be essential to plants, 13 of which are obtained from the soil, including nitrogen, phosphorus, potassium,
calcium Calcium is a chemical element with the symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar ...
, magnesium, and sulfur, all used in relatively large amounts (Buckman and Brady 1969).Buckman, H.O.; Brady, N.C. 1969. The Nature and Properties of Soils, 7th ed. Macmillan NY. 653 p. Nutrient concentrations in conifers also vary with season, age, and kind of tissue sampled, and analytical technique. The ranges of concentrations occurring in well-grown plants provide a useful guide by which to assess the adequacy of particular nutrients, and the ratios among the major nutrients are helpful guides to nutritional imbalances.


Economic importance

The softwood derived from conifers is of great economic value, providing about 45% of the world's annual lumber production. Other uses of the timber include the production of paper and plastic from chemically treated wood pulp. Some conifers also provide foods such as pine nuts and
Juniper berries A juniper berry is the female seed cone produced by the various species of junipers. It is not a true berry, but a cone with unusually fleshy and merged scales, which gives it a berry-like appearance. The cones from a handful of species, especial ...
, the latter used to flavor gin.


References


Bibliography

*


External links

* *
tolweb.org
Conifers


World list of conifer species from Conifer Database by A. Farjon in the Catalogue of Life

Tree browser for conifer families and genera via the Catalogue of Life

Royal Horticultural Society Encyclopedia of Conifers: A Comprehensive Guide to Cultivars and Species

DendroPress: Conifers Around the World
* {{Authority control Plant divisions Extant Pennsylvanian first appearances