Conifer: Difference between revisions

Latest revision as of 21:26, 29 October 2025

Template:Short description Template:Good article Script error: No such module "other uses". Template:Use dmy dates Template:Use British English Template:Automatic taxobox

Conifers (Template:IPAc-en) are a group of seed plants, a subset of gymnosperms. They are mainly evergreen trees with a regular branching pattern, reproducing with male and female cones, usually on the same tree. They are wind-pollinated and the seeds are usually dispersed by the wind. Scientifically, they make up the division Pinophyta, also known as Coniferae. All extant conifers except for the Gnetophytes are perennial woody plants with secondary growth. There are over 600 living species.

Conifers first appear in the fossil record over 300 million years ago in the Carboniferous. They became dominant land plants in the Mesozoic, until flowering plants took over many ecosystems in the Cretaceous. Many conifers today are relict species, surviving in a small part of their former ranges. Such relicts include Wollemia, known only from a small area of Australia, and Metasequoia glyptostroboides, known from Cretaceous fossils and surviving in a small area of China.

Although the total number of species is relatively small, conifers are ecologically important. They are the dominant plants over the taiga of the Northern Hemisphere. Boreal conifers have multiple adaptations to survive winters, including a conical shape to shed snow, strong tracheid vessels to tolerate ice pressure, and a waxy covering on the needle leaves to minimise water loss. Several fungi form ectomycorrhizal associations with conifers. Other fungi cause diseases such as needle cast, especially harmful to young trees. Conifers are affected by pest insects such as wood-boring longhorn beetles and by bark beetles, which make galleries just under the bark. Conifers are of great economic value for timber and paper production.

Evolution

Fossil history

The earliest conifers appear in the fossil record during the Late Carboniferous (Pennsylvanian) over 300 million years ago. Conifers are thought to be most closely related to the Cordaitales, a group of extinct Carboniferous-Permian trees and clambering plants whose reproductive structures had 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) to lowlands due to increasing aridity. Walchian conifers were gradually replaced by more advanced voltzialean or "transition" conifers.^[1] Conifers were largely unaffected by the Permian–Triassic extinction event,^[2] and were dominant land plants of the Mesozoic era. Modern groups of conifers emerged from the Voltziales during the Late Permian through Jurassic.^[3] Conifers underwent a major decline in the Late Cretaceous corresponding to the explosive adaptive radiation of flowering plants.^[4]

Voltziales: Walchia laxifolia foliage, Cisuralian, Germany

Voltziales: Walchia laxifolia foliage, Cisuralian, Germany
Araucaria cone, Jurassic, Argentina

Araucaria cone, Jurassic, Argentina
Elatides foliage, Late Cretaceous, N. America

Elatides foliage, Late Cretaceous, N. America
Base of conifer trunk with roots, Early Miocene, Lesbos, Greece

Base of conifer trunk with roots, Early Miocene, Lesbos, Greece

Relict species

Several extant conifers have relict taxon status, surviving in small areas or in very small numbers where they once may have been common and widespread. One such is Wollemia nobilis, discovered in 1994 in some narrow, steep-sided, sandstone gorges in Australia.^[5] The wild population consisted of under 60 adult trees with essentially no genetic variability, implying a genetic bottleneck some thousands of years ago.^[6] The extant gnetophytes consist of three relict genera, namely Ephedra, Gnetum, and Welwitschia. Fossils definitely of the group date back to the Late Jurassic, with many species in the Cretaceous.^[7] Conifers as a whole, too, declined markedly after the angiosperms (flowering plants) diversified during the Cretaceous, coming to dominate most terrestrial ecosystems. Many conifer species became extinct, leaving 30 out of 80 genera with just one extant species, and 11 more with just two or three species. The popular phrase "living fossils" could, the Dutch botanist Aljos Farjon states, fittingly be applied to many of these. Thus, Metasequoia glyptostroboides, the dawn redwood, is known from fossils of Late Cretaceous and Miocene age, and was found also as an extant tree with a small relict range in China.^[8]

Wollemia nobilis is a relict taxon known only from a small area in Australia.

Wollemia nobilis is a relict taxon known only from a small area in Australia.
Welwitschia mirabilis is one of the gnetophytes, all relict taxa very unlike other conifers.

Welwitschia mirabilis is one of the gnetophytes, all relict taxa very unlike other conifers.
Metasequoia glyptostroboides survives in a small part of China, and is known from fossils from the Late Cretaceous onwards.

Metasequoia glyptostroboides survives in a small part of China, and is known from fossils from the Late Cretaceous onwards.

External phylogeny

The cladogram summarizes the group's external phylogeny. The conifers are gymnosperms, sister to a clade consisting of the ginkgos and cycads.^[9]^[10]^[11]^[12]

Template:Clade

Internal phylogeny

The Gnetophyta, despite their distinct appearances, were long viewed as outside the conifer group, but phylogenomic analysis indicates that they are part of the conifer clade, sister to the pine family (the 'gnepine' hypothesis). If so, the gnetophytes once shared the distinctive characters of the conifers, and have lost them.^[13] The cladogram summarizes the conifers' internal phylogeny:^[14]

Template:Clade

Taxonomy

The name conifer, meaning 'cone-bearing', derives from Latin Script error: No such module "Lang". (cone) and Script error: No such module "Lang". (to bear).^[15] As recently as 1999, the botanist Aljos Farjon wrote that while the Coniferae had up to the early 20th century been considered "a natural family",^[8] comparable to the Rosaceae, he doubted that the conifers or the gymnosperms formed natural groups (clades).^[8] By 2016, the conifers were recognised as a clade, with six families (not including the gnetophytes),^[16] 65–70 genera, and over 600 living species (Template:Circa).^[17]Template:Rp^[18]^[19] Depending on interpretation, the Cephalotaxaceae may or may not be included within the Taxaceae, while some authors recognize Phyllocladaceae as distinct from Podocarpaceae. The family Taxodiaceae is here included in the family Cupressaceae.^[20]

Description

All living conifers (except the gnetophytes) are woody plants, and most are trees with narrow leaves, often needle-like. There are separate male and female reproductive structures, the cones. Pollination is always by wind; the seeds are mostly winged. The trees have a regular branching pattern. Many conifers have distinctly scented resin.^[21] The world's tallest and oldest living trees are conifers. The tallest is a coast redwood (Sequoia sempervirens), with a height of Template:Convert.^[22] Among the smallest conifers is the pygmy pine (Lepidothamnus laxifolius) of New Zealand, which is seldom taller than 30 cm when mature.^[23] The oldest non-clonal living tree is a Great Basin bristlecone pine (Pinus longaeva), 4,700 years old.^[24] Boreal conifers have multiple adaptations to survive winters, including the tree's conical shape to shed snow, strong tracheid vessels to tolerate ice pressure, and a waxy covering on the needle leaves to minimise water loss.^[25]

Tallest: Sequoia sempervirens can reach a height of Template:Convert.[22]

Tallest: Sequoia sempervirens can reach a height of Template:Convert.^[22]
Oldest: Pinus longaeva can reach an age of 4,700 years.[24]

Oldest: Pinus longaeva can reach an age of 4,700 years.^[24]
The narrow conical shape of boreal conifers, and their downward-drooping limbs, help them shed snow.[25]

The narrow conical shape of boreal conifers, and their downward-drooping limbs, help them shed snow.^[25]

Foliage

Most conifers are evergreens.^[26] In many species such as pines, firs, and cedars, the leaves are long, thin and needle-like. Others like cypresses have flat, triangular scale-like leaves.^[27] In the majority of conifers, the leaves are arranged spirally, the exceptions being most of Cupressaceae and one genus in Podocarpaceae, where they are arranged in decussate opposite pairs or whorls of 3 or 4. In many species with spirally arranged leaves, such as Abies grandis, 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). The stomata 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 latitudes or under forest canopy shade. Conifers from lower latitudes with high sunlight levels (e.g. Turkish pine Pinus brutia) often have yellower-green leaves, while others (e.g. blue spruce, Picea pungens) may develop blue or silvery leaves to reflect ultraviolet light. In the great majority of genera the leaves remain on the plant for several (2–40) years before falling, but five genera (Larix, Pseudolarix, Glyptostrobus, Metasequoia and Taxodium) are deciduous, shedding their leaves in autumn.^[26] The seedlings of some conifers, including pines, have a distinct juvenile foliage period where the leaves are different from the typical adult leaves.^[28]

Pinaceae: needle-like leaves of Scots pine (Pinus sylvestris)

Pinaceae: needle-like leaves of Scots pine (Pinus sylvestris)
Araucariaceae: awl-like leaves of Cook pine (Araucaria columnaris)

Araucariaceae: awl-like leaves of Cook pine (Araucaria columnaris)
In Abies grandis and many other species with spirally arranged leaves, each leaf is twisted near its base to maximize light capture.

In Abies grandis and many other species with spirally arranged leaves, each leaf is twisted near its base to maximize light capture.
Cupressaceae: scale leaves of Lawson's cypress (Chamaecyparis lawsoniana); scale in mm

Cupressaceae: scale leaves of Lawson's cypress (Chamaecyparis lawsoniana); scale in mm

Wood

Conifer wood consists of two types of cells: parenchyma, which have an oval or polyhedral shape, 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 walls. Then, the first tracheids of the transition zone are formed, where the radial size of cells and the thickness of their cell walls changes considerably. Finally, latewood tracheids are formed, with small radial sizes and greater cell wall thickness. This is the basic pattern of the internal cell structure of conifer tree rings.^[29]

Vertical (tangential) section of Abies concolor wood (xylem), showing tracheids as long overlapping tubes. Perforation pits (small circles) allow water to move from one tracheid to the next.

Vertical (tangential) section of Abies concolor wood (xylem), showing tracheids as long overlapping tubes. Perforation pits (small circles) allow water to move from one tracheid to the next.
Transverse section of wood, cutting across the tracheid tubes, showing tree rings of fast (big cells, earlywood) and slow seasonal growth

Transverse section of wood, cutting across the tracheid tubes, showing tree rings of fast (big cells, earlywood) and slow seasonal growth

Reproduction

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Conifers produce their seeds inside a protective cone called a strobilus. Most species are monoecious, with male and female cones on the same tree. All conifers are wind-pollinated. In conifers such as pines, the cones are woody, and when mature the scales usually spread open allowing the seeds, which are often winged, to fall out and be dispersed by the wind. In others such as firs and cedars, the cones disintegrate to release the seeds.^[30] Some conifers produce nut-like seeds, such as pine nuts, which are dispersed by birds, in particular, nutcrackers, and jays, which break up the cones.^[31]^[32] In fire-adapted pines such as Pinus radiata, the seeds may be stored in closed cones for many years, being released only when a fire opens the cones.^[33] In families such as Taxaceae, the cone scales are much modified as edible arils, resembling berries. These are eaten by fruit-eating birds, which then pass the seeds in their droppings.^[34]

Pinaceae: unopened female cones of subalpine fir (Abies lasiocarpa)

Pinaceae: unopened female cones of subalpine fir (Abies lasiocarpa)
Northern nutcracker with nut of Pinus sibirica

Northern nutcracker with nut of Pinus sibirica
Taxaceae: the fleshy aril that surrounds each seed in the European yew is a highly modified seed cone scale.

Taxaceae: the fleshy aril that surrounds each seed in the European yew is a highly modified seed cone scale.
Pinaceae: pollen cone of a Japanese larch (Larix kaempferi)

Pinaceae: pollen cone of a Japanese larch (Larix kaempferi)

Life cycle

File:Gymnosperm life cycle diagram-en.svg

Life cycle of a pine tree

Conifers are heterosporous, generating two different types of spores: male microspores and female megaspores.^[35] These spores develop on separate male and female sporophylls on separate male and female cones, usually on the same tree.Template:Sfn

In the male cones, microspores are produced from microsporocytes by meiosis. The microspores develop into pollen grains, which contain the male (micro)gametophytes. Large amounts of pollen are released and carried by the wind. Some pollen grains land on female cones, pollinating them. 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.Template:Sfn

The female cone develops two ovules, each of which contains haploid megaspores. A megasporocyte is divided by meiosis in each ovule. The female gametophytes grow to produce two or more haploid eggs. The fertilized egg, the (diploid) zygote, gives rise to the embryo, and a seed is produced. The female cone then opens, releasing the seeds which grow into seedlings. Some seedlings survive to grow into trees.Template:Sfn

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 have one of three reproductive cycles that differ in the time to complete female strobilus development from initiation to seed maturation. The cycle is one year in genera such as Abies, Picea, Cedrus, and Tsuga; two years in most pine species and in Sequoiadendron; and three years in three pine species including Pinus pinea. All three types have a long gap between pollination and fertilization.Template:Sfn

Distribution and ecology

Conifers are the dominant plants over the taiga forest of the Northern Hemisphere,^[26] forming the world's largest terrestrial biome. The taiga consists mainly of larches, pines, and spruces.^[36] Larch is the most common tree in Russia, and by volume of timber, easily the most abundant tree genus worldwide.^[37] The larch species Larix gmelinii is the world's most northerly-growing tree, at 75° north in the Taymyr Peninsula.^[8] Conifers are widespread also in southern Europe, the Middle East, the Himalayas, Southeast Asia, and Japan. Conifers are not confined to the Northern Hemisphere: around 200 conifer species live only in the tropics, and others live in Australasia, Africa (including Madagascar), and Central and South America.^[38] Species richness decreases with latitude; a northern country like Canada has just 9 species, whereas Mexico has 43, and the tropical island of New Caledonia has 42 endemic species.^[8]

Since conifers cannot regrow their leaves rapidly like hardwoods, leaf diseases can seriously damage coniferous plantations, especially dense stands of young trees. Needle cast diseases, often caused by ascomycete fungi in the Rhytismataceae family, result in leaf fall.^[39] Another ascomycete, Rhizosphaera (Sphaeropsidales), causes severe defoliation and shoot blight, for instance in spruces.^[40]

At least 20 species of roundheaded wood-boring longhorn beetles (Cerambycidae) feed on the wood of spruces, firs, and hemlocks.^[41] Bark beetles (Scolytinae, in the Curculionidae) are destructive pests of commercial forestry; major pests of spruce and other conifers include Ips typographus in Eurasia^[42] and Dendroctonus rufipennis in North America.^[43]

The basidiomycete fungus Boletus pinophilus is among the fungi that form an ectomycorrhizal association with conifers, in its case with pines such as Pinus sylvestris.^[44]

Some conifers introduced for forestry including Pinus radiata have become invasive species in New Zealand,^[45] South Africa,^[46] and Australia.^[47]^[48]

Taiga coniferous forest, mostly larches, pines, and spruces, covers a large area of Siberia (pictured) and Canada.[36]

Taiga coniferous forest, mostly larches, pines, and spruces, covers a large area of Siberia (pictured) and Canada.^[36]
Galleries of Ips typographus bark beetles weaken conifers such as Norway spruce, and can seriously harm commercial forestry.

Galleries of Ips typographus bark beetles weaken conifers such as Norway spruce, and can seriously harm commercial forestry.
The pine bolete Boletus pinophilus forms an ectomycorrhizal association with several pines.

The pine bolete Boletus pinophilus forms an ectomycorrhizal association with several pines.
Pinus radiata (radiata or Monterey pine) is an invasive species in Australia (pictured), New Zealand, and South Africa.

Pinus radiata (radiata or Monterey pine) is an invasive species in Australia (pictured), New Zealand, and South Africa.

Economic importance

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The softwood derived from conifers is more easily worked than hardwood from broadleaved (angiosperm) trees. This makes it widely used and of great economic value, its many uses including construction, furniture, telegraph poles and fencing.^[49] A large part of production is used for paper.^[49]^[50] In the United Kingdom, the 48% of the woodland that is coniferous yields over 90% of the timber; the top species is sitka spruce, yielding about half of the timber produced.^[51] Worldwide, wood products reached a value of $100 billion by the end of the 20th century.^[8] Template:Annotated image 4

Conifers such as fir, cedar, cypress, juniper, spruce, pine, yew and false cedar have been selected by plant breeders for ornamental purposes. Plants with unusual growth habits, sizes, and colours are propagated and planted in parks and gardens throughout the world.^[52]

Commercial forestry using sitka spruce

Commercial forestry using sitka spruce
Globosa, an ornamental cultivar of Scots pine

Globosa, an ornamental cultivar of Scots pine
Pinus thunbergii 'Kotobuki' as a 65-year-old bonsai

Pinus thunbergii 'Kotobuki'
as a 65-year-old bonsai

References

Template:Reflist

External links

Template:Sister project

Conifers at the Tree of Life Web Project
Gymnosperms at Michigan State University
Conifers of the World: Resources for Conifer Research - some 37,000 herbarium records

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Conifer: Difference between revisions

Latest revision as of 21:26, 29 October 2025

Contents

Evolution

Fossil history

Relict species

External phylogeny

Internal phylogeny

Taxonomy

Description

Foliage

Wood

Reproduction

Life cycle

Distribution and ecology

Economic importance

References

External links

Navigation menu

@@ Line 1: / Line 1: @@
-{{Short description|Group of cone-bearing seed plants}}
+{{Short description|Group of seed plants}}
+{{good article}}
 {{Other uses}}
 {{Use dmy dates|date=June 2022}}
-{{EngvarB|date = June 2022}}
+{{Use British English|date=September 2025}}
 {{Automatic taxobox
 | fossil_range = {{fossil range|307|0}}[[Carboniferous]]–[[Holocene|Present]]
 | image = Sapins pectinés.jpg
-| image_caption = Large conifer [[forest]] composed of ''[[Abies alba]]'' at [[Vosges]], Eastern [[France]]
+| image_alt = Conifer forest
+| image_caption = Large conifer [[forest]] of silver fir (''[[Abies alba]]'') at [[Vosges]], Eastern [[France]]
 | display_parents = 2
 | taxon = Pinopsida
@@ Line 30: / Line 32: @@
 * [[Palissyales]] †
 * [[Voltziales]] †
-*[[Cordaitales]] †
+* [[Cordaitales]] †
 | synonyms = * Coniferophyta
 * Coniferae
@@ Line 36: / Line 38: @@
 }}
-'''Conifers''' ({{IPA|ˈkɒnɪfər}}) are a group of [[conifer cone|cone-bearing]] [[Spermatophyte|seed plants]], a subset of [[gymnosperm]]s. Scientifically, they make up the [[phylum|division]] '''Pinophyta''' ({{IPAc-en|p|ɪ|ˈ|n|ɒ|f|ᵻ|t|ə|,_|ˈ|p|aɪ|n|oʊ|f|aɪ|t|ə}}), also known as '''Coniferophyta''' ({{IPAc-en|ˌ|k|ɒ|n|ᵻ|f|ə|ˈ|r|ɒ|f|ᵻ|t|ə|,_|-|oʊ|f|aɪ|t|ə}}) or '''Coniferae'''. The division contains a single extant [[class (biology)|class]], '''Pinopsida'''. All [[Neontology|extant]] conifers are [[perennial plant|perennial]] [[woody plant]]s with [[secondary growth]].{{Efn|This depends on the placement of [[Gnetophytes]], which have been traditionally excluded from the conifers, though recent molecular evidence suggest gnetophytes are the sister to the Pinaceae. See text for details.}} The majority are [[tree]]s, though a few are [[shrub]]s. Examples include [[Cedrus|cedars]], [[Pseudotsuga|Douglas-firs]], [[Cupressaceae|cypresses]], [[fir]]s, [[juniper]]s, [[Agathis|kauri]], [[larch]]es, [[pine]]s, [[Tsuga|hemlocks]], [[Sequoioideae|redwoods]], [[spruce]]s, and [[Taxaceae|yews]].<ref name="Campbell-2005">Campbell, Reece, "Phylum Coniferophyta". ''Biology''. 7th ed. 2005. Print. p. 595.</ref> As of 2002,{{Update inline|date=December 2024}} Pinophyta contained seven families, 60 to 65 genera, and more than 600 living species.
+'''Conifers''' ({{IPAc-en|ˈ|k|ɒ|n|ɪ|f|ər}}) are a group of [[Spermatophyte|seed plants]], a subset of [[gymnosperm]]s. They are mainly [[evergreen]] trees with a regular branching pattern, reproducing with male and female [[conifer cone|cones]], usually [[monoecy|on the same tree]]. They are [[wind-pollinated]] and the seeds are usually dispersed by the wind.
+Scientifically, they make up the [[phylum|division]] '''Pinophyta''', also known as '''Coniferae'''. All [[Neontology|extant]] conifers except for the [[Gnetophytes]] are [[perennial plant|perennial]] [[woody plant]]s with [[secondary growth]].<!--<ref name="Campbell-2005">--> There are over 600 living species.
-Although the total number of species is relatively small, conifers are [[ecology|ecologically]] 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]]s have more [[biodiversity]] and turnover, the immense conifer forests of the world represent the largest terrestrial [[carbon sink]]. Conifers are of great economic value for [[softwood]] [[lumber]] and [[paper]] production.
+Conifers first appear in the fossil record over 300 million years ago in the [[Carboniferous]]. They became dominant land plants in the [[Mesozoic]], until [[flowering plants]] took over many ecosystems in the [[Cretaceous]]. Many conifers today are [[Relict (biology)|relict species]], surviving in a small part of their former ranges. Such relicts include ''[[Wollemia]]'', known only from a small area of Australia, and ''[[Metasequoia glyptostroboides]]'', known from Cretaceous fossils and surviving in a small area of China.
-== Etymology ==
+Although the total number of species is relatively small, conifers are [[ecology|ecologically]] important. They are the dominant plants over the [[taiga]] of the [[Northern Hemisphere]]. Boreal conifers have multiple adaptations to survive winters, including a conical shape to shed snow, strong tracheid vessels to tolerate ice pressure, and a waxy covering on the needle leaves to minimise water loss. Several fungi form [[ectomycorrhiza]]l associations with conifers. Other fungi cause diseases such as [[needle cast]], especially harmful to young trees. Conifers are affected by pest insects such as wood-boring [[longhorn beetle]]s and by [[bark beetle]]s, which make galleries just under the bark.  Conifers are of great economic value for [[timber]] and [[paper]] production.
-''Conifer'' is from Latin, meaning "cone-bearing", from Latin ''conus'' (cone) and ''ferre'' (to bear).<ref>{{cite web |title=conifer (n.) |url=https://www.etymonline.com/word/conifer |publisher=Etymology Online |access-date=23 April 2025}}</ref>
+== Evolution ==
-The division name Pinophyta conforms to the rules of the ''[[International Code of Nomenclature for algae, fungi, and plants]]'' (ICN), which state (Article 16.1) that the names of higher [[taxon|taxa]] 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).
+=== Fossil history ===
+The earliest conifers appear in the fossil record during the Late [[Carboniferous]] ([[Pennsylvanian (geology)|Pennsylvanian]]) over 300 million years ago. Conifers are thought to be most closely related to the [[Cordaitales]]'','' a group of extinct Carboniferous-Permian trees and clambering plants whose reproductive structures had some similarities to those of conifers. The most primitive conifers belong to the paraphyletic assemblage of "[[Walchia|walchian conifers]]", which were small trees, and probably originated in dry upland habitats. The range of conifers expanded during the Early [[Permian]] ([[Cisuralian]]) to lowlands due to increasing aridity. Walchian conifers were gradually replaced by more advanced [[Voltziales|voltzialean]] or "transition" conifers.<ref name="Feng-2017">{{Cite journal |last=Feng |first=Zhuo |date=September 2017 |title=Late Palaeozoic plants |journal=[[Current Biology]] |volume=27 |issue=17 |pages=R905–R909 |bibcode=2017CBio...27.R905F |doi=10.1016/j.cub.2017.07.041 |pmid=28898663 |doi-access=free}}</ref> Conifers were largely unaffected by the [[Permian–Triassic extinction event]],<ref>{{Cite journal |last1=Nowak |first1=Hendrik |last2=Schneebeli-Hermann |first2=Elke |last3=Kustatscher |first3=Evelyn |date=2019-01-23 |title=No mass extinction for land plants at the Permian–Triassic transition |journal=[[Nature Communications]] |volume=10 |issue=1 |page=384 |bibcode=2019NatCo..10..384N |doi=10.1038/s41467-018-07945-w |pmc=6344494 |pmid=30674875 |doi-access=free}}</ref> and were dominant land plants of the [[Mesozoic]] era. Modern groups of conifers emerged from the Voltziales during the Late Permian through [[Jurassic]].<ref name="Leslie-2018">{{Cite journal |last1=Leslie |first1=Andrew B. |last2=Beaulieu |first2=Jeremy |last3=Holman |first3=Garth |last4=Campbell |first4=Christopher S. |last5=Mei |first5=Wenbin |last6=Raubeson |first6=Linda R. |last7=Mathews |first7=Sarah |date=September 2018 |title=An overview of extant conifer evolution from the perspective of the fossil record |journal=[[American Journal of Botany]] |language=en |volume=105 |issue=9 |pages=1531–1544 |doi=10.1002/ajb2.1143 |pmid=30157290 |doi-access=free}}</ref> Conifers underwent a major decline in the [[Late Cretaceous]] corresponding to the explosive [[adaptive radiation]] of [[flowering plant]]s.<ref>{{cite journal |last1=Condamine |first1=Fabien L. |last2=Silvestro |first2=Daniele |last3=Koppelhus |first3=Eva B. |last4=Antonelli |first4=Alexandre |title=The rise of angiosperms pushed conifers to decline during global cooling |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |date=17 November 2020 |volume=117 |issue=46 |pages=28867–28875 |doi=10.1073/pnas.2005571117 |bibcode=2020PNAS..11728867C |pmc=7682372 |pmid=33139543 |doi-access=free }}</ref>
+<gallery class=center mode=nolines widths=180 heights=180>
+File:Conifer fossil.jpg|[[Voltziales]]: ''[[Walchia|Walchia laxifolia]]'' foliage, [[Cisuralian]], Germany
+File:Araucaria mirabilis (fossil cone) (Jurassic; Argentina) (49021443726).jpg|''Araucaria'' cone, [[Jurassic]], Argentina
+File:Elatides sp. (fossil conifer) (Judith River Group, Upper Cretaceous; Montana or Canada) (25210755717).jpg|''Elatides'' foliage, [[Late Cretaceous]], N. America
+File:Σίγρι1.jpg|Base of conifer trunk with roots, [[Early Miocene]], [[Lesbos]], Greece
+</gallery>
-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]]s" may also be used at any [[rank (botany)|rank]] above family. Both are allowed.
+=== Relict species ===
-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, Coniferae, or [[Coniferales]].
+Several [[Neontology|extant]] conifers have [[relict taxon]] status, surviving in small areas or in very small numbers where they once may have been common and widespread. One such is ''[[Wollemia nobilis]]'', discovered in 1994 in some narrow, steep-sided, [[sandstone]] [[gorge]]s in Australia.<ref>{{cite web |title=Wollemia nobilis: The Australian Botanic Garden, Mount Annan – April |url=https://www.rbgsyd.nsw.gov.au/annan/the_garden/Plant_of_the_Month/wollemia_nobilis |archive-url=https://web.archive.org/web/20151019130835/https://www.rbgsyd.nsw.gov.au/annan/the_garden/Plant_of_the_Month/wollemia_nobilis |archive-date=19 October 2015 |access-date=30 October 2015 |publisher=[[Royal Botanic Garden, Sydney]]}}</ref> The wild population consisted of under 60 adult trees with essentially no genetic variability, implying a genetic bottleneck some thousands of years ago.<ref>{{cite report |last1=Stevenson |first1=Dennis Wm. |last2=Ramakrishnan |first2=Srividya |last3=Alves |first3=Cristiane de Santis |last4=Coelho |first4=Laís Araujo |last5=Kramer |first5=Melissa |last6=Goodwin |first6=Sara |last7=Ramos |first7=Olivia Mendevil |last8=Eshel |first8=Gil |last9=Sondervan |first9=Veronica M. |last10=Frangos |first10=Samantha |display-authors=6 |year=2023 |title=The genome of the Wollemi pine, a critically endangered "living fossil" unchanged since the Cretaceous, reveals extensive ancient transposon activity |type=preprint |pmid=37662366 |pmc=10473749 |doi=10.1101/2023.08.24.554647 |url=https://www.biorxiv.org/content/10.1101/2023.08.24.554647v1.full |access-date=2023-09-15 |url-status=live |archive-url=https://web.archive.org/web/20230916150916/https://www.biorxiv.org/content/10.1101/2023.08.24.554647v1.full |archive-date=2023-09-16}}</ref> The extant [[gnetophyte]]s consist of three relict genera, namely ''[[Ephedra (plant)|Ephedra]]'', ''[[Gnetum]]'', and ''[[Welwitschia]]''. Fossils definitely of the group date back to the [[Late Jurassic]], with many species in the Cretaceous.<ref name="Coiro-2022">{{Cite journal |last1=Coiro |first1=Mario |last2=Roberts |first2=Emily A. |last3=Hofmann |first3=Christa-Ch. |last4=Seyfullah |first4=Leyla J. |date=14 December 2022 |title=Cutting the long branches: Consilience as a path to unearth the evolutionary history of Gnetales |journal=[[Frontiers in Ecology and Evolution]] |volume=10 |article-number=1082639 |doi=10.3389/fevo.2022.1082639 |doi-access=free }}</ref> Conifers as a whole, too, declined markedly after the angiosperms (flowering plants) diversified during the Cretaceous, coming to dominate most [[terrestrial ecosystem]]s. Many conifer species became [[Extinction|extinct]], leaving 30 out of 80 genera with just one extant species, and 11 more with just two or three species. The popular phrase "[[living fossil]]s" could, the Dutch botanist [[Aljos Farjon]] states, fittingly be applied to many of these. Thus, ''[[Metasequoia glyptostroboides]]'', the dawn redwood, is known from fossils of Late Cretaceous and [[Miocene]] age, and was found also as an extant tree with a small relict range in China.<ref name="Farjon 1999"/>
-== Taxonomy and evolution ==
+<gallery class=center mode=nolines widths=180 heights=180>
-Conifers are by far the largest and most economically important group of gymnosperms. 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]] (Taxaceae only) and [[Pinales]] (the rest), but recent research into [[molecular biology|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 (biology)|morphology]].
+File:Wakehurst Place woodland Wollemi pine.jpg|''[[Wollemia nobilis]]'' is a [[relict taxon]] known only from a small area in Australia.
+File:Welwitschia at Ugab River basin.jpg|''[[Welwitschia mirabilis]]'' is one of the [[gnetophyte]]s, all relict taxa very unlike other conifers.
+File:Metasequoia glyptostroboides Autumn leaf color.jpg|''[[Metasequoia glyptostroboides]]'' survives in a small part of China, and is known from [[fossil]]s from the [[Late Cretaceous]] onwards.
+</gallery>
+=== External phylogeny===
+The cladogram summarizes the group's external phylogeny. The conifers are [[gymnosperm]]s, sister to a [[clade]] consisting of the [[Ginkgoidae|ginkgos]] and  [[Cycadidae|cycads]].<ref name="Leslie 2018">{{Cite journal |last=Leslie |first=Andrew B. |last2=Beaulieu |first2=Jeremy |last3=Holman |first3=Garth |last4=Campbell |first4=Christopher S. |last5=Mei |first5=Wenbin |last6=Raubeson |first6=Linda R. |last7=Mathews |first7=Sarah |display-authors=et al. |year=2018 |title=An overview of extant conifer evolution from the perspective of the fossil record |journal=[[American Journal of Botany]] |volume=105 |issue=9 |pages=1531–1544 |doi=10.1002/ajb2.1143 |pmid=30157290 |s2cid=52120430}}</ref><ref name="Leslie appendix">{{Cite journal |last=Leslie |first=Andrew B. |display-authors=et al. |year=2018 |title=ajb21143-sup-0004-AppendixS4 |url=https://bsapubs.onlinelibrary.wiley.com/action/downloadSupplement?doi=10.1002%2Fajb2.1143&file=ajb21143-sup-0004-AppendixS4.pdf |journal=[[American Journal of Botany]] |volume=105 |issue=9 |pages=1531–1544 |doi=10.1002/ajb2.1143 |pmid=30157290 |s2cid=52120430}}</ref><ref name="Stull 2021">{{Cite journal |last=Stull |first=Gregory W. |last2=Qu |first2=Xiao-Jian |last3=Parins-Fukuchi |first3=Caroline |last4=Yang |first4=Ying-Ying |last5=Yang |first5=Jun-Bo |last6=Yang |first6=Zhi-Yun |last7=Hu |first7=Yi |last8=Ma |first8=Hong |last9=Soltis |first9=Pamela S. |last10=Soltis |first10=Douglas E. |last11=Li |first11=De-Zhu |last12=Smith |first12=Stephen A. |last13=Yi |first13=Ting-Shuang |display-authors=et al. |year=2021 |title=Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms |url=https://www.nature.com/articles/s41477-021-00964-4 |journal=[[Nature Plants]] |volume=7 |issue=8 |pages=1015–1025 |bibcode=2021NatPl...7.1015S |biorxiv=10.1101/2021.03.13.435279 |doi=10.1038/s41477-021-00964-4 |pmid=34282286 |s2cid=232282918}}</ref><ref name="Stull matrix">{{Cite report |url=https://figshare.com/articles/dataset/Gene_duplications_and_genomic_conflict_underlie_major_pulses_of_phenotypic_evolution_in_gymnosperms/14547354 |title=main.dated.supermatrix.tree.T9.tre |last=Stull |first=Gregory W. |year=2021 |publisher=Figshare |doi=10.6084/m9.figshare.14547354.v1 |display-authors=et al.}}</ref>
+{{clade
+|label1=[[Gymnosperm]]s
+|1={{clade
+   |1={{clade
+      |1=[[Ginkgoidae]] [[File:Addisonia (PLATE 362) - colored illustrations and popular descriptions of plants (1916-(1964)) (16150512234).jpg|60px]]
+      |2=[[Cycadidae]] [[File:Cycas circinalis(draw).jpg|60px]]
+      }}
+   |2='''Conifers''' [[File:Abies alba - Köhler–s Medizinal-Pflanzen-001.jpg|60px]]
+   }}
+}}
-There were seven families of conifers {{Circa|2011}},<ref name="GymnoData">{{cite web|url=http://www.conifers.org/zz/pinales.htm|title=Pinidae (conifers) description – The Gymnosperm Database|url-status=dead|archive-url=https://web.archive.org/web/20160220110331/http://www.conifers.org/zz/pinales.htm|archive-date=2016-02-20}}</ref> with 65–70 genera and over 600 living species ({{Circa|2002|lk=no}}).<ref name="Judd-2002">{{cite book |last1=Judd |first1=W.S |title=Plant systematics, a phylogenetic approach |last2=Campbell |first2=C.S. |last3=Kellogg |first3=E.A. |last4=Stevens |first4=P.F. |last5=Donoghue |first5=M.J. |date=2002 |publisher=Sinauer Associates |isbn=0-87893-403-0 |edition=2nd |location=Sunderland, Massachusetts}}</ref>{{rp|205}}<ref>{{cite journal |last1=Lott |first1=John N.A. |last2=Liu |first2=Jessica C. |last3=Pennell |first3=Kelly A |last4=Lesage |first4=Aude |last5=West |first5=M Marcia |year=2002 |title=Iron-rich particles and globoids in embryos of seeds from phyla Coniferophyta, Cycadophyta, Gnetophyta, and Ginkgophyta: characteristics of early seed plants |journal=Canadian Journal of Botany |volume=80 |issue=9 |pages=954–961 |doi=10.1139/b02-083 |bibcode=2002CaJB...80..954L }}</ref>{{Update inline|date=December 2023|reason=The taxobox shows six conifer families and probably reflects a more recent taxonomy (where gnetophytes are included within Pinophyta and ''Cephalotaxus'' is included in Taxaceae).}} The seven most distinct families are linked in the box above right and phylogenetic diagram left. In other interpretations, the [[Cephalotaxaceae]] may be better included within the Taxaceae, and some authors additionally recognize [[Phyllocladus|Phyllocladaceae]] as distinct from Podocarpaceae (in which it is included here). The family [[Taxodiaceae]] is here included in the 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&nbsp;al.<ref name="Christenhusz-2011">{{cite journal |last1=Christenhusz |first1=M.J.M. |last2=Reveal |first2=J. |last3=Farjon |first3=A.|last4=Gardner.|first4=M.F. |last5=Mill |first5=R.R. |last6=Chase |first6=M.W. |year=2011|title=A new classification and linear sequence of extant gymnosperms |journal=Phytotaxa |volume=19 |issue=1 |pages=55–70 |doi=10.11646/phytotaxa.19.1.3 |bibcode=2011Phytx..19...55C }}</ref>
+=== Internal phylogeny ===
-The conifers are an ancient group, with a [[fossil]] record extending back about 300&nbsp;million years to the [[Paleozoic]] in the late [[Carboniferous]] period; even many of the modern genera are recognizable from fossils 60–120&nbsp;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 [[herb]]aceous conifers with no woody stems.<ref>{{cite journal |last1=Rothwell |first1=Gar W. |last2=Grauvogel-Stamm |first2=Léa |last3=Mapes |first3=Gene |title=An herbaceous fossil conifer: Gymnospermous ruderals in the evolution of Mesozoic vegetation |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |date=February 2000 |volume=156 |issue=1–2 |pages=139–145 |doi=10.1016/S0031-0182(99)00136-4 |bibcode=2000PPP...156..139R }}</ref> Major fossil orders of conifers or conifer-like plants include the [[Cordaitales]], [[Voltziales]] and perhaps also the [[Czekanowskiales]] (possibly more closely related to the [[Ginkgo]]phyta). The cladogram summarizes conifer phylogeny:<ref>Derived from papers by A. Farjon and C. J. Quinn & R. A. Price in the Proceedings of the Fourth International Conifer Conference, ''Acta Horticulturae'' 615 (2003)</ref>
+The [[Gnetophyta]], despite their distinct appearances, were long viewed as outside the conifer group, but phylogenomic analysis indicates that they are part of the conifer clade, sister to the pine family (the 'gnepine' hypothesis). If so, the gnetophytes once shared the distinctive characters of the conifers, and have lost them.<ref name="Chaw 1997">{{cite journal |last1=Chaw |first1=S. M. |last2=Aharkikh |first2=A. |last3=Sung |first3=H. M. |last4=Lau |first4=T. C. |last5=Li |first5=W. H. |year=1997 |title=Molecular phylogeny of extant gymnosperms and seed plant evolution: Analysis of nuclear 18S rRNA sequences |journal=[[Molecular Biology and Evolution]] |volume=14 |issue=1 |pages=56–68 |pmid=9000754 |doi=10.1093/oxfordjournals.molbev.a025702 |doi-access=free}}</ref> The cladogram summarizes the conifers' internal phylogeny:<ref name="Ran 2018">{{cite journal |last1=Ran |first1=Jin-Hua |last2=Shen |first2=Ting-Ting |last3=Wang |first3=Ming-Ming |last4=Wang |first4=Xiao-Quan |title=Phylogenomics resolves the deep phylogeny of seed plants and indicates partial convergent or homoplastic evolution between Gnetales and angiosperms |journal=[[Proceedings of the Royal Society B: Biological Sciences]] |volume=285 |issue=1881 |date=27 June 2018 |pmid=29925623 |pmc=6030518 |doi=10.1098/rspb.2018.1012 |doi-access=free |article-number=20181012 |url=https://royalsocietypublishing.org/doi/pdf/10.1098/rspb.2018.1012 |access-date=20 September 2025}}</ref>
 {{clade|style=line-height:100%;
+|label1='''Pinophyta'''
+|sublabel1=(Coniferae)
 |1={{clade
-    |1=[[Pinaceae]]
+    |1={{clade
-   |2={{clade
        |1={{clade
-          |1=[[Araucariaceae]]
+         |label1=[[Pinaceae]] |sublabel1=pine family
-          |2=[[Podocarpaceae]]
+          |1=[[File:Abies alba - Köhler–s Medizinal-Pflanzen-001.jpg|60px]]
+         |label2=[[Gnetophyta]] |sublabel2=(3 relict genera)
+          |2=[[File:Gnetum gnemon.jpg|60px]]
           }}
        |2={{clade
-          |1=[[Sciadopityaceae]]
+          |1={{clade
+            |label1=[[Araucariaceae]] |sublabel1=monkey puzzle family
+            |1=[[File:Araucaria brasiliana SZ138.jpg|60px]]
+            |label2=[[Podocarpaceae]] |sublabel2=podocarps
+            |2=[[File:Podocarpus macrophyllus SZ134.png|60px]]
+            }}
           |2={{clade
-             |1=[[Cupressaceae]]
+            |label1=[[Sciadopityaceae]] |sublabel1=umbrella pines
+             |1=[[File:Sciadopitys verticillata SZ102.jpg|60px]]
              |2={{clade
-                |1=[[Cephalotaxaceae]]
+               |label1=[[Cupressaceae]] |sublabel1=cypress family
-                |2=[[Taxaceae]]
+                |1=[[File:Cupressus sempervirens (cropped).tiff|60px]]
+                |2={{clade
+                  |label1=[[Cephalotaxaceae]] |sublabel1=plum yew family
+                  |1=[[File:Die winterharten Nadelhölzer Mitteleuropas - ein Handbuch für Gärtner und Gartenfreunde (1909) (20959624281).jpg|50px]]
+                  |label2=[[Taxaceae]] |sublabel2=yew family
+                  |2=[[File:Nouvelle iconographie fourragère (cropped).jpg|60px]]
+                  }}
                 }}
              }}
@@ Line 79: / Line 124: @@
 }}
-Multiple studies indicate that the [[Gnetophyta]] belong within the conifers despite their distinct appearances, either placing them as a [[sister group]] to [[Pinales]] (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.<ref>{{cite journal |last1=Stull |first1=Gregory W. |last2=Qu |first2=Xiao-Jian |last3=Parins-Fukuchi |first3=Caroline |last4=Yang |first4=Ying-Ying |last5=Yang |first5=Jun-Bo |last6=Yang |first6=Zhi-Yun |last7=Hu |first7=Yi |last8=Ma |first8=Hong |last9=Soltis |first9=Pamela S. |last10=Soltis |first10=Douglas E. |last11=Li |first11=De-Zhu |last12=Smith |first12=Stephen A. |last13=Yi |first13=Ting-Shuang |title=Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms |journal=Nature Plants |date=19 July 2021 |volume=7 |issue=8 |pages=1015–1025 |doi=10.1038/s41477-021-00964-4 |pmid=34282286 |bibcode=2021NatPl...7.1015S }}</ref><ref>{{cite journal |last1=Ran |first1=Jin-Hua |last2=Shen |first2=Ting-Ting |last3=Wang |first3=Ming-Ming |last4=Wang |first4=Xiao-Quan |title=Phylogenomics resolves the deep phylogeny of seed plants and indicates partial convergent or homoplastic evolution between Gnetales and angiosperms |journal=Proceedings of the Royal Society B: Biological Sciences |date=27 June 2018 |volume=285 |issue=1881 |pages=20181012 |doi=10.1098/rspb.2018.1012 |pmc=6030518 |pmid=29925623}}</ref><ref>{{cite journal |last1=Farjon |first1=Aljos |title=The Kew Review: Conifers of the World |journal=Kew Bulletin |date=April 2018 |volume=73 |issue=1 |page=8 |doi=10.1007/s12225-018-9738-5 |bibcode=2018KewBu..73....8F |doi-access=free}}
+=== Taxonomy ===
-</ref>
-[[File:Abies lasiocarpa 5922.JPG|thumb|upright|The narrow conical shape of northern conifers, and their downward-drooping limbs, help them shed snow.]]
+The name ''conifer'', meaning 'cone-bearing', derives from Latin {{lang|la|laconus}} (cone) and {{lang|la|ferre}} (to bear).<ref>{{cite web |title=Conifer (n.) |url=https://www.etymonline.com/word/conifer |publisher=[[Online Etymology Dictionary]] |access-date=23 April 2025}}</ref> As recently as 1999, the botanist [[Aljos Farjon]] wrote that while the Coniferae had up to the early 20th century been considered "a natural family",<ref name="Farjon 1999"/><!--p. 159--> comparable to the [[Rosaceae]], he doubted that the conifers or the gymnosperms formed natural groups ([[clade]]s).<ref name="Farjon 1999"/><!--p. 160--> By 2016, the conifers were recognised as a clade, with six families (not including the gnetophytes),<ref name="GymnoData">{{cite web |url=http://www.conifers.org/zz/pinales.htm |title=Pinidae (conifers) description – The Gymnosperm Database |archive-url=https://web.archive.org/web/20160220110331/http://www.conifers.org/zz/pinales.htm|archive-date=2016-02-20}}</ref> 65–70 genera, and over 600 living species ({{Circa|2002|lk=no}}).<ref name="Judd-2002">{{cite book |last1=Judd |first1=W. S. |title=Plant systematics, a phylogenetic approach |last2=Campbell |first2=C. S. |last3=Kellogg |first3=E. A. |last4=Stevens |first4=P. F. |last5=Donoghue |first5=M. J. |date=2002 |publisher=[[Sinauer Associates]] |isbn=0-87893-403-0 |edition=2nd |location=Sunderland, Massachusetts}}</ref>{{rp|205}}<ref>{{cite journal |last1=Lott |first1=John N. A. |last2=Liu |first2=Jessica C. |last3=Pennell |first3=Kelly A. |last4=Lesage |first4=Aude |last5=West |first5=M Marcia |year=2002 |title=Iron-rich particles and globoids in embryos of seeds from phyla Coniferophyta, Cycadophyta, Gnetophyta, and Ginkgophyta: characteristics of early seed plants |journal=[[Canadian Journal of Botany]] |volume=80 |issue=9 |pages=954–961 |doi=10.1139/b02-083 |bibcode=2002CaJB...80..954L }}</ref><ref>{{cite book |last1=Díaz-Sala |first1=Carmen |last2=Cabezas |first2=José Antonio |last3=de Simón |first3=Brígida Fernández |last4=Abarca |first4=Dolores |last5=Guevara |first5=M. Ángeles |last6=de Miguel |first6=Marina |last7=Cadahía |first7=Estrella |last8=Aranda |first8=Ismael |last9=Cervera |first9=María-Teresa |display-authors=5 |title=From Plant Genomics to Plant Biotechnology |chapter=The uniqueness of conifers |publisher=[[Elsevier]] |year=2013 |isbn=978-1-907568-29-9 |doi=10.1533/9781908818478.67 |page=67–96}}</ref> Depending on interpretation, the [[Cephalotaxaceae]] may or may not be included within the Taxaceae, while some authors recognize [[Phyllocladus|Phyllocladaceae]] as distinct from Podocarpaceae. The family [[Taxodiaceae]] is here included in the family Cupressaceae.<ref name="Christenhusz-2011">{{cite journal |last1=Christenhusz |first1=M. J. M. |last2=Reveal |first2=J. |last3=Farjon |first3=Aljos |author3-link=Aljos Farjon |last4=Gardner |first4=M. F. |last5=Mill |first5=R. R. |last6=Chase |first6=M. W. |year=2011 |title=A new classification and linear sequence of extant gymnosperms |journal=[[Phytotaxa]] |volume=19 |issue=1 |pages=55–70 |doi=10.11646/phytotaxa.19.1.3 |bibcode=2011Phytx..19...55C }}</ref>
-=== Fossil history ===
+== Description ==
-The earliest conifers appear in the fossil record during the Late [[Carboniferous]] ([[Pennsylvanian (geology)|Pennsylvanian]]), over 300 million years ago. Conifers are thought to be most closely related to the [[Cordaitales]]'','' a group of extinct Carboniferous-Permian trees and clambering plants whose reproductive structures had some similarities to those of conifers. The most primitive conifers belong to the paraphyletic assemblage of "[[Walchia|walchian conifers]]", which were small trees, and probably originated in dry upland habitats. The range of conifers expanded during the Early [[Permian]] ([[Cisuralian]]) to lowlands due to increasing aridity. Walchian conifers were gradually replaced by more advanced [[Voltziales|voltzialean]] or "transition" conifers.<ref name="Feng-2017">{{Cite journal |last=Feng |first=Zhuo |date=September 2017 |title=Late Palaeozoic plants |journal=Current Biology |volume=27 |issue=17 |pages=R905–R909 |bibcode=2017CBio...27.R905F |doi=10.1016/j.cub.2017.07.041 |pmid=28898663 |doi-access=free}}</ref> Conifers were largely unaffected by the [[Permian–Triassic extinction event]],<ref>{{Cite journal |last1=Nowak |first1=Hendrik |last2=Schneebeli-Hermann |first2=Elke |last3=Kustatscher |first3=Evelyn |date=2019-01-23 |title=No mass extinction for land plants at the Permian–Triassic transition |journal=Nature Communications |language=en |volume=10 |issue=1 |pages=384 |bibcode=2019NatCo..10..384N |doi=10.1038/s41467-018-07945-w |pmc=6344494 |pmid=30674875 |doi-access=free}}</ref> and were dominant land plants of the [[Mesozoic]] era. Modern groups of conifers emerged from the Voltziales during the Late Permian through [[Jurassic]].<ref name="Leslie-2018">{{Cite journal |last1=Leslie |first1=Andrew B. |last2=Beaulieu |first2=Jeremy |last3=Holman |first3=Garth |last4=Campbell |first4=Christopher S. |last5=Mei |first5=Wenbin |last6=Raubeson |first6=Linda R. |last7=Mathews |first7=Sarah |date=September 2018 |title=An overview of extant conifer evolution from the perspective of the fossil record |journal=American Journal of Botany |language=en |volume=105 |issue=9 |pages=1531–1544 |doi=10.1002/ajb2.1143 |pmid=30157290 |doi-access=free}}</ref> Conifers underwent a major decline in the [[Late Cretaceous]] corresponding to the explosive [[adaptive radiation]] of [[flowering plant]]s.<ref>{{cite journal |last1=Condamine |first1=Fabien L. |last2=Silvestro |first2=Daniele |last3=Koppelhus |first3=Eva B. |last4=Antonelli |first4=Alexandre |title=The rise of angiosperms pushed conifers to decline during global cooling |journal=Proceedings of the National Academy of Sciences |date=17 November 2020 |volume=117 |issue=46 |pages=28867–28875 |doi=10.1073/pnas.2005571117 |bibcode=2020PNAS..11728867C |pmc=7682372 |pmid=33139543 |doi-access=free }}</ref>
+All living conifers (except the gnetophytes) are woody plants, and most are trees with narrow leaves, often needle-like. There are separate male and female reproductive structures, the cones. Pollination is always by wind; the seeds are mostly winged. The trees have a regular branching pattern. Many conifers have distinctly scented [[resin]].<ref name="Mitchell 1985">{{cite book |last1=Mitchell |first1=Alan F. |last2=Edlin |first2=Herbert L. |author-link2=Herbert L. Edlin |title=Conifers: Forestry Commission Booklet No. 15 |year=1985 |orig-date=1966 |edition=3rd |publisher=[[HMSO]] |pages=4–5 |url=https://cdn.forestresearch.gov.uk/1985/03/fcbk015_3ed.pdf |isbn=978-0-11-710040-4}}</ref>
+The world's tallest and oldest living trees are conifers. The tallest is a [[Sequoia sempervirens|coast redwood]] (''Sequoia sempervirens''), with a height of {{convert|116.07|m|ft}}.<ref name="Ghose">{{cite news |last=Ghose |first=Tia |date=May 23, 2022 |title=What is the world's tallest tree? |url=https://www.livescience.com/28729-tallest-tree-in-world.html |publisher=[[LiveScience]]}}</ref> Among the smallest conifers is the [[Lepidothamnus laxifolius|pygmy pine]] (''Lepidothamnus laxifolius'') of New Zealand, which is seldom taller than 30 cm when mature.<ref>{{cite web |last=Wassilieff |first=Maggy |title=Conifers |publisher=[[Te Ara: The Encyclopedia of New Zealand]] |date=1 Mar 2009 |url=http://www.teara.govt.nz/en/conifers/6/5 |access-date=17 December 2012 |archive-date=1 March 2010 |archive-url=https://web.archive.org/web/20100301031106/http://www.teara.govt.nz/en/conifers/6/5 |url-status=live }}</ref> The oldest non-clonal living tree is a Great Basin bristlecone pine (''[[Pinus longaeva]]''), 4,700 years old.<ref name="Dallimore 1967">{{cite book |last1=Dallimore |first1=W. |last2=Jackson |first2=A. B. |last3=Harrison |first3=S. G. |year=1967 |title=A handbook of Coniferae and Ginkgoaceae |edition=4th |location=New York |publisher=[[St. Martin's Press]] |page=xix}}</ref>
+[[wikt:Boreal|Boreal]] conifers have multiple [[adaptation]]s to survive winters, including the tree's conical shape to shed snow, strong tracheid vessels to tolerate ice pressure, and a waxy covering on the needle leaves to minimise water loss.<ref name="Michigan">{{cite web |title=Winter Adaptations of Trees |url=https://mff.forest.mtu.edu/Environment/WinterTrees.htm |publisher=[[Michigan Technological University]] |access-date=20 September 2025}}</ref>
-==Description==
+<gallery class=center mode=nolines widths=200 heights=200>
+File:US 199 Redwood Highway.jpg|Tallest: ''[[Sequoia sempervirens]]'' can reach a height of {{convert|116.07|m|ft}}.<ref name="Ghose"/>
+File:Big_bristlecone_pine_Pinus_longaeva.jpg|Oldest: ''[[Pinus longaeva]]'' can reach an age of 4,700 years.<ref name="Dallimore 1967"/>
+File:Snow falling at Tower (f6d906ff-4e12-4375-97d6-a21bedf95d8b).jpg|The narrow conical shape of [[wikt:Boreal|boreal]] conifers, and their downward-drooping limbs, help them shed snow.<ref name="Michigan"/>
+</gallery>
-{{refimprove section|date=April 2025}}
+=== Foliage ===
-[[File: 00 29 0496 Waipoua Forest NZ - Kauri Baum Tane Mahuta.jpg|thumb|right|[[Tāne Mahuta]], the biggest [[kauri]] (''Agathis australis'') tree alive, in the [[Waipoua Forest]] of the Northland Region of [[New Zealand]].]]
+Most conifers are evergreens.<ref name="Campbell-2005">{{cite book |last=Campbell |first=Reece |chapter=Phylum Coniferophyta |title=Biology |edition=7th |year=2005 |page=595}}</ref> In many species such as pines, firs, and [[cedrus|cedar]]s, the [[leaf|leaves]] are long, thin and needle-like. Others like [[Cupressaceae|cypresses]] have flat, triangular scale-like leaves.<ref>{{cite web |title=Conifer Tree ID by  Leaf and Needle Shape |url=https://www.treeguideuk.co.uk/conifer-key/ |website=Treeguide |access-date=20 September 2025}}</ref> In the majority of conifers, the leaves are arranged spirally, the exceptions being most of Cupressaceae and one genus in Podocarpaceae, where they are arranged in [[Decussation|decussate]] opposite pairs or [[Whorl (botany)|whorl]]s of 3 or 4. In many species with spirally arranged leaves, such as ''[[Abies grandis]]'', 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]]''). The [[stoma]]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]]s or under forest canopy shade. Conifers from lower latitudes with high sunlight levels (e.g. Turkish pine ''[[Pinus brutia]]'') often have yellower-green leaves, while others (e.g. [[blue spruce]], ''Picea pungens'') may develop blue or silvery leaves to reflect [[ultraviolet]] light. In the great majority of genera the leaves remain on the plant for several (2–40) years before falling, but five genera (''[[larch|Larix]]'', ''[[Pseudolarix]]'', ''[[Glyptostrobus]]'', ''[[Metasequoia]]'' and ''[[Taxodium]]'') are [[deciduous]], shedding their leaves in autumn.<ref name="Campbell-2005"/> The seedlings of some conifers, including pines, have a distinct juvenile foliage period where the leaves are different from the typical adult leaves.<ref name="RHS-Dict-1992">{{cite book |title=Royal Horticultural Society Dictionary of Gardening |publisher=[[Macmillan Press]]; Stockton Press |date=1992 |isbn=1-56159-001-0 |volume=3 |pages=582–594}}</ref>
-All living conifers are woody plants, and most are trees, the majority having a 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 [[fungus|fungal]] infection of wounds. Fossilized resin hardens into [[amber]], which has been commercially exploited historically (for example, in New Zealand's 19th-century [[kauri gum]] industry).
+<gallery class=center mode=nolines widths=180 heights=180>
+File:20160118Pinus sylvestris1.jpg|[[Pinaceae]]: needle-like leaves of Scots pine (''[[Pinus sylvestris]]'')
+File:Araucaria Leaves.JPG|[[Araucariaceae]]: awl-like leaves of Cook pine (''[[Araucaria columnaris]]'')
+File:Abies grandis 5359.JPG|In ''[[Abies grandis]]'' and many other species with spirally arranged leaves, each leaf is twisted near its base to maximize light capture.
+File:C lawsoniana Lge.jpg|[[Cupressaceae]]: scale leaves of [[Chamaecyparis lawsoniana|Lawson's cypress]] (''Chamaecyparis lawsoniana''); scale in mm
+</gallery>
-The size of mature conifers varies from less than one metre to over 100 metres in height.<ref>{{cite book |last1=Enright |first1=Neal J. |last2=Hill |first2=Robert S. |title=Ecology of the Southern Conifers |date=1995 |publisher=Melbourne University Press |isbn=978-0-522-84566-2 }}{{pn|date=January 2025}}</ref> The world's tallest, thickest, largest, and oldest living trees are all conifers. The tallest is a [[Sequoia sempervirens|coast redwood]] (''Sequoia sempervirens''), with a height of 115.55 metres (although one mountain ash, ''[[Eucalyptus regnans]]'', allegedly grew to a height of 140 metres,<ref>{{Cite news |date=1872-02-22 |title=STATE FOREST OF THE WATTS RIVER. |url=http://nla.gov.au/nla.news-article197448140 |access-date=2024-04-06 |work=Age}}</ref> the tallest living [[Flowering plant|angiosperms]] are significantly smaller at around 100 metres.<ref>{{Cite journal |last1=Shenkin |first1=Alexander |last2=Chandler |first2=Chris J. |last3=Boyd |first3=Doreen S. |last4=Jackson |first4=Toby |last5=Disney |first5=Mathias |last6=Majalap |first6=Noreen |last7=Nilus |first7=Reuben |last8=Foody |first8=Giles |last9=bin Jami |first9=Jamiluddin |last10=Reynolds |first10=Glen |last11=Wilkes |first11=Phil |last12=Cutler |first12=Mark E. J. |last13=van der Heijden |first13=Geertje M. F. |last14=Burslem |first14=David F. R. P. |last15=Coomes |first15=David A. |date=2019 |title=The World's Tallest Tropical Tree in Three Dimensions |journal=Frontiers in Forests and Global Change |volume=2 |page=32 |doi=10.3389/ffgc.2019.00032 |doi-access=free |bibcode=2019FrFGC...2...32S |hdl=2164/12435 |hdl-access=free }}</ref><ref>{{Cite news |date=2018-12-11 |title=100 metres and growing: Australia's tallest tree leaves all others in the shade |url=https://www.abc.net.au/news/2018-12-12/new-milestone-for-australias-tallest-tree-centurion/10604588 |access-date=2024-04-06 |work=ABC News |language=en-AU}}</ref>) The thickest (that is, the [[Árbol del Tule|tree with the greatest trunk diameter]]) is a [[Taxodium mucronatum|Montezuma cypress]] (''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.<ref>{{cite book |last=Vidaković |first=Mirko |title=Conifers: Morphology and Variation |date=1991 |publisher=CAB International |isbn=978-86-399-0279-7 }}{{pn|date=January 2025}}</ref> The smallest is the [[Lepidothamnus laxifolius|pygmy pine]] (''Lepidothamnus laxifolius'') of New Zealand, which is seldom taller than 30&nbsp;cm when mature.<ref>{{cite web |last=Wassilieff |first=Maggy |title=Conifers |publisher=Te Ara&nbsp;– the Encyclopedia of New Zealand updated 1-Mar-09 |url=http://www.teara.govt.nz/en/conifers/6/5 |access-date=17 December 2012 |archive-date=1 March 2010 |archive-url=https://web.archive.org/web/20100301031106/http://www.teara.govt.nz/en/conifers/6/5 |url-status=live }}</ref> The oldest non-clonal living tree is a Great Basin bristlecone pine (''[[Pinus longaeva]]''), 4,700 years old.<ref>{{cite book|last1=Dallimore|first1=W.|first2=A.B.|last2=Jackson|first3=S.G.|last3=Harrison|year=1967|title=A handbook of Coniferae and Ginkgoaceae|edition=4th|location=New York|publisher=St. Martin's Press|page=xix}}</ref>
+=== Wood ===
-===Foliage===
+Conifer [[xylem|wood]] consists of two types of [[cell (biology)|cells]]: [[parenchyma]], which have an oval or polyhedral shape, and strongly elongated [[tracheid]]s. 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]]s. Then, the first tracheids of the transition zone are formed, where the radial size of cells and the thickness of their cell walls changes considerably. Finally, latewood tracheids are formed, with small radial sizes and greater cell wall thickness. This is the basic pattern of the internal cell structure of conifer [[tree ring]]s.<ref name="Ledig-1982">{{cite journal |last1=Ledig |first1=F. Thomas |last2=Porterfield |first2=Richard L. |date=1982 |title=Tree Improvement in Western Conifers: Economic Aspects |journal=[[Journal of Forestry]] |volume=80 |issue=10 |pages=653–657 |doi=10.1093/jof/80.10.653 |osti=5675533 }}</ref>
-[[File:Pseudotsuga menziesii 06280.JPG|left|thumb|upright|[[Pinaceae]]: needle-like leaves and vegetative buds of Coast Douglas fir ([[Pseudotsuga menziesii var. menziesii|''Pseudotsuga menziesii'' var. ''menziesii'']])]]
-[[File:Araucaria Leaves.JPG|thumb|[[Araucariaceae]]: awl-like leaves of Cook pine (''[[Araucaria columnaris]]'')]]
-[[File:Abies grandis 5359.JPG|left|thumb|upright|In ''[[Abies grandis]]'' (''grand fir''), and many other species with spirally arranged leaves, leaf bases are twisted to flatten their arrangement and maximize light capture.]]
-[[File:C lawsoniana Lge.jpg|thumb|[[Cupressaceae]]: scale leaves of [[Chamaecyparis lawsoniana|Lawson's cypress]] (''Chamaecyparis lawsoniana''); scale in mm]]
-Since most conifers are evergreens,<ref name="Campbell-2005" /> the [[leaf|leaves]] of many conifers are long, thin and have a needle-like appearance, but others, including most of the [[Cupressaceae]] and some of the [[Podocarpaceae]], have flat, triangular scale-like leaves. Some, notably ''[[Agathis]]'' in Araucariaceae and ''[[Nageia]]'' in Podocarpaceae, have broad, flat strap-shaped leaves. Others such as ''[[Araucaria columnaris]]'' have leaves that are awl-shaped. In the majority of conifers, the leaves are arranged spirally, the exceptions being most of Cupressaceae and one genus in Podocarpaceae, where they are arranged in decussate opposite pairs or whorls of 3 (−4).
+<gallery class=center mode=nolines widths=180 heights=180>
+File:Abies concolor tangential.jpg|Vertical (tangential) section of ''[[Abies concolor]]'' wood ([[xylem]]), showing [[tracheid]]s as long overlapping tubes. Perforation pits (small circles) allow water to move from one tracheid to the next.
+File:Report on the relation of railroads to forest supplies and forestry - together with appendices on the structure of some timber ties, their behavior, and the cause of their decay in the road bed, on (14755970324).jpg|Transverse section of wood, cutting across the [[tracheid]] tubes, showing [[tree ring]]s of fast (big cells, earlywood) and slow seasonal growth
+</gallery>
-In many species with spirally arranged leaves, such as ''[[Abies grandis]]'' (pictured), the leaf bases are twisted to present the leaves in a very flat plane for maximum light capture. Leaf size varies from 2&nbsp;mm in many scale-leaved species, up to 400&nbsp;mm long in the needles of some pines (e.g. Apache pine, ''[[Pinus engelmannii]]''). The [[stoma]]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]]s or under forest canopy shade.
+=== Reproduction ===
-Conifers from lower latitudes with high sunlight levels (e.g. Turkish pine ''[[Pinus brutia]]'') often have yellower-green leaves, while others (e.g. [[blue spruce]], ''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 (''[[larch|Larix]]'', ''[[Pseudolarix]]'', ''[[Glyptostrobus]]'', ''[[Metasequoia]]'' and ''[[Taxodium]]'') are [[deciduous]], shedding their leaves in autumn.<ref name="Campbell-2005" /> 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===
-[[File:Report on the relation of railroads to forest supplies and forestry - together with appendices on the structure of some timber ties, their behavior, and the cause of their decay in the road bed, on (14755970324).jpg|thumb|A thin transverse section showing the internal structure of conifer wood]]
-[[Dendrochronology#Growth rings|Tree rings]] are records of the [[wikt:influence|influence]] of [[Ecology#Physical environments|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 [[cell (biology)|cells]]: '''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]]s. Then, the first tracheids of the transition zone are formed, where the radial size of cells and the thickness of their cell walls changes considerably. Finally, latewood tracheids are formed, with small radial sizes and greater cell wall thickness. This is the basic pattern of the internal cell structure of conifer tree rings.<ref name="Ledig-1982">{{cite journal |last1=Ledig |first1=F. Thomas |last2=Porterfield |first2=Richard L. |date=1982 |title=Tree Improvement in Western Conifers: Economic Aspects |journal=Journal of Forestry |volume=80 |issue=10 |pages=653–657 |doi=10.1093/jof/80.10.653 |osti=5675533 }}</ref>
-===Reproduction===
 {{Main|Conifer cone}}
-Most conifers are [[Plant reproductive morphology#Terminology|monoecious]], but some are [[Plant reproductive morphology#Terminology|subdioecious]] or [[Plant reproductive morphology#Terminology|dioecious]]; all are [[Anemophily|wind-pollinated]]. 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 {{Convert|2 to 600|mm|frac=8}} long.
+Conifers produce their seeds inside a [[conifer cone|protective cone]] called a strobilus. Most species are [[Plant reproductive morphology#Terminology|monoecious]], with male and female cones on the same tree. All conifers are [[Anemophily|wind-pollinated]]. In conifers such as pines, the cones are [[wood]]y, and when mature the scales usually spread open allowing the seeds, which are often winged, to fall out and be dispersed by the [[wind]]. In others such as firs and cedars, the cones disintegrate to release the seeds.<ref name="Treeguide">{{cite web |title=Conifer Life Cycle |url=https://www.treeguideuk.co.uk/conifer-life-cycle/ |website=Tree Guide UK |access-date=20 September 2025}}</ref>
+Some conifers produce nut-like seeds, such as [[pine nut]]s, which are dispersed by [[bird]]s, in particular, [[Nutcracker (bird)|nutcracker]]s, and [[jay]]s, which break up the cones.<ref>{{cite book |last=Lanner |first=Ronald M. |title=Made for each other: A symbiosis of birds and pines |date=1996 |publisher=[[Oxford University Press]] |location=Oxford |isbn=0-19-508-903-0 |pages=61–75}}</ref><ref>{{cite book |last1=Tomback |first1=Diana F. |author-link=Diana Tomback |editor1-last=Sekercioglu |editor1-first=Cagan |editor2-last=Wenny |editor2-first=Daniel G. |editor3-last=Whelan |editor3-first=Christopher J. |title=Why birds matter: avian ecological function and ecosystem services |date=2016 |publisher=[[University of Chicago Press]] |location=Chicago |isbn=0-226-38263-X |page=201 |chapter=7}}</ref>
+In fire-adapted pines such as ''Pinus radiata'', the seeds may be stored in closed cones for many years, being released only when a [[Pyriscence|fire opens the cones]].<ref name="Rushforth-1987">{{cite book |last=Rushforth |first=Keith |title=Conifers |publisher=[[Christopher Helm Publishers]] |publication-place=London |date=1987-01-01 |isbn=0-7470-2801-X |pages=158–192}}</ref>
+In families such as [[Taxaceae]], the cone scales are much modified as edible [[aril]]s, resembling berries. These are eaten by fruit-eating birds, which then pass the seeds in their droppings.<ref>{{cite web |title=Tree ID: Yew tree |url=https://parks.wa.gov/about/news-center/field-guide-blog/tree-id-yew-tree |publisher=[[Washington State Parks]] |access-date=20 September 2025}}</ref>
-In [[Pinaceae]], [[Araucariaceae]], [[Sciadopityaceae]] and most [[Cupressaceae]], the cones are [[wood]]y, and when mature the scales usually spread open allowing the seeds to fall out and be dispersed by the [[wind]]. In some (e.g. [[fir]]s and [[Cedrus|cedar]]s), the cones disintegrate to release the seeds, and in others (e.g. the [[pine]]s that produce [[pine nut]]s) the nut-like seeds are dispersed by [[bird]]s (mainly [[Nutcracker (bird)|nutcracker]]s, and [[jay]]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.
+<gallery class=center mode=nolines widths=180 heights=180>
-In the families [[Podocarpaceae]], [[Cephalotaxaceae]], [[Taxaceae]], and one [[Cupressaceae]] genus (''[[Juniper]]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]]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 [[sporangium|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 [[wikt:micropyle|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 plant]]s 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 [[mitochondria]]l [[organelle]]s to the [[embryo]],{{citation needed|date=June 2021}} a sort of [[meiosis|meiotic]] drive that perhaps explains why ''[[Pinus]]'' and other conifers are so productive, and perhaps also has bearing on observed sex-ratio bias.{{citation needed|date=August 2021}}
-<gallery class="center">
 File:Abies lasiocarpa 6972.JPG|Pinaceae: unopened female cones of [[Abies lasiocarpa|subalpine fir]] (''Abies lasiocarpa'')
-Taxus baccata MHNT.jpg|Taxaceae: the fleshy aril that surrounds each seed in the [[Taxus baccata|European yew]] (''Taxus baccata'') is a highly modified seed cone scale
+File:Spotted nutcracker with pine nut (cropped).jpg|[[Northern nutcracker]] with nut of ''[[Pinus sibirica]]''
+Taxus baccata MHNT.jpg|Taxaceae: the fleshy [[aril]] that surrounds each seed in the [[Taxus baccata|European yew]] is a highly modified seed cone scale.
 Japanese Larch pollen cone, Cardiff, Wales.jpg|Pinaceae: pollen cone of a [[Japanese larch]] (''Larix kaempferi'')
 </gallery>
-===Life cycle===
+=== Life cycle ===
-Conifers are [[heterosporous]], generating two different types of spores: male [[microspore]]s and female [[megaspore]]s. These spores develop on separate male and female [[sporophylls]] on separate male and female cones. In the male cones, microspores are produced from microsporocytes by [[meiosis]]. The microspores develop into pollen grains, which contain the 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 develop into a female [[multicellular]] gametophyte. The female gametophytes grow to produce two or more [[archegonia]], each of which contains an egg. Upon fertilization, the [[diploid]] egg gives rise to the embryo, and a seed is produced. The female cone then opens, releasing the seeds which grow to a young [[seedling]].
+[[File:Gymnosperm life cycle diagram-en.svg|thumb|upright=1.35|[[Biological life cycle|Life cycle]] of a pine tree]]
-# To fertilize the ovum, the male cone releases [[pollen]] that is carried in the wind to the female cone. This is [[pollination]]. (Male and female cones usually occur on the same plant.)
+Conifers are [[heterosporous]], generating two different types of spores: male [[microspore]]s and female [[megaspore]]s.<ref>{{cite book |last=Williams |first=Claire G. |title=Conifer Reproductive Biology |publisher=[[Springer Science+Business Media]] |publication-place=Dordrecht |year=2009 |isbn=978-1-4020-9601-3 |page=9}}</ref> These spores develop on separate male and female [[sporophylls]] on separate male and female cones, usually on the same tree.{{sfn|Williams|2009|pp=25–35}}
-# The pollen fertilizes the female gamete (located in the female cone). Fertilization in some species does not occur until 15 months after pollination.<ref>{{Cite web |url=http://bioserv.fiu.edu/~biolab/labs/1011/Spring%202009/TA%20notes%20and%20pictures/Week%205%20-%20Seed%20Plants.htm |title=Gymnosperms |access-date=2014-05-11 |archive-date=2015-05-27 |archive-url=https://web.archive.org/web/20150527001621/http://bioserv.fiu.edu/~biolab/labs/1011/Spring%202009/TA%20notes%20and%20pictures/Week%205%20-%20Seed%20Plants.htm |url-status=dead }}</ref>
-# 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 ====
+In the male cones, microspores are produced from microsporocytes by [[meiosis]]. The microspores develop into pollen grains, which contain the male (micro)gametophytes. Large amounts of pollen are released and carried by the wind. Some pollen grains land on female cones, pollinating them. 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.{{sfn|Williams|2009|pp=25–35}}
-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]].
-'''One year reproductive cycle''': The genera include ''[[Abies]]'', ''[[Picea]]'', ''[[Cedrus]]'', ''[[Pseudotsuga]],'' ''[[Tsuga]]'', ''[[Keteleeria]]'' ''([[Pinaceae]])'' and ''[[Cupressus]], [[Thuja]], [[Cryptomeria]], [[Cunninghamia]]'' and ''[[Sequoia (genus)|Sequoia]] ([[Cupressaceae]])''. Female strobili are initiated in late summer or fall of 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.<ref name="Singh-1978">{{cite book |last=Singh |first=Hardev |title=Embryology of Gymnosperms |date=1978 |publisher=Gerbrüder Borntraeger |isbn=978-3-443-14011-3 }}{{pn|date=January 2025}}</ref>
+The female cone develops two ovules, each of which contains haploid megaspores. A megasporocyte is divided by meiosis in each ovule. The female gametophytes grow to produce two or more haploid eggs. The fertilized egg, the ([[diploid]]) [[zygote]], gives rise to the [[embryo]], and a [[seed]] is produced. The female cone then opens, releasing the seeds which grow into [[seedling]]s. Some seedlings survive to grow into trees.{{sfn|Williams|2009|pp=25–35}}
-'''Two-year reproductive cycle''': The genera includes ''[[Widdringtonia]]'', ''[[Sequoiadendron]]'' (''[[Cupressaceae]]'') 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 second year [[archegonia]] form in the conelet. Fertilization of the archegonia occurs by early summer of the second year, so the pollination-fertilization interval exceeds a year. After fertilization, the conelet is considered an immature cone. Maturation occurs by autumn of the second year, at which time seeds are shed. In summary, the one-year and the two-year cycles differ mainly in the duration of the pollination-fertilization interval.<ref name="Singh-1978"/>
+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 have one of three reproductive cycles that differ in the time to complete female strobilus development from initiation to seed maturation. The cycle is one year in genera such as ''Abies'', ''Picea'', ''Cedrus'', and ''Tsuga''; two years in most pine species and in ''[[Sequoiadendron]]''; and three years in three pine species including ''[[Pinus pinea]]''. All three types have a long gap between [[pollination]] and [[fertilization]].{{sfn|Williams|2009|pp=101–102}}
-'''Three-year reproductive cycle''': Three of the conifer species are [[pine]] species (''[[Pinus pinea]]'', ''[[Pinus leiophylla]]'', ''[[Pinus torreyana]]'') which have pollination and fertilization events separated by a two-year interval. Female strobili initiated during late summer or autumn of a year, then overwinter until the following spring. Female [[strobili]] emerge then pollination occurs in spring of the second year then the pollinated strobili become conelets in the same year (i.e. the second year). The female [[gametophytes]] in the conelet develop so slowly that the [[megaspore]] does not go through free-nuclear divisions until autumn of the third year. The conelet then overwinters again in the free-nuclear female gametophyte stage. Fertilization takes place by early summer of the fourth year and seeds mature in the cones by autumn of the fourth year.<ref name="Singh-1978"/>
+== Distribution and ecology ==
-==== Tree development ====
+Conifers are the dominant plants over the [[taiga]] forest of the [[Northern Hemisphere]],<ref name="Campbell-2005"/> forming the world's largest terrestrial [[biome]]. The taiga consists mainly of larches, pines, and spruces.<ref name="Berkeley">{{cite web |url=http://www.ucmp.berkeley.edu/exhibits/biomes/forests.php#boreal |title=The forest biome |publisher=[[University of California Museum of Paleontology]] |location=Berkeley |access-date=12 May 2019 |archive-url=https://web.archive.org/web/20190620145416/https://ucmp.berkeley.edu/exhibits/biomes/forests.php#boreal |archive-date=20 June 2019 }}</ref> Larch is the most common tree in Russia, and by volume of timber, easily the most abundant tree genus worldwide.<ref name="Tsepliaev">{{cite book |last=Tsepliaev |first=Vasilii P. |date=1965 |title=The Forests of the U.S.S.R. |location=Jerusalem |publisher=Israel Program for Scientific Translations |page=289 (Table 86)}}</ref> The larch species ''[[Larix gmelinii]]'' is the world's most northerly-growing tree, at 75° north in the [[Taymyr Peninsula]].<ref name="Farjon 1999"/> Conifers are widespread also in southern Europe, the [[Middle East]], the [[Himalayas]], [[Southeast Asia]], and Japan. Conifers are not confined to the Northern Hemisphere: around 200 conifer species live only in the tropics, and others live in Australasia, Africa (including Madagascar), and Central and South America.<ref>{{cite web |title=Conifers of the World: Resources for Conifer Research |url=https://herbaria.plants.ox.ac.uk/bol/conifers |publisher=[[Oxford University Herbaria]] |access-date=20 September 2025}}</ref> Species richness decreases with latitude; a northern country like Canada has just 9 species, whereas Mexico has 43, and the tropical island of [[New Caledonia]] has 42 [[Endemism|endemic]] species.<ref name="Farjon 1999">{{cite journal |last=Farjon |first=Aljos |author-link=Aljos Farjon |title=Introduction to the Conifers |journal=[[Curtis's Botanical Magazine]] |volume=16 |issue=3 |year=1999 |pages=158–172 |jstor=45065379}}</ref>
-The growth and form of a forest tree are the result of activity in the primary and secondary [[meristem]]s, influenced by the distribution of photosynthate from its needles and the hormonal gradients controlled by the apical meristems.<ref name="Fraser-1964">{{cite journal|last1=Fraser|first1=D.A.|last2=Belanger|first2=L.|last3=McGuire|first3=D.|last4=Zdrazil|first4=Z.|year=1964|title=Total growth of the aerial parts of a white spruce tree at Chalk River, Ontario, Canada|journal=Can. J. Bot.|volume=42|issue=2 |pages=159–179|doi=10.1139/b64-017 |bibcode=1964CaJB...42..159F }}</ref> 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 [[herb]]s and [[shrub]]s 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&nbsp;million weighing 14.25&nbsp;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)<ref name="Fraser-1964" /> speculated that if the photosynthate used in making apical growth in 1961 was manufactured the previous year, then the 4&nbsp;million needles that were produced up to 1960 manufactured food for about 600,000&nbsp;mm of apical growth or 730 g dry weight, over 12&nbsp;million mm<sup>3</sup> of wood for the 1961 annual ring, plus 1&nbsp;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&nbsp;mg dry weight of apical growth, 3&nbsp;mm<sup>3</sup> wood, one-quarter of a new needle, plus an unknown amount of branch wood, bark and roots.
+Since conifers cannot regrow their leaves rapidly like hardwoods, leaf<!--foliar--> diseases can seriously damage coniferous plantations, especially dense stands of young trees. [[Needle cast]] diseases, often caused by [[Ascomycota|ascomycete fungi]] in the [[Rhytismataceae]] family, result in leaf fall.<ref name="Worrall 2025">{{cite web |last=Worrall |first=J. |title=Foliage Diseases |url=https://forestpathology.org/foliage/ |website=Forest Pathology |access-date=20 September 2025}}</ref> Another ascomycete, ''Rhizosphaera'' ([[Sphaeropsidales]]), causes severe defoliation and shoot blight, for instance in spruces.<ref>{{cite web |title=Rhizosphaera Needle Cast |url=https://hort.extension.wisc.edu/articles/rhizosphaera-needle-cast/ |publisher=[[University of Wisconsin–Madison]]: Wisconsin Horticulture |access-date=7 September 2025}}</ref>
-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),<ref name="Fraser-1964" /> the needles constituted 17.5% of the over-day weight. Undoubtedly, the proportions change with time.
+At least 20 species of roundheaded wood-boring [[longhorn beetle]]s (Cerambycidae) feed on the wood of spruces, firs, and hemlocks.<ref name="Rose1985">Rose, A.H.; Lindquist, O.H. 1985. Insects of eastern spruces, fir and, hemlock, revised edition. Government of Canada, [[Canadian Forest Service]], Ottawa, Forestry Technical Report 23.</ref> [[Bark beetle]]s (Scolytinae, in the [[Curculionidae]]) are destructive pests of commercial forestry; major pests of spruce and other conifers include ''[[Ips typographus]]'' in Eurasia<ref name="Hlasny-2019">{{cite book |last=Hlasny |first=Tomas |display-authors=etal |title=Living with bark beetles: impacts, outlook and management options |date=2019 |publisher=[[European Forest Institute]] |isbn=978-952-5980-75-2 |pages=8-11 |url=https://efi.int/sites/default/files/files/publication-bank/2019/efi_fstp_8_2019_0.pdf}}</ref> and ''[[Dendroctonus rufipennis]]'' in North America.<ref>http://www.na.fs.fed.us/spfo/pubs/fidls/sprucebeetle/sprucebeetle.htm {{Webarchive|url=https://web.archive.org/web/20150217073724/http://www.na.fs.fed.us/spfo/pubs/fidls/sprucebeetle/sprucebeetle.htm |date=2015-02-17 }} USFS Spruce Beetle</ref>
-===Seed-dispersal mechanism===
+The [[basidiomycete]] fungus ''[[Boletus pinophilus]]'' is among the fungi that form an [[ectomycorrhizal]] association with conifers, in its case with pines such as ''[[Pinus sylvestris]]''.<ref name="Gallardi-2020">{{cite book |last=Gallardi |first=Matteo |chapter=Diversity, Biogeographic Distribution, Ecology, and Ectomycorrhizal Relationships of the Edible Porcini Mushrooms (''Boletus'' s. str., Boletaceae) Worldwide: State of the Art and an Annotated Checklist |title=Mushrooms, Humans and Nature in a Changing World: Perspectives from Ecological, Agricultural and Social Sciences |editor=Pérez-Moreno, Jesús |editor2=Guerin-Laguette, Alexis |editor3=Arzú, Roberto Flores |editor4=Yu, Fu-Qiang |date=2020 |publisher=Springer |location=[[Cham, Switzerland]] |isbn=978-3-030-37378-8 |pages=236–237 |chapter-url=https://books.google.com/books?id=M3DgDwAAQBAJ&pg=PA237}}</ref>
-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 and long-distance dispersal. Long-distance dispersal distances range from {{convert|11.9|-|33.7|km}} from the source.<ref>{{cite journal|last1=Williams|first1=C.G.|last2=LaDeau|first2=S.L. |last3=Oren|first3=R. |last4=Katul|first4=G.G. |year=2006 |title=Modeling seed dispersal distances: implications for transgenic Pinus taeda |journal=Ecological Applications |volume=16|issue=1 |pages=117–124|doi=10.1890/04-1901 |pmid=16705965 |bibcode=2006EcoAp..16..117W }}</ref>
+[[Wilding conifer|Some conifers introduced for forestry]] including ''[[Pinus radiata]]'' have become [[invasive species]] in New Zealand,<ref name="NZDeptConservation 2001">{{cite web |url=http://csl.doc.govt.nz/publications/conservation/threats-and-impacts/weeds/south-island-wilding-conifer-strategy/ |title=South Island wilding conifer strategy |publisher=[[Department of Conservation (New Zealand)]] |year=2001 |access-date=19 April 2009 |archive-date=14 August 2011 |archive-url=https://web.archive.org/web/20110814032140/http://csl.doc.govt.nz/publications/conservation/threats-and-impacts/weeds/south-island-wilding-conifer-strategy/ }}</ref> South Africa,<ref>{{cite conference |url=http://invasive.org/publications/xsymposium/proceed/13pg941.pdf |title=Biological Control of Alien, Invasive Pine Trees (Pinus species) in South Africa |journal=The X International Symposium on Biological Control of Weeds |date=4–14 July 1999 |location=Montana State University, Bozeman, Montana |editor-first=Neal R.|editor-last=Spencer |pages=941–953 |first1=V. C. |last1=Moran |first2=J. H. |last2=Hoffmann |first3=D. |last3=Donnelly |first4=B. W. |last4=van Wilgen |first5=H. G. |last5=Zimmermann |access-date=28 June 2016 |archive-date=6 October 2016 |archive-url=https://web.archive.org/web/20161006014151/http://www.invasive.org/publications/xsymposium/proceed/13pg941.pdf|url-status=live}}</ref> and Australia.<ref name="Lindemayer 2007">{{cite journal |last1=Lindenmayer |first1=D. B. |last2=Hobbs |first2=R. J. |title=Fauna conservation in Australian plantation forests – a review |journal=[[Biological Conservation (journal)|Biological Conservation]] |date=September 2004 |volume=119 |issue=2 |pages=151–168 |doi=10.1016/j.biocon.2003.10.028 |bibcode=2004BCons.119..151L }}</ref><ref>{{cite web |url=https://keyserver.lucidcentral.org/weeds/data/media/Html/pinus_radiata.htm |title=Pinus radiata |publisher=keyserver.lucidcentral.org |work=Weeds of Australia |date=2016 |access-date=22 August 2018 |archive-date=19 June 2017 |archive-url=https://web.archive.org/web/20170619101113/http://keyserver.lucidcentral.org/weeds/data/media/Html/pinus_radiata.htm |url-status=live}}</ref>
-Birds of the crow family, [[Corvidae]], are the primary distributor of the conifer seeds. These birds are known to [[Hoarding (animal behavior)|cache]] 32,000 pine seeds and transport the seeds as far as {{convert|12|-|22|km|abbr=on}} from the source. The birds store the seeds in the soil at depths of {{convert|2|–|3|cm|abbr=on|frac=4}} under conditions which favor [[germination]].<ref>{{cite journal |last1=Tomback |first1=D. |author-link=Diana Tomback |first2=Y. |last2=Linhart |year=1990|title=The evolution of bird-dispersed pines |journal=Evolutionary Ecology |volume=4|issue=3|pages=185–219 |doi=10.1007/BF02214330 |bibcode=1990EvEco...4..185T }}</ref>
-==Distribution and habitat==
+<gallery class=center mode=nolines heights=180 widths=180>
+File:Siberian autumn in taiga..JPG|[[Taiga]] coniferous forest, mostly larches, pines, and spruces, covers a large area of [[Siberia]] (pictured) and Canada.<ref name="Berkeley"/>
+File:Kuuse-kooreürask ja tegutsemisjäljed Ips typographus.jpg|Galleries of ''[[Ips typographus]]'' bark beetles weaken conifers such as [[Picea abies|Norway spruce]], and can seriously harm commercial forestry.
+File:Boletus pinophilus3.JPG|The pine bolete ''[[Boletus pinophilus]]'' forms an [[ectomycorrhizal]] association with several pines.
+File:Prospect Pine Forest, Sydney.jpg|''[[Pinus radiata]]'' (radiata or Monterey pine) is an [[invasive species]] in Australia (pictured), New Zealand, and South Africa.
+</gallery>
-Conifers are the dominant plants over large areas of land, most notably the [[taiga]] of the [[Northern Hemisphere]],<ref name="Campbell-2005" /> but also in similar cool climates in mountains further south.
+== Economic importance ==
-==Ecology==
+{{further|Forestry|Silviculture}}
-=== As an invasive species ===
+The [[softwood]] derived from conifers is more easily worked than [[hardwood]] from broadleaved ([[angiosperm]]) trees. This makes it widely used and of great economic value, its many uses including construction, furniture, telegraph poles and fencing.<ref name="Edlin 1966">{{cite book |last=Edlin |first=Herbert L. |author-link=Herbert L. Edlin |title=Know Your Conifers: Forestry Commission Booklet No. 15 |date=1966 |publisher=[[HMSO]] |pages=5–6 |url=https://cdn.forestresearch.gov.uk/1966/03/fcbk015.pdf}}</ref> A large part of [[Pulp and paper industry|production is used for paper]].<ref name="Edlin 1966"/><ref>{{cite book |last1=Mleziva |first1=M. M. |last2=Wang |first2=J. H. |chapter=Paper |title=Polymer Science: A Comprehensive Reference |date=2012 |pages=397–410 |isbn=978-0-08-087862-1 |doi=10.1016/B978-0-444-53349-4.00274-0}}</ref> In the United Kingdom, the 48% of the woodland that is coniferous yields over 90% of the timber; the top species is [[sitka spruce]], yielding about half of the timber produced.<ref name="Willoughby 2025">{{cite journal |last1=Willoughby |first1=Ian H. |last2=Dhanda |first2=Rajni |last3=Clarke |first3=Toni |last4=Reynolds |first4=Chris |title=Seventeen coniferous tree species show early promise for future commercial timber production in the UK |journal=[[Forestry journal|Forestry]]: An International Journal of Forest Research |date=10 August 2025 |doi=10.1093/forestry/cpaf048 |doi-access=free}}</ref> Worldwide, wood products reached a value of $100 billion by the end of the 20th century.<ref name="Farjon 1999"/>
-{{Main|Wilding conifer}}
+{{Annotated image 4
-[[File:Prospect Pine Forest, Sydney.jpg|thumb|A Monterey pine forest in [[Sydney]], Australia]]
+| header          = Conifer [[wood]]
+| image           = 4 conifer wood samples.jpg
+| align           = center
+| image-width     = 400
+| width           = 400
+| height          = 120
+| annot-font-size = 12
+| annot-color     = f
+| annotations =
+{{Annotation| 35|90|&nbsp;[[Pine]]&nbsp;}}
+{{Annotation|125|90|&nbsp;[[Spruce]]&nbsp;}}
+{{Annotation|230|90|&nbsp;[[Larch]]&nbsp;}}
+{{Annotation|325|90|&nbsp;[[Juniper]]&nbsp;}}
+}}
-A number of conifers originally introduced for forestry have become [[invasive species]] in parts of [[New Zealand]], including radiata pine (''[[Pinus radiata]]''), lodgepole pine (''[[Pinus contorta|P. contorta]]''), [[Douglas fir]] (''Pseudotsuga mensiezii'') and European larch (''[[Larix decidua]]'').<ref name="NZDeptConservation-2001">{{cite web |url=http://csl.doc.govt.nz/publications/conservation/threats-and-impacts/weeds/south-island-wilding-conifer-strategy/ |title=South Island wilding conifer strategy |publisher=[[Department of Conservation (New Zealand)]] |year=2001 |access-date=2009-04-19 |archive-date=14 August 2011 |archive-url=https://web.archive.org/web/20110814032140/http://csl.doc.govt.nz/publications/conservation/threats-and-impacts/weeds/south-island-wilding-conifer-strategy/ |url-status=dead }}</ref>
+Conifers such as fir, cedar, cypress, juniper, spruce, pine, yew and [[Thuja|false cedar]] have been selected by plant breeders for ornamental purposes. Plants with unusual growth habits, sizes, and colours are propagated and planted in parks and gardens throughout the world.<ref name="Farjon-2010">{{cite book |last=Farjon |first=Aljos |author-link=Aljos Farjon |title=A Handbook of the World's Conifers |date=2010 |isbn=978-90-474-3062-9 |chapter=The economic importance of conifers |pages=25–29 |doi=10.1163/9789047430629}}</ref>
-In parts of [[South Africa]], maritime pine (''[[Pinus pinaster]]''), patula pine (''[[Pinus patula|P. patula]]'') and radiata pine have been declared invasive species.<ref>{{cite conference|url = http://invasive.org/publications/xsymposium/proceed/13pg941.pdf|title = Biological Control of Alien, Invasive Pine Trees (Pinus species) in South Africa|journal = The X International Symposium on Biological Control of Weeds|date = 4–14 July 1999|location = Montana State University, Bozeman, Montana, USA|editor-first = Neal R.|editor-last = Spencer|pages = 941–953|first1 = V. C.|last1 = Moran|first2 = J. H.|last2 = Hoffmann|first3 = D.|last3 = Donnelly|first4 = B. W.|last4 = van Wilgen|first5 = H. G.|last5 = Zimmermann|access-date = 28 June 2016|archive-date = 6 October 2016|archive-url = https://web.archive.org/web/20161006014151/http://www.invasive.org/publications/xsymposium/proceed/13pg941.pdf|url-status = live}}</ref> These [[wilding conifers]] are a serious environmental issue causing problems for pastoral farming and for [[conservation (ethic)|conservation]].<ref name="NZDeptConservation-2001"/>
+<gallery class=center mode=nolines heights=180 widths=240>
+File:Young Sitka spruce plantation - geograph.org.uk - 949091.jpg|[[Forestry|Commercial forestry]] using [[sitka spruce]]
+File:2016.04.12 18.14.33 DSC03322 - Flickr - andrey zharkikh.jpg|''Globosa'', an ornamental [[cultivar]] of Scots pine
+File:JBP_Kotobuki.jpg|''[[Pinus thunbergii]]'' 'Kotobuki'<br/>as a 65-year-old [[bonsai]]
+</gallery>
-Radiata pine was introduced to Australia in the 1870s. It is "the dominant tree species in the Australian plantation estate"<ref name="Lindemayer-2007">{{cite journal |last1=Lindenmayer |first1=D.B. |last2=Hobbs |first2=R.J. |title=Fauna conservation in Australian plantation forests – a review |journal=Biological Conservation |date=September 2004 |volume=119 |issue=2 |pages=151–168 |doi=10.1016/j.biocon.2003.10.028 |bibcode=2004BCons.119..151L }}</ref> – 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<ref>{{cite web|url=https://keyserver.lucidcentral.org/weeds/data/media/Html/pinus_radiata.htm|title=Pinus radiata|publisher=keyserver.lucidcentral.org|work=Weeds of Australia|date=2016|access-date=22 August 2018|archive-date=19 June 2017|archive-url=https://web.archive.org/web/20170619101113/http://keyserver.lucidcentral.org/weeds/data/media/Html/pinus_radiata.htm|url-status=live}}</ref> and the removal of individual plants beyond plantations is encouraged.<ref>{{cite web |url=http://www.bmcc.nsw.gov.au/sustainableliving/weedmanagement/factsheets |title=Blue Mountains City Council – Fact Sheets &#91;Retrieved 1 August 2015&#93;|access-date=22 August 2018|archive-date=24 June 2015|archive-url=https://web.archive.org/web/20150624041550/http://www.bmcc.nsw.gov.au/sustainableliving/weedmanagement/factsheets|url-status=dead}}</ref>
+== References ==
-=== Predators ===
+{{reflist}}
-At least 20 species of roundheaded borers of the family [[Longhorn beetle|Cerambycidae]] feed on the wood of [[spruce]], [[fir]], and [[Tsuga|hemlock]] (Rose and Lindquist 1985).<ref name="Rose1985">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)</ref> 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 [[Monochamus scutellatus|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 two weeks and the tiny [[larva]]e 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 tunnelling 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&nbsp;mm long, [[pupa]]te 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.
+== External links ==
-==Cultivation==
-{{See also|Silviculture}}
-[[File:2016.04.12 18.14.33 DSC03322 - Flickr - andrey zharkikh.jpg|thumb|''Globosa'', a [[cultivar]] of ''[[Pinus sylvestris]]'', a northern European species, in the North American [[Red Butte Garden]]]]
-Conifers – notably ''[[Abies]]'' (fir), ''[[Cedrus]]'', ''[[Chamaecyparis lawsoniana]]'' (Lawson's cypress), ''[[Cupressus]]'' (cypress), [[juniper]], ''[[Picea]]'' (spruce), ''[[Pinus]]'' (pine), ''[[Taxus]]'' (yew), ''[[Thuja]]'' (cedar)&nbsp;– have been the subject of selection for ornamental purposes. Plants with unusual growth habits, sizes, and colours are propagated and planted in parks and gardens throughout the world.<ref name="Farjon-2010">{{cite book |doi=10.1163/9789047430629 |title=A Handbook of the World's Conifers |date=2010 |last1=Farjon |first1=Aljos |isbn=978-90-474-3062-9 }}{{pn|date=January 2025}}</ref>
-==Conditions for growth==
-Conifers [[Plant nutrition |can absorb nitrogen]] in either the [[ammonium]] (NH<sub>4</sub><sup>+</sup>) or [[nitrate]] (NO<sub>3</sub><sup>−</sup>) 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).<ref name="Durzan-1967">{{cite journal |last1=Durzan |first1=D.J. |last2=Steward |first2=F.C. |year=1967 |title=The nitrogen metabolism of ''Picea glauca'' (Moench) Voss and ''Pinus banksiana'' Lamb. as influenced by mineral nutrition |journal=Can. J. Bot. |volume=45 |issue=5  |pages=695–710 |doi=10.1139/b67-077  |bibcode=1967CaJB...45..695D }}</ref> Ammonium nitrogen was shown to foster [[arginine]] and [[amide]]s and lead to a large increase of free [[guanidine]] 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 a higher nitrogen content after 5 weeks<ref name="McFee-1968">{{cite journal |last1=McFee |first1=W.W. |last2=Stone |first2=E.L. |year=1968 |title=Ammonium and nitrate as nitrogen sources for ''Pinus radiata ''and ''Picea glauca'' |journal=Soil Sci. Soc. Am. Proc. |volume=32 |issue=6 |pages=879–884 |doi=10.2136/sssaj1968.03615995003200060045x  |bibcode=1968SSASJ..32..879M }}</ref> than did the same amount of nitrate nitrogen. Swan  found the same effect in 105-day-old white spruce.<ref name="Swan-1960">{{cite report |last=Swan |first=H.S.D. |year=1960 |title=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 |publisher=Pulp Paper Res. Instit. Can. |location=Montreal QC |series=Woodlands Res. Index Number 116 |id=Tech. Rep. 168}}</ref>
-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).<ref name="Armson-1961">{{cite book |last1=Armson |first1=KA |last2=Carman |first2=RD |year=1961 |title=Forest tree nursery soil management |publisher=Ont. Dep. Lands & Forests, Timber Branch |location=Ottawa ON}}</ref> Over a longer period, root growth is also stimulated. Many [[Plant nursery |nursery]] managers were long reluctant to apply nitrogenous [[fertilizer]]s 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 [[Picea abies |Norway spruce]] and white spruce in his nursery operation by fertilizing with 50–80&nbsp;lb/ac (56–90&nbsp;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.<ref name="Eastman-1980">{{cite conference |last=Eastman |first=B |title=The Western Maine Forest Nursery Company |pages=291–295 |book-title=Proc. of the North American Forest Tree Nursery Soils Workshop |date=July 28 – August 1, 1980 |location=Syracuse, New York |publisher=Environment Canada, Canadian Forestry Service, USDA For. Serv.}}</ref>
-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]], [[magnesium]], and [[sulfur]], all used in relatively large amounts.<ref name="Buckman-1969">{{cite book |last1=Buckman |first1=H.O. |last2=Brady |first2=N.C. |year=1969 |title=The Nature and Properties of Soils |edition=7th |publisher=Macmillan |location=New York}}</ref>
-==Economic importance==
-The [[softwood]] derived from conifers is of great economic value, providing about 45% of the world's annual lumber production.{{fact|date=January 2025}} Other uses of the timber include the [[Pulp and paper industry|production of paper]]<ref>{{cite book |doi=10.1016/B978-0-444-53349-4.00274-0 |quote=There are generally two types of trees used in papermaking: softwoods or coniferous woods (gymnosperms) and hardwoods (angiosperms). |chapter=Paper |title=Polymer Science: A Comprehensive Reference |date=2012 |last1=Mleziva |first1=M.M. |last2=Wang |first2=J.H. |pages=397–410 |isbn=978-0-08-087862-1 }}</ref> and plastic from chemically treated wood pulp. Some conifers also provide foods such as [[pine nuts]] and [[juniper berries]], the latter used to flavor [[gin]].
-==References==
-{{Reflist}}
-{{notelist}}
-==External links==
 {{Commons category}}
 {{Wikispecies|Pinophyta}}
 * [http://tolweb.org/tree?group=Conifers&contgroup=Spermatopsida Conifers] at the Tree of Life Web Project
-* [http://www.livescience.com/animals/070504_chicago_cave.html 300 million-year-old conifer in Illinois – 4/2007]
+* [https://s10.lite.msu.edu/res/msu/botonl/b_online/library/knee/hcs300/gymno.htm Gymnosperms] at Michigan State University
-* [http://www.catalogueoflife.org/col/browse/classification/order/Pinales/fossil/0/match/1 World list of conifer species from Conifer Database by A. Farjon] in the [[Catalogue of Life]] ({{Webarchive|url=https://web.archive.org/web/20170619200712/http://www.catalogueoflife.org/col/browse/classification/order/Pinales/fossil/0/match/1 |date=2017-06-19 }})
+* [https://herbaria.plants.ox.ac.uk/bol/conifers Conifers of the World: Resources for Conifer Research] - some 37,000 herbarium records
-* [http://www.catalogueoflife.org/col/browse/tree/id/844f3ce878b11544be19b2da1c2f03dc Tree browser for conifer families and genera via the Catalogue of Life] ({{Webarchive|url=https://web.archive.org/web/20191220201038/http://www.catalogueoflife.org/col/browse/tree/id/844f3ce878b11544be19b2da1c2f03dc |date=2019-12-20 }})
-* [https://web.archive.org/web/20121113094003/http://www.conifer-encyclopedia.com/ ''Royal Horticultural Society Encyclopedia of Conifers: A Comprehensive Guide to Cultivars and Species'']
-* [http://conifersaroundtheworld.com DendroPress: Conifers Around the World].
-* {{cite web |last=Knee |first= Michael |title=Gymnosperms |access-date=14 January 2016 |url=https://s10.lite.msu.edu/res/msu/botonl/b_online/library/knee/hcs300/gymno.htm}}
 {{Pinophyta}}
 {{Plant classification}}
 {{Life on Earth}}
 {{Taxonbar|from=Q132825}}
 {{Authority control}}

Conifer: Difference between revisions

Latest revision as of 21:26, 29 October 2025

Evolution

Fossil history

Relict species

External phylogeny

Internal phylogeny

Taxonomy

Description

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Distribution and ecology

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