Conifer

{{Short description|Group of seed plants}} {{Other uses}} {{good article}} {{Use dmy dates|date=June 2022}} {{Automatic taxobox | fossil_range = {{fossil range|307|0}}Carboniferous–Present | image = Sapins pectinés.jpg | image_alt = Conifer forest | image_caption = Large conifer forest of silver fir (''Abies alba'') at Vosges, Eastern France | display_parents = 2 | taxon = Pinopsida | authority = | subdivision_ranks = Subclasses, orders, and families | subdivision = * Cupressidae ** Araucariales *** Araucariaceae *** Podocarpaceae ** Cupressales *** Sciadopityaceae *** Cupressaceae *** Taxaceae * Pinidae ** Pinales *** Pinaceae *** Cheirolepidiaceae † *** Arctopityaceae † * Gnetidae<ref>{{cite journal |last=Yang|first=Yong|last2=Ferguson|first2=David Kay|last3=Liu|first3=Bing|last4=Mao|first4=Kang-Shan|last5=Gao|first5=Lian-Ming|last6=Zhang|first6=Shou-Zhou|last7=Wan|first7=Tao|last8=Rushforth|first8=Keith|last9=Zhang|first9=Zhi-Xiang|title=Recent advances on phylogenomics of gymnosperms and a new classification|journal=Plant Diversity|volume=44|issue=4|year=2022|pages=340-350|url=https://www.sciencedirect.com/science/article/pii/S2468265922000440}}</ref> ** Gnetales *** Gnetaceae ** Welwitschiales *** Welwitschiaceae ** Ephedrales *** Ephedraceae * Palissyales † * Voltziales † * Cordaitales † | synonyms = * Coniferophyta * Coniferae * Pinophytina }}

'''Conifers''' ({{IPAc-en|ˈ|k|ɒ|n|ɪ|f|ər}}) are a group of vascular plants and a subset of gymnosperms. They are primarily perennial, woody trees and shrubs, mostly evergreen 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. Taxonomically, 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 minimize water loss. Several fungi form ectomycorrhizal associations with conifers, while other fungi cause diseases such as needle cast, which is 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. As of 2025, 94 conifer species are listed as endangered and 30 as critically endangered. Conifers are of great economic value for timber and paper production.

== 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.<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 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 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> 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 minimize 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>

<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 boreal conifers, and their downward-drooping limbs, help them shed snow.<ref name="Michigan"/> </gallery>

=== Foliage ===

Most conifers are evergreens, retaining functional foliage throughout the year.<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 cedars, the leaves are long, thin and needle-like. Others like 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 decussate (crossing) 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 flat plane for maximum light absorption. Leaf size varies from 2 mm in many scale-leaved species, up to {{convert|600|mm|abbr=on}} long in the needles of some pines (e.g. longleaf pine, ponderosa pine). 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 color, 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 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.<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>

<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 absorption. File:C lawsoniana Lge.jpg|Cupressaceae: scale leaves of Lawson's cypress (''Chamaecyparis lawsoniana''); ruler in mm </gallery>

=== Wood ===

The microscopic structure of conifer wood (xylem) is homogeneous,<ref name="softwood anatomy">{{cite journal |last1=Esteban |first1=Luis G. |last2=de Palacios |first2=Paloma |last3=Heinz |first3=Immo |last4=Gasson |first4=Peter |last5=García-Iruela |first5=Alberto |last6=García-Fernández |first6=Francisco |year=2023 |title=Softwood Anatomy: A Review |journal=Forests |volume=14|issue=2|url=https://doi.org/10.3390/f14020323}}</ref> and 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.<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>

Conifers produce growth rings, normally gently curved in their outline, but in Taxus, Juniperus, and Cupressaceae are undulating.<ref name="softwood anatomy"/> 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.<ref name="Ledig-1982"/> Frost rings have been observed in Cupressaceae.<ref name="softwood anatomy"/>

Conifer wood contains medullary rays, primarily composed of parenchymal cells, and involved in radial gas exchange, as a conduit for storage of water and nutrients, and as a pathway for diffusing heartwood substances.<ref name="softwood anatomy"/> The latewood in particular also contains resin canals. These are surrounded by specialised epithelial cells that secrete resin into the canal. Radial and axial resin canals are interconnected throughout the wood.<ref>{{cite journal |last=Koddenberg |first=T. |title=A new dimension in wood anatomy education: exploring softwood and hardwood structures in 3D |journal=European Journal of Wood and Wood Products |volume=83 |page=201 |date=6 December 2025 |doi=10.1007/s00107-025-02329-6}}</ref>

Conifers secrete oleoresin, a composite of turpentine and rosin. When an insect or a fungus attacks, oleoresin flows into the wound. It traps invading insects and blocks fungi. The rosin hardens and seals the wound, protecting against damage.<ref>{{cite journal |last1=Trapp |first1=Susan |last2=Croteau |first2=Rodney |title=Defensive Resin Biosynthesis in Conifers |journal=Annual Review of Plant Physiology and Plant Molecular Biology |volume=52 |pages=689–724 |year=2001 |doi=10.1146/annurev.arplant.52.1.689}}</ref> During wet weather, the Gitxsan harvest conifer rosin for use as fire starters.<ref>{{cite journal|last1=Turner|first1=Nancy J.|last2=Ari|first2=Yilmaz|last3=Berkes|first3=Fikret|last4=Davidson-Hunt|first4=Iain|last5=Ertug|first5=Z. Fusun|last6=Miller|first6=Andrew|title=Cultural management of living trees: an international perspective|journal=Journal of Ethnobiology|volume=29|issue=2|year=2009|pages=237-270}}</ref>

<gallery class=center mode=nolines widths=180 heights=180> File:Abies concolor tangential.jpg|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. 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 rings of fast (big cells, earlywood) and slow seasonal growth File:Resin Canals.png|Resin canals appear as white circles in a section of pine wood. </gallery>

=== Roots === Saplings initially develop a primary taproot, while mature conifers develop an extensive network of coarse roots for mechanical support.<ref>{{cite journal|last1=Dūmiņš|first1=Kārlis|last2=Žīgure|first2=Sindija|last3=Celma|first3=Santa|last4=Štāls|first4=Toms Artūrs|last5=Vendiņa|first5=Viktorija|last6=Zuševica|first6=Austra|last7=Lazdiņa|first7=Dagnija|title=Impact of Soil Preparation Method and Stock Type on Root Architecture of Scots Pine, Norway Spruce, Silver Birch and Black Alder|journal=Forests|volume=16|issue=5|pages=830|year=2025|publisher=Latvian State Forest Research Institute SILAVA|doi=10.3390/f16050830|doi-access=free}}</ref> Structural integrity and failure is strongly influenced by the radial symmetry of the root system, spoke-like root systems are more stable than asymmetrical supports, especially in shallower soils.<ref>{{cite journal|last1=Coutts|first1=M. P.|last2=Nielsen|first2=C. C. N.|last3=Nicoll|first3=B. C.|title=The development of symmetry, rigidity and anchorage in the structural root system of conifers|journal=Plant and Soil|volume=217|pages=1–15|year=1999|publisher=Kluwer Academic Publishers|location=Netherlands|doi=10.1023/A:1004578032481}}</ref>

Conifers also develop a near-surface fine root system, concentrated on lateral branches arising from the course roots, which are colonized by mycorrhizal fungi that enhance water and mineral absorption.<ref>{{cite journal|last1=Pregitzer|first1=Kurt S.|last2=DeForest|first2=Jared L.|last3=Burton|first3=Andrew J.|last4=Allen|first4=Michael F.|last5=Ruess|first5=Roger W.|last6=Hendrick|first6=Ronald L.|title=Fine root architecture of nine North American trees|journal=Ecological Monographs|volume=72|issue=2|pages=293–309|year=2002|publisher=Southern Research Station|doi=10.1890/0012-9615(2002)072[0293:fraonn]2.0.co;2}}</ref>

=== Reproduction ===

Conifers produce their seeds inside a protective cone called a strobilus. Most species are monoecious, with male and female cones on the same tree.<ref name="Principles of Horticulture"/><ref name="Treeguide"/> All conifers are wind-pollinated.<ref>{{cite journal |last=Owens |first=J. N. |year=1998 |title=Pollination in conifers |journal=Trends in Ecology & Evolution |volume=13 |issue=12 |pages=… |doi=10.1016/S1360-1385(98)01337-5 |bibcode=1998TPS.....3..479O |url=https://www.sciencedirect.com/science/article/pii/S1360138598013375 |access-date=25 January 2026}}</ref> 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.<ref name="Principles of Horticulture">{{cite book |last1=Adams |first1=Charles R. |last2=Bamford |first2=Katherine M. |last3=Early |first3=Mike P. |title=Principles of Horticulture |edition=6th |publisher=Routledge |year=2012 |doi=10.4324/9780080969589 |isbn=978-0080969572}}</ref><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 nuts, which are dispersed by birds, in particular, nutcrackers, and jays, 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=978-0-226-38263-0 |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 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 arils, resembling berries. These are eaten by 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>

<gallery class=center mode=nolines widths=180 heights=180> File:Abies lasiocarpa 6972.JPG|Pinaceae: unopened female cones of subalpine fir (''Abies lasiocarpa'') 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 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 ===

[[File:Gymnosperm life cycle diagram-en.svg|thumb|upright=1.35|Life cycle of a pine tree]]

Conifers are heterosporous, generating two different types of spores: male microspores and female megaspores.<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}}

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}}

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.{{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 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}}

== Evolution ==

=== Fossil history ===

The earliest conifers appear in the fossil record during the Late Carboniferous (Pennsylvanian) over 300 million years ago.<ref name="Feng-2017"/><ref name="Leslie Beaulieu 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 |title=An overview of extant conifer evolution from the perspective of the fossil record |journal=American Journal of Botany |volume=105 |issue=10 |pages=1531–1544 |year=2018 |doi=10.1002/ajb2.1143 |pmid=30157290 |bibcode=2018AmJB..105.1531L }}</ref> 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.<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 |bibcode=2018AmJB..105.1531L }}</ref> Conifers underwent a major decline in the Late Cretaceous corresponding to the explosive adaptive radiation of flowering plants.<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 laxifolia'' foliage, Early Permian, 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>

=== 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.<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 bioRxiv |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 |biorxiv=10.1101/2023.08.24.554647 }}</ref> 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.<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 |bibcode=2022FrEEv..1082639C }}</ref> 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.<ref name="Farjon 1999"/>

<gallery class=center mode=nolines widths=180 heights=180> 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 gnetophytes, 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 fossils from the Late Cretaceous onwards. </gallery>

=== External phylogeny ===

The cladogram summarizes the group's external phylogeny. The conifers are gymnosperms, sister to a clade consisting of the ginkgos and cycads.<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 |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 |bibcode=2018AmJB..105.1531L |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 |bibcode=2018AmJB..105.1531L |s2cid=52120430}}</ref><ref name="Stull 2021">{{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 |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=Gymnosperms |1={{clade |1={{clade |1=Ginkgoidae 60px |2=Cycadidae 60px }} |2='''Conifers''' 60px }} }}

=== Internal phylogeny ===

The Gnetophyta, despite distinct appearances, were long viewed as outside the conifer group, but phylogenomic analysis indicates that the group is 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 |bibcode=2018PBioS.28581012R }}</ref>

{{clade|style=line-height:100%; |label1='''Pinophyta''' |sublabel1=(Coniferae) |1={{clade |1={{clade |1={{clade |label1=Pinaceae |sublabel1=pine family |1=60px |label2=Gnetophyta |sublabel2=(3 relict genera) |2=60px }} |2={{clade |1={{clade |label1=Araucariaceae |sublabel1=monkey puzzle family |1=60px |label2=Podocarpaceae |sublabel2=podocarps |2=60px }} |2={{clade |label1=Sciadopityaceae |sublabel1=umbrella pines |1=60px |2={{clade |label1=Cupressaceae |sublabel1=cypress family |1=60px |2={{clade |label1=Cephalotaxaceae |sublabel1=plum yew family |1=50px |label2=Taxaceae |sublabel2=yew family |2=60px }} }} }} }} }} }} }}

=== Taxonomy ===

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"/> comparable to the Rosaceae, he doubted that the conifers or the gymnosperms formed natural groups (clades).<ref name="Farjon 1999"/> By 2016, the conifers were recognized 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 |pages=67–96}}</ref> 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.<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>

== Distribution and ecology ==

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 northern-ranging tree species, at 75° north in the Taymyr Peninsula.<ref name="Farjon 1999"/> Conifers are widespread also in southern Europe, Western Asia, 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 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 |doi=10.1111/1467-8748.00215 |jstor=45065379}}</ref>

Conifers are adapted to acidic, nutrient-poor soils, low temperatures, and seasonal water limitations. Their ecology is strongly shaped by subsurface interactions within the rhizosphere. In these environments, conifers associate with ectomycorrhizal fungi, forming symbioses that help them to acquire nutrients. Alongside fungi, plant growth–promoting rhizobacteria are key members of the rhizosphere microbial community, contributing to nutrient cycling and plant health.<ref>{{cite book |last1=Cardoso |first1=E. J. |last2=Vasconcellos |first2=R. L. |last3=Ribeiro |first3=C. M. |last4=Miyauchi |first4=M. Y. |chapter=PGPR in coniferous trees |title=Bacteria in Agrobiology: Crop Ecosystems |year=2011 |publisher=Springer |location=Berlin, Heidelberg |pages=345–359 |url=https://www.researchgate.net/publication/341179236_Bacteria_in_Agrobiology_Crop_Ecosystems#page=359 |isbn=978-3-642-18356-0}}</ref>

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.<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>

At least 20 species of roundheaded wood-boring longhorn beetles (Cerambycidae) feed on the wood of spruces, firs, and hemlocks.<ref name="Rose 1985">{{cite report |last1=Rose |first1=Arthur H. |last2=Lindquist |first2=O. H. |department=Natural Resources Canada, Canadian Forest Service |title=Insects of Eastern Spruces, Fir and Hemlock |year=1994 |url=https://publications.gc.ca/collections/collection_2014/rncan-nrcan/Fo64-23-1994-eng.pdf |access-date=19 February 2026 |publisher=Government of Canada |series=Cat. No. Fo64‑23‑1994‑eng |isbn=0-660-15112-X }}</ref> Bark beetles (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>{{cite web |title=USFS Spruce Beetle |url=http://www.na.fs.fed.us/spfo/pubs/fidls/sprucebeetle/sprucebeetle.htm |archive-url=https://web.archive.org/web/20150217073724/http://www.na.fs.fed.us/spfo/pubs/fidls/sprucebeetle/sprucebeetle.htm |archive-date=17 February 2015 |access-date=20 February 2026}}</ref>

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>

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 |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>

<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 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>

== Interaction with humans ==

=== Conservation ===

As of 2025, 83 species of conifer are listed as vulnerable, 94 are endangered, and 30 are critically endangered.<ref>{{cite web |title=IUCN Red List: Search for "Conifers" |url=https://www.iucnredlist.org/search?query=Conifers&searchType=species |publisher=IUCN |access-date=21 February 2026}}</ref> Among the most endangered species is ''Abies beshanzuensis'', the Baishan fir, which is at risk from flooding; only some 600 individual trees remain in the wild in Southeast China.<ref>{{cite web |title=Threats: Extreme weather events |url=https://threatenedconifers.rbge.org.uk/threats/extreme-weather-events |publisher=Royal Botanic Garden Edinburgh |access-date=19 February 2026}}</ref> The causes of decline of other species include habitat loss through agriculture and land development, over-exploitation, habitat degradation, invasive species, and climate change.<ref>{{cite book |last1=Farjon |first1=Aljos |last2=Page |first2=Christopher N. |title=Conifers: Status Survey and Conservation Action Plan |publisher=IUCN |year=1999 |series=IUCN/SSC Conifer Specialist Group |isbn=2-8317-0465-0 |url=https://portals.iucn.org/library/node/7565}}</ref>

=== Economic importance ===

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 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: An International Journal of Forest Research |date=10 August 2025 |article-number=cpaf048 |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"/> {{Annotated image 4 |header = Conifer wood cross sections |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| Pine }} {{Annotation|125|90| Spruce }} {{Annotation|230|90| Larch }} {{Annotation|325|90| Juniper }} }}

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.<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>

<gallery class=center mode=nolines heights=180 widths=240> File:Young Sitka spruce plantation - geograph.org.uk - 949091.jpg|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>

Conifers provide numerous non-wood products including ornamental and cultural items such as Christmas trees, bonsai, and topiary; foliage and wreaths for decorative greenery, mulch, and craft materials; and bark and roots traditionally used for food, medicine, natural dyes, and niche uses for compounds like taxol. Resins tapped from conifers yield commercial products such as turpentine and rosin, while essential oils extracted from foliage or wood are used in fragrances and industrial applications. Seeds, fruits, and cones include edible pine nuts and juniper berries used as a spice.<ref>{{cite book |last=Ciesla |first=William M. |title=Non-wood Forest Products from Conifers |year=1998 |publisher=Food and Agriculture Organization |isbn=92-5-104212-8 |series=Non-Wood Forest Products |number=37 |url=https://www.fao.org/4/x0453e/x0453e.pdf}}</ref><ref>{{cite web |last=Alexander |first=Rebecca |title=Edible Uses of Conifers |url=https://arboretumfoundation.org/wp-content/uploads/alexander_edible-conifers.pdf |publisher=Arboretum Foundation |work=Washington Park Arboretum Bulletin |access-date=19 February 2026}}</ref>

=== As food and drink ===

The Sámi and indigenous North American peoples used to eat conifer cambium. They peeled the bark and removed, dried, and ground the cambium into flour.<ref>{{cite journal |last1=Östlund |first1=Lars |last2=Ahlberg |first2=Lisa |last3=Zackrisson |first3=Olle |last4=Bergman |first4=Ingela |last5=Arno |first5=Steve |title=Bark-Peeling, Food Stress and Tree Spirits – The Use of Pine Inner Bark for Food in Scandinavia and North America |journal=Journal of Ethnobiology |volume=29 |issue=1 |year=2009 |pages=94–112 |doi=10.2993/0278-0771-29.1.94}}</ref>

Spring tips of pine, fir, and spruce have been used to flavour beer in the absence of hops.<ref>{{cite book |editor-last=Oliver |editor-first=Garrett |title=The Oxford Companion to Beer |publisher=Oxford University Press |year=2011 |isbn=978-0-19-536713-3 |entry=Pine, fir, and spruce tips}}</ref> Spruce beer was common in the colonial United States and eastern Canada, made from red or black spruce.<ref>{{cite book |title=Wines & beers of old New England: a how-to-do-it history |author=Sanborn Conner Brown |author2=Ed Lindlof |author3=Martin Kaufman |others=Ed Lindlof (illus.) |publisher=UPNE |year=1978 |isbn=0-87451-148-8 |page=[https://archive.org/details/universityofverm00kauf/page/67 67] |url-access=registration |url=https://archive.org/details/universityofverm00kauf/page/67 }}</ref> Crush produced spruce-beer soda until 2000.<ref>{{cite magazine |last=Faber |first=Rachel |title=A tall, cool glass of spruce |magazine=Maclean's |volume=128 |issue=37 |date=21 September 2015 |page=55}}</ref> Sahti, a beer using juniper, is brewed in rural areas in Finland.<ref>{{cite journal |last1=Ekberg |first1=Jukka |last2=Gibson |first2=Brian |last3=Joensuu |first3=Jussi J. |last4=Krogerus |first4=Kristoffer |last5=Magalhães |first5=Frederico |last6=Mikkelson |first6=Atte |last7=Seppänen-Laakso |first7=Tuulikki |last8=Wilpola |first8=Arvi |title=Physicochemical characterization of sahti, 'ancient' beer style indigenous to Finland |journal=Journal of the Institute of Brewing |date=11 September 2015 |doi=10.1002/jib.246 }}</ref>

The Abenaki in North America drank pine needle tea with teaberry and honey,<ref>{{cite journal |last=Benner |first=D. |title=Forage in Winter: Don’t let the snow stop you from enjoying the fresh food nature has to offer |journal=Mother Earth News |year=2025 |issue=333 |pages=27–29 |url=https://search.ebscohost.com/login.aspx?direct=true&db=f6h&AN=188868783&site=eds-live&scope=site |access-date=2026-02-21}}</ref> while in winter, the Iroquois made hemlock tea by boiling the past season's growth.<ref>{{cite book |last1=Kuhnlein |first1=Harriet |last2=Turner |first2=Nancy J. |title=Traditional Plant Foods of Canadian Indigenous Peoples: Nutrition, Botany and Use |publisher=Routledge |year=2020 |url=https://www.scribd.com/doc/127950233/Traditional-Plant-Foods-of-Canadian-Indigenous-Peoples-Nutrition-Botany-and-Use}}</ref> In Korea, Solip-cha is a needle tea made mostly from Korean red pine or Manchurian red pine.<ref name="Jeong">{{cite book |title=Cha Saenghwal Munhwa Daejeon |trans-title=Tea Life Culture Festival |publisher=Hong Ik Jae |year=2012 |isbn=9788971433515 |editor-last=정 |editor-first=동효 |location=Seoul |language=ko |script-title=ko:차생활문화대전 |chapter=sollip-cha (Pine Needle Tea) |script-chapter=ko:솔잎차 |access-date=9 June 2017 |editor-last2=윤 |editor-first2=백현 |editor-last3=이 |editor-first3=영희 |chapter-url=http://terms.naver.com/entry.nhn?docId=1635012&cid=48181&categoryId=48262 |via=Naver}}</ref>

=== In culture ===

Across many cultures, coniferous traits such as being evergreen, longevity, and endurance provide metaphors for elevation, immortality, and community resilience. Their persistent greenery has served as an emblem of life's continuity, to express intangible human values, and link the physical world to spiritual and cosmological ideas. Within Iroquois tradition, the eastern white pine was elevated to the ''Tree of Peace'', symbolizing the unity and enduring harmony of the Haudenosaunee Confederacy. Such an interpretation resonates with mythological motifs such as the World Tree or Tree of Life that represent unity, the sacred, and the continuity of life in diverse cultural contexts.<ref name="tree of peace">{{cite report |last=Schroeder |first=Herbert W. |title=The tree of peace: Symbolic and spiritual values of the white pine |publisher=USDA Forest Service, Northern Research Station |type=Symposium proceedings |pages=73–83 |location=Duluth, Minnesota |date=16–18 September 1992 |url=https://research.fs.usda.gov/treesearch/13474/ |access-date=7 February 2026}}</ref>

== References ==

== External links ==

* [http://tolweb.org/tree?group=Conifers&contgroup=Spermatopsida Conifers] at the Tree of Life Web Project * [https://s10.lite.msu.edu/res/msu/botonl/b_online/library/knee/hcs300/gymno.htm Gymnosperms] at Michigan State University * [https://herbaria.plants.ox.ac.uk/bol/conifers Conifers of the World: Resources for Conifer Research] - some 37,000 herbarium records

Category:Conifers Category:Extant Pennsylvanian first appearances Category:Plant divisions