View source for Gymnosperm

{{Short description|Clade of non-flowering, naked-seeded vascular plants}}
{{Distinguish|text=the flowering plant genera ''[[Gymnosperma]]'' and ''[[Gymnospermium]]''}}
{{Automatic taxobox
| name =
| fossil_range = {{fossil range|Bashkirian|0|earliest=320|[[Carboniferous]]–[[Holocene|Present]]}}
| image = Gymnospermae.jpg
| image_caption = Various gymnosperms.
| taxon = Gymnosperms
| authority =
| subdivision_ranks = Living orders<ref name="Phylogenomics of gymnosperms" />
| subdivision = *[[Cycadopsida]]
** [[Cycadales]]
* [[Ginkgoopsida]]
** [[Ginkgoales]]
* [[Pinopsida]]
** [[Cupressidae]]
*** [[Araucariales]]
*** [[Cupressales]]
** [[Pinidae]]
*** [[Pinales]]
* [[Gnetopsida]]
** [[Ephedrales]]
** [[Welwitschiales]]
** [[Gnetales]]
}}

The '''gymnosperms''' ({{IPAc-en|'|dʒ|ɪ|m|n|ə|ˌ|s|p|ɜːr|m|z|,_|-|n|oʊ|-|audio=en-us-gymnosperm.ogg}} {{respell||nə|spurmz|,_|-|noh|-}}; from [[Ancient Greek]] γυμνός (''gumnós''), meaning "naked", and σπέρμα (''spérma''), meaning "seed", and thus, "naked seed") are a group of woody, perennial [[Seed plant|seed-producing plants]], typically lacking the protective outer covering which surrounds the seeds in flowering plants, that include [[Pinophyta|conifers]], [[cycad]]s, [[Ginkgo]], and [[gnetophyta|gnetophyte]]s, forming the clade '''Gymnospermae'''.<ref name="DK Pub.-2012">{{Cite book |title=The Ultimate Visual Family Dictionary |publisher=[[DK Pub.]] |year=2012 |isbn=978-0-1434-1954-9 |location=New Delhi |pages=122–125 |language=en}}</ref> The name is based on the unenclosed condition of their seeds (called [[ovule]]s in their unfertilized state). The non-encased condition of their seeds contrasts with the seeds and ovules of flowering plants ([[angiosperms]]), which are enclosed within an [[Ovary (botany)|ovary]]. Gymnosperm seeds develop either on the surface of scales or [[Leaf|leaves]], which are often modified to form [[Conifer cone|cones]], or on their own as in [[Taxus|yew]], ''[[Torreya]]'', and ''[[Ginkgo]]''.

The life cycle of a gymnosperm involves [[alternation of generations]], with a dominant [[diploid]] [[sporophyte]] phase, and a reduced [[haploid]] [[gametophyte]] phase, which is dependent on the sporophytic phase.<ref>{{Cite journal |last1=Pandey |first1=Saurabh |last2=Moradi |first2=Amir Bahram |last3=Dovzhenko |first3=Oleksandr |last4=Touraev |first4=Alisher |last5=Palme |first5=Klaus |last6=Welsch |first6=Ralf |date=2022-01-13 |title=Molecular Control of Sporophyte-Gametophyte Ontogeny and Transition in Plants |journal=Frontiers in Plant Science |volume=12 |article-number=789789 |bibcode=2022FrPS...1289789P |doi=10.3389/fpls.2021.789789 |issn=1664-462X |pmc=8793881 |pmid=35095963 |doi-access=free}}</ref> The term "gymnosperm" is often used in [[paleobotany]] to refer to (the [[paraphyletic]] group of) all non-angiosperm seed plants. In that case, to specify the modern [[monophyletic]] group of gymnosperms, the term '''Acrogymnospermae''' is sometimes used.<ref>Coniferae, Gnetophyta. "1 Relationships of Angiosperms to Other Seed Plants."</ref>

The gymnosperms and [[angiosperms]] together constitute the [[spermatophytes]] or seed plants. The spermatophytes are subdivided into five [[Division (botany)|divisions]], the angiosperms and four divisions of gymnosperms: the [[Cycadophyta]], [[Ginkgophyta]], [[Gnetophyta]], and [[Pinophyta]] (also known as Coniferophyta). Newer classification place the gnetophytes among the conifers.<ref>{{Cite journal |last1=Yang |first1=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 |date=2022-07-01 |title=Recent advances on phylogenomics of gymnosperms and a new classification |journal=Plant Diversity |language=en |volume=44 |issue=4 |pages=340–350 |bibcode=2022PlDiv..44..340Y |doi=10.1016/j.pld.2022.05.003 |issn=2468-2659 |pmc=9363647 |pmid=35967253}}</ref> Numerous extinct seed plant groups are recognised including those considered [[Pteridospermatophyta|pteridosperms/seed ferns]], as well other groups like the Bennettitales.<ref name="Raven-2013">{{Cite book |last=Raven |first=P.H. |title=Biology of Plants |publisher=New York: W.H. Freeman and Co. |year=2013}}</ref>

By far the largest group of living gymnosperms are the conifers (pines, cypresses, and relatives), followed by cycads, gnetophytes (''[[Gnetum]]'', ''[[Ephedra (genus)|Ephedra]]'' and ''[[Welwitschia]]''), and ''[[Ginkgo biloba]]'' (a single living species). About 65% of gymnosperms are [[Dioecy|dioecious]],<ref>{{Cite journal |last1=Walas |first1=Łukasz |last2=Mandryk |first2=Wojciech |last3=Thomas |first3=Peter A. |last4=Tyrała-Wierucka |first4=Żanna |last5=Iszkuło |first5=Grzegorz |date=2018-09-01 |title=Sexual systems in gymnosperms: A review |url=https://www.sciencedirect.com/science/article/abs/pii/S1439179117304498 |journal=Basic and Applied Ecology |language=en |volume=31 |pages=1–9 |bibcode=2018BApEc..31....1W |doi=10.1016/j.baae.2018.05.009 |issn=1439-1791 |s2cid=90740232 |url-access=subscription}}</ref> but conifers are almost all [[Monoecy|monoecious]].<ref name="Walasetal">{{Cite journal |vauthors=Walas Ł, Mandryk W, Thomas PA, Tyrała-Wierucka Ż, Iszkuło G |date=2018 |title=Sexual systems in gymnosperms: A review |url=http://eprints.keele.ac.uk/4961/1/29052018_1-s2.0-S1439179117304498-main.pdf |journal=Basic and Applied Ecology |volume=31 |pages=1–9 |bibcode=2018BApEc..31....1W |doi=10.1016/j.baae.2018.05.009 |s2cid=90740232}}</ref> Some genera have [[ectomycorrhiza]] fungal associations with roots (''Pinus''),<ref>{{Cite journal |last1=Gehring |first1=Catherine A. |last2=Theimer |first2=Tad C. |last3=Whitham |first3=Thomas G. |last4=Keim |first4=Paul |date=1998 |title=Ectomycorrhizal Fungal Community Structure of Pinyon Pines Growing in Two Environmental Extremes |url=https://esajournals.onlinelibrary.wiley.com/doi/abs/10.1890/0012-9658%281998%29079%5B1562%3AEFCSOP%5D2.0.CO%3B2 |journal=Ecology |language=en |volume=79 |issue=5 |pages=1562–1572 |doi=10.1890/0012-9658(1998)079[1562:EFCSOP]2.0.CO;2 |issn=1939-9170 |url-access=subscription}}</ref> while in some others (''Cycas'') small specialised roots called coralloid roots are associated with nitrogen-fixing [[cyanobacteria]].<ref>{{Cite journal |last1=Chang |first1=Aimee Caye G. |last2=Chen |first2=Tao |last3=Li |first3=Nan |last4=Duan |first4=Jun |date=2019-08-14 |title=Perspectives on Endosymbiosis in Coralloid Roots: Association of Cycads and Cyanobacteria |journal=Frontiers in Microbiology |language=English |volume=10 |article-number=1888 |doi=10.3389/fmicb.2019.01888 |issn=1664-302X |pmc=6702271 |pmid=31474965 |doi-access=free}}</ref>

==Diversity and origin==

[[File:Encephalartos sclavoi reproductive cone.jpg|thumb|upright=0.5|''[[Encephalartos sclavoi]]'' cone, about 30 cm long]]

Over 1,000 living species of gymnosperm exist.<ref name="TPL">{{Cite web |title=Gymnosperms on The Plant List |url=http://www.theplantlist.org/browse/G/ |archive-url=https://web.archive.org/web/20130824024109/http://www.theplantlist.org/browse/G/ |archive-date=2013-08-24 |access-date=2013-07-24 |publisher=Theplantlist.org}}</ref> It was previously widely accepted that the gymnosperms originated in the [[Late Carboniferous]] period, replacing the [[Lycopodiophyta|lycopsid]] rainforests of the tropical region, but more recent phylogenetic evidence indicates that they diverged from the ancestors of [[Flowering plant|angiosperms]] during the [[Early Carboniferous]].<ref>{{Cite journal |last1=Li |first1=Hong-Tao |last2=Yi |first2=Ting-Shuang |last3=Gao |first3=Lian-Ming |last4=Ma |first4=Peng-Fei |last5=Zhang |first5=Ting |last6=Yang |first6=Jun-Bo |last7=Gitzendanner |first7=Matthew A. |last8=Fritsch |first8=Peter W. |last9=Cai |first9=Jie |last10=Luo |first10=Yang |last11=Wang |first11=Hong |date=May 2019 |title=Origin of angiosperms and the puzzle of the Jurassic gap |journal=Nature Plants |language=en |volume=5 |issue=5 |pages=461–470 |bibcode=2019NatPl...5..461L |doi=10.1038/s41477-019-0421-0 |pmid=31061536 |s2cid=146118264}}</ref><ref>{{Cite journal |last1=Morris |first1=Jennifer L. |last2=Puttick |first2=Mark N. |last3=Clark |first3=James W. |last4=Edwards |first4=Dianne |last5=Kenrick |first5=Paul |last6=Pressel |first6=Silvia |last7=Wellman |first7=Charles H. |last8=Yang |first8=Ziheng |last9=Schneider |first9=Harald |last10=Donoghue |first10=Philip C. J. |date=2018-03-06 |title=The timescale of early land plant evolution |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=115 |issue=10 |pages=E2274–E2283 |bibcode=2018PNAS..115E2274M |doi=10.1073/pnas.1719588115 |pmc=5877938 |pmid=29463716 |doi-access=free}}</ref> The radiation of gymnosperms during the late Carboniferous appears to have resulted from a whole [[Gene duplication|genome duplication]] event around {{ma|319}}.<ref name="Jiao2011">{{Cite journal |last1=Jiao |first1=Yuannian |last2=Wickett |first2=Norman J. |last3=Ayyampalayam |first3=Saravanaraj |last4=Chanderbali |first4=André S. |last5=Landherr |first5=Lena |last6=Ralph |first6=Paula E. |last7=Tomsho |first7=Lynn P. |last8=Hu |first8=Yi |last9=Liang |first9=Haiying |last10=Soltis |first10=Pamela S. |author-link10=Pamela S. Soltis |last11=Soltis |first11=Douglas E. |author-link11=Douglas E. Soltis |date=2011-04-10 |title=Ancestral polyploidy in seed plants and angiosperms |journal=Nature |volume=473 |issue=7345 |pages=97–100 |bibcode=2011Natur.473...97J |doi=10.1038/nature09916 |pmid=21478875 |s2cid=4313258}}</ref> Early characteristics of seed plants are evident in fossil [[progymnosperms]] of the late [[Devonian]] period around 383 million years ago. It has been suggested that during the mid-Mesozoic era, pollination of some extinct groups of gymnosperms was by extinct species of [[Mecoptera|scorpionflies]] that had specialized [[proboscis|proboscides]] for feeding on pollination drops. The scorpionflies likely engaged in pollination mutualisms with gymnosperms, long before the similar and independent coevolution of nectar-feeding insects on angiosperms.<ref>{{Cite journal |last1=Ollerton |first1=J. |last2=Coulthard |first2=E. |year=2009 |title=Evolution of Animal Pollination |journal=Science |volume=326 |issue=5954 |pages=808–809 |bibcode=2009Sci...326..808O |doi=10.1126/science.1181154 |pmid=19892970 |s2cid=856038}}</ref><ref name="Ren">{{Cite journal |last1=Ren |first1=D |last2=Labandeira |first2=CC |last3=Santiago-Blay |first3=JA |last4=Rasnitsyn |first4=A |last5=Shih |first5=CK |last6=Bashkuev |first6=A |last7=Logan |first7=MA |last8=Hotton |first8=CL |last9=Dilcher |first9=D. |display-authors=4 |year=2009 |title=A Probable Pollination Mode Before Angiosperms: Eurasian, Long-Proboscid Scorpionflies |journal=Science |volume=326 |issue=5954 |pages=840–847 |bibcode=2009Sci...326..840R |doi=10.1126/science.1178338 |pmc=2944650 |pmid=19892981}}</ref> Evidence has also been found that mid-Mesozoic gymnosperms were pollinated by [[Kalligrammatidae|Kalligrammatid lacewings]], a now-extinct family with members which (in an example of [[convergent evolution]]) resembled the modern butterflies that arose far later.<ref>{{Cite journal |last1=Labandeira |first1=Conrad C. |last2=Yang |first2=Qiang |last3=Santiago-Blay |first3=Jorge A. |last4=Hotton |first4=Carol L. |last5=Monteiro |first5=Antónia |last6=Wang |first6=Yong-Jie |last7=Goreva |first7=Yulia |last8=Shih |first8=ChungKun |last9=Siljeström |first9=Sandra |last10=Rose |first10=Tim R. |last11=Dilcher |first11=David L. |last12=Ren |first12=Dong |year=2016 |title=The evolutionary convergence of mid-Mesozoic lacewings and Cenozoic butterflies |journal=Proceedings of the Royal Society B: Biological Sciences |volume=283 |issue=1824 |article-number=20152893 |doi=10.1098/rspb.2015.2893 |pmc=4760178 |pmid=26842570}}</ref>
[[File:Zamia integrifolia02.jpg|thumb|''Zamia integrifolia,'' a cycad native to Florida]]

All gymnosperms are [[perennial]] [[woody plant]]s.<ref>{{Cite journal |last=Bond |first=W. J. |date=March 1989 |title=The tortoise and the hare: ecology of angiosperm dominance and gymnosperm persistence |journal=Biological Journal of the Linnean Society |volume=36 |issue=3 |pages=227–249 |doi=10.1111/j.1095-8312.1989.tb00492.x}}</ref>
Unlike in other extant gymnosperms the soft and highly [[Ground tissue#Parenchyma|parenchymatous]] wood in cycads is poorly lignified,<ref>{{Cite journal |last1=Fisher |first1=Jack B. |last2=Lindström |first2=Anders |last3=Marler |first3=Thomas E. |date=2009-06-01 |title=Tissue Responses and Solution Movement After Stem Wounding in Six Cycas Species |url=https://journals.ashs.org/hortsci/view/journals/hortsci/44/3/article-p848.xml |journal=HortScience |volume=44 |issue=3 |pages=848–851 |doi=10.21273/HORTSCI.44.3.848 |issn=0018-5345 |s2cid=83644706 |doi-access=free}}</ref> and their main structural support comes from an armor of sclerenchymatous leaf bases covering the stem,<ref>{{Cite book |last1=Bell |first1=Peter R. |url=https://books.google.com/books?id=HYkTvGq_RccC&q=armor%2520sclerenchymatous%2520leaf%2520bases&pg=PA247 |title=Green Plants: Their Origin and Diversity |last2=Bell |first2=Peter R. |last3=Hemsley |first3=Alan R. |date=2000 |publisher=Cambridge University Press |isbn=978-0-521-64673-4 |page=247}}</ref> with the exception of species with underground stems.<ref>{{Cite book |last1=Cleal |first1=Christopher J. |url=https://books.google.com/books?id=KDGdDwAAQBAJ&q=%2522exclusively%2520underground%2C%2520like%2520a%2520tuber%2520%28e.g.%2520some%2520Zamia%2520species%29%2522&pg=PA179 |title=Introduction to Plant Fossils |last2=Thomas |first2=Barry A. |date=2019 |publisher=Cambridge University Press |isbn=978-1-108-48344-5 |page=179}}</ref> There are no [[Herbaceous plant|herbaceous]] gymnosperms and compared to angiosperms they occupy fewer [[ecological niche]]s, but have evolved both parasites (''[[Parasitaxus]]''), [[epiphyte]]s (''[[Zamia pseudoparasitica]]'') and [[rheophyte]]s (''[[Retrophyllum minus]]'').<ref>{{cite journal |last=Mill |first=R. R. |date=2016-06-22 |title=A Monographic Revision of Retrophyllum (Podocarpaceae) |journal=Edinburgh Journal of Botany |volume=73 |issue=2 |pages=171–261 |doi=10.1017/S0960428616000081 |doi-access=free}}</ref>

[[Conifer]]s are by far the most abundant extant group of gymnosperms with six to eight families, with a total of 65–70 genera and 600–630 species (696 accepted names).<ref name="Catalogue">{{cite web |date=2006 |editor-last=A. Farjon |title=Conifer database |url=http://www.catalogueoflife.org/show_database_details.php?database_name=Conifer+Database |archive-url=https://web.archive.org/web/20090115190412/http://www.catalogueoflife.org/show_database_details.php?database_name=Conifer+Database |archive-date=January 15, 2009 |website=Catalogue of Life: 2008 Annual checklist}}</ref> Most conifers are [[evergreen]]s.<ref>Campbell, Reece, "Phylum Coniferophyta."Biology. 7th. 2005. Print. P.595</ref> The [[leaf|leaves]] of many conifers are long, thin and needle-like, while other species, including most [[Cupressaceae]] and some [[Podocarpaceae]], have flat, triangular scale-like leaves. ''[[Agathis]]'' in Araucariaceae and ''[[Nageia]]'' in Podocarpaceae have broad, flat strap-shaped leaves.{{cn|date=June 2024}}

[[Cycad]]s, small palm-like trees,<ref name="DK Pub.-2012" /> are the next most abundant group of gymnosperms, with two or three families, 11 genera, and approximately 338 species. A majority of cycads are native to tropical climates and are most abundantly found in regions near the equator. The other extant groups are the 95–100 species of [[Gnetophyta|''Gnetophytes'']] and one species of ''[[Ginkgo]]''. The ginkgo or maidenhair trees are tall and have bilobed leaves, while gnetophytes are a diverse groups of plants and shrubs including the horizontally growing [[welwitschia]]<ref name="Raven-2013" />

Gymnosperms are the most threatened of all plant groups.<ref>{{cite journal |last=Gilbert |first=Natasha |date=2010-09-28 |title=Threats to the world's plants assessed |url=https://www.nature.com/articles/news.2010.499 |journal=Nature |doi=10.1038/news.2010.499 |url-access=subscription}}</ref>

==Classification==

{{cladogram|title=Phylogeny of Gymnosperms<ref>{{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 |s2cid=52120430}}</ref><ref>{{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>{{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>{{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|style=font-size:90%;line-height:80%;width:400px
|1={{Clade
  |1={{Clade
    |label1='''[[Ginkgoidae]]'''
    |1={{Clade
      |label1='''[[Ginkgoales]]'''
      |1={{Clade
        |label1=[[Ginkgoaceae]]
        |1=''[[Ginkgo]]''
         }}
       }}
    |label2='''[[Cycadidae]]'''
    |2={{Clade
      |label1='''[[Cycadales]]'''
      |1={{Clade
        |label1=[[Cycadaceae]]
        |1=''[[Cycas]]''
        |label2=[[Zamiaceae]]
        |2={{Clade
          |label1=Diooideae
          |1=''[[Dioon]]''
          |label2=Zamioideae
          |2={{Clade
            |1={{Clade
              |1=''[[Bowenia]]''
              |2={{Clade
                |1=''[[Macrozamia]]''
                |2={{Clade
                  |1=''[[Encephalartos]]''
                  |2=''[[Lepidozamia]]''
                   }}
                 }}
               }}
            |2={{Clade
              |1={{Clade
                |1=''[[Ceratozamia]]''
                |2=''[[Stangeria]]''
                }}
              |2={{Clade
                |1=''[[Microcycas]]''
                |2=''[[Zamia]]''
                 }}
               }}
             }}
           }}
         }}
       }}
     }}
  |2={{Clade
  |label1x='''[[Pinidae]]'''
  |1={{Clade
    |1={{Clade
      |label1='''[[Gnetales]]'''
      |1={{Clade
        |1={{Clade
          |label1=[[Ephedraceae]]
          |1=''[[Ephedra (plant)|Ephedra]]''
           }}
        |2={{Clade
          |label1=[[Gnetaceae]]
          |1=''[[Gnetum]]''
          |label2=[[Welwitschiaceae]]
          |2=''[[Welwitschia]]''
           }}
         }}
      |label2='''[[Pinales]]'''
      |2={{Clade
        |label1=[[Pinaceae]]
        |1={{Clade
          |label1=Pinoideae
          |1={{Clade
            |1={{Clade
              |1=''[[Larix]]''
              |2=''[[Pseudotsuga]]''
               }}
            |2={{Clade
              |1=''[[Pinus]]''
              |2={{Clade
                |1=''[[Cathaya]]''
                |2=''[[Picea]]''
                 }}
               }}
             }}
          |label2=Abietoideae
          |2={{Clade
            |1=''[[Cedrus]]''
            |2={{Clade
              |1={{Clade
                |1=''[[Abies]]''
                |2=''[[Keteleeria]]''
                 }}
              |2={{Clade
                |1=''[[Pseudolarix]]''
                |2={{Clade
                  |1=''[[Nothotsuga]]''
                  |2=''[[Tsuga]]''
                   }}
                 }}
               }}
             }}
           }}
         }}
       }}
    |label2=
    |2={{Clade
      |label1='''[[Araucariales]]'''
      |1={{Clade
        |label1=[[Araucariaceae]]
        |1={{Clade
          |label1=Araucarieae
          |1=''[[Araucaria]]''
          |label2=Agathideae
          |2={{Clade
            |1=''[[Agathis]]''
            |2=''[[Wollemia]]''
             }}
           }}
        |label2=[[Podocarpaceae]]
        |2={{Clade
          |1={"Phyllocladoideae"}
          |2={"Podocarpoideae"}
           }}
         }}
      |label2='''[[Cupressales]]'''
      |2={{Clade
        |1={{Clade
          |label1=[[Sciadopityaceae]]
          |1=''[[Sciadopitys]]''
           }}
        |2={{Clade
          |label1=[[Taxaceae]]
          |1={{Clade
            |label1=Cephalotaxeae
            |1=''[[Cephalotaxus]]''
            |label2=Taxoideae
            |2={{Clade
              |1={{Clade
                |1=''[[Amentotaxus]]''
                |2=''[[Torreya]]''
                 }}
              |2={{Clade
                |1=''[[Austrotaxus]]''
                |2={{Clade
                  |1=''[[Pseudotaxus]]''
                  |2=''[[Taxus]]''
                   }}
                 }}
               }}
             }}
          |label2=[[Cupressaceae]]
          |2={{Clade
            |1={{Clade
              |label1=Cunninghamioideae
              |1=''[[Cunninghamia]]''
               }}
            |2={{Clade
              |1={{Clade
                |label1=Taiwanioideae
                |1=''[[Taiwania]]''
                 }}
              |2={{Clade
                |1={{Clade
                  |label1=Athrotaxidoideae
                  |1=''[[Athrotaxis]]''
                   }}
                |2={{Clade
                  |1={{Clade
                    |label1=Sequoioideae
                    |1={{Clade
                      |1=''[[Metasequoia]]''
                      |2={{Clade
                        |1=''[[Sequoiadendron]]''
                        |2=''[[Sequoia (genus)|Sequoia]]''
                         }}
                       }}
                     }}
                  |2={{Clade
                    |1={{Clade
                      |label1=Taxodioideae
                      |1={{Clade
                        |1=''[[Cryptomeria]]''
                        |2={{Clade
                          |1=''[[Glyptostrobus]]''
                          |2=''[[Taxodium]]''
                           }}
                         }}
                       }}
                    |2={{Clade
                      |1={"Actinostroboideae"}
                      |2={"Cupressoideae"}
                       }}
                     }}
                   }}
                 }}
               }}
             }}
           }}
         }}
       }}
     }}
   }}
   }}
|targetA={"Phyllocladoideae"}
|subcladeA={{Clade
|label1 =Phyllocladoideae
|1={{Clade hidden |id=1 |mode=left
  |1=''[[Halocarpus]]''
  |2={{Clade
    |1={{Clade
      |1=''[[Pectinopitys]]''
      |2={{Clade
        |1=''[[Prumnopitys]]''
        |2=''[[Sundacarpus]]''
         }}
       }}
    |2={{Clade
      |1={{Clade
        |1=''[[Lepidothamnus]]''
        |2=''[[Phyllocladus]]''
         }}
      |2={{Clade
        |1=''[[Parasitaxus]]''
        |2={{Clade
          |1=''[[Lagarostrobos]]''
          |2=''[[Manoao]]''
           }}
         }}

       }}
     }}
   }}
           }}
|targetB={"Podocarpoideae"}
|subcladeB={{Clade
|label1 =Podocarpoideae
|1={{Clade hidden |id=2 |mode=left
  |1=''[[Saxegothaea]]''
  |2={{Clade
    |1=''[[Microcachrys]]''
    |2={{Clade
      |1=''[[Pherosphaera]]''
      |2={{Clade
        |1=''[[Acmopyle]]''
        |2={{Clade
          |1={{Clade
            |1=''[[Dacrycarpus]]''
            |2={{Clade
              |1=''[[Dacrydium]]''
              |2=''[[Falcatifolium]]''
               }}
             }}
          |2={{Clade
            |1=''[[Podocarpus]]''
            |2={{Clade
              |1=''[[Retrophyllum]]''
              |2={{Clade
                |1=''[[Afrocarpus]]''
                |2=''[[Nageia]]''
                 }}
               }}
             }}
           }}
         }}
       }}
     }}
   }}
           }}
|targetC={"Actinostroboideae"}
|subcladeC={{Clade
|label1 =Actinostroboideae
|1={{Clade hidden |id=3 |mode=left
  |1=''[[Papuacedrus]]''
  |2={{Clade
    |1=''[[Austrocedrus]]''
    |2={{Clade
      |1={{Clade
        |1=''[[Libocedrus]]''
        |2=''[[Pilgerodendron]]''
         }}
      |2={{Clade
        |1={{Clade
          |1=''[[Callitris]]''
          |2=''[[Neocallitropsis]]''
           }}
        |2={{Clade
          |1=''[[Widdringtonia]]''
          |2={{Clade
            |1=''[[Diselma]]''
            |2=''[[Fitzroya]]''
             }}
           }}
         }}
       }}
     }}
   }}
           }}
|targetD={"Cupressoideae"}
|subcladeD={{Clade
|label1 =Cupressoideae
|1={{Clade hidden |id=4 |mode=left
  |1={{Clade
    |1=''[[Thuja]]''
    |2=''[[Thujopsis]]''
     }}
  |2={{Clade
    |1=''[[Chamaecyparis]]''
    |2={{Clade
      |1={{Clade
        |1=''[[Calocedrus]]''
        |2={{Clade
          |1=''[[Tetraclinis]]''
          |2={{Clade
            |1=''[[Microbiota decussata|Microbiota]]''
            |2=''[[Platycladus]]''
             }}
           }}
         }}
      |2={{Clade
        |1=''[[Juniperus]]''
        |2={{Clade
          |1=''[[Cupressus]]''
          |2={{Clade
            |1=''[[Hesperocyparis]]''
            |2=''[[Xanthocyparis]]''
             }}
           }}
         }}
       }}
     }}
   }}
 }}
}}
}}

{{further|Spermatophyte}}
A formal classification of the living gymnosperms is the "Acrogymnospermae", which form a [[monophyletic group]] within the [[spermatophyte]]s.{{sfn|Cantino|2007}}<ref name="Christenhusz-2011">{{Cite journal |last1=Christenhusz |first1=M.J.M. |last2=Reveal |first2=J.L. |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 |url=http://www.mapress.com/phytotaxa/content/2011/f/pt00019p070.pdf |journal=Phytotaxa |volume=19 |issue=1 |pages=55–70 |bibcode=2011Phytx..19...55C |doi=10.11646/phytotaxa.19.1.3 |s2cid=86797396}}</ref> The wider "Gymnospermae" group includes extinct gymnosperms and is thought to be [[Paraphyly|paraphyletic]]. The fossil record of gymnosperms includes many distinctive [[taxa]] that do not belong to the four modern groups, including seed-bearing trees that have a somewhat [[fern]]-like vegetative morphology (the so-called "seed ferns" or [[pteridosperm]]s).<ref>{{Cite journal |last1=Hilton |first1=Jason |last2=Bateman |first2=Richard M. |date=January 2006 |title=Pteridosperms are the backbone of seed-plant phylogeny 1 |journal=The Journal of the Torrey Botanical Society |language=en |volume=133 |issue=1 |pages=119–168 |doi=10.3159/1095-5674(2006)133[119:PATBOS]2.0.CO;2 |s2cid=86395036}}</ref> When fossil gymnosperms such as these and the [[Bennettitales]], [[glossopterid]]s, and ''[[Caytonia]]'' are considered, it is clear that angiosperms are nested within a larger gymnospermae clade, although which group of gymnosperms is their closest relative remains unclear.

The extant gymnosperms include 12 main families and 83 genera which contain more than 1000 known species.<ref name="TPL" /><ref name="Christenhusz-2011" /><ref name="Christenhusz-Byng2016">{{Cite journal |last1=Christenhusz, M. J. M. |last2=Byng, J. W. |year=2016 |title=The number of known plants species in the world and its annual increase |url=http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 |journal=Phytotaxa |volume=261 |issue=3 |pages=201–217 |bibcode=2016Phytx.261..201C |doi=10.11646/phytotaxa.261.3.1 |doi-access=free}}</ref>

'''Subclass [[Cycadidae]]'''
* Order '''[[Cycadales]]'''
** Family '''[[Cycadaceae]]''': ''[[Cycas]]''
** Family '''[[Zamiaceae]]''': ''[[Dioon]]'', ''[[Bowenia]]'', ''[[Macrozamia]]'', ''[[Lepidozamia]]'', ''[[Encephalartos]]'', ''[[Stangeria]]'', ''[[Ceratozamia]]'', ''[[Microcycas]]'', ''[[Zamia]]''

'''Subclass [[Ginkgoidae]]'''
* Order '''[[Ginkgoales]]'''
** Family '''[[Ginkgoaceae]]''': ''[[Ginkgo]]''

'''Subclass [[Gnetidae]]'''
* Order '''[[Welwitschiales]]'''
** Family '''[[Welwitschiaceae]]''': ''[[Welwitschia]]''
* Order '''[[Gnetales]]'''
** Family '''[[Gnetaceae]]''': ''[[Gnetum]]''
* Order '''[[Ephedrales]]'''
** Family '''[[Ephedraceae]]''': ''[[Ephedra (plant)|Ephedra]]''

'''Subclass [[Pinidae]]'''
* Order '''[[Pinales]]'''
** Family '''[[Pinaceae]]''': ''[[Cedrus]]'', ''[[Pinus]]'', ''[[Cathaya]]'', ''[[Picea]]'', ''[[Pseudotsuga]]'', ''[[Larix]]'', ''[[Pseudolarix]]'', ''[[Tsuga]]'', ''[[Nothotsuga]]'', ''[[Keteleeria]]'', ''[[Abies]]''
* Order '''[[Araucariales]]'''
** Family '''[[Araucariaceae]]''': ''[[Araucaria]]'', ''[[Wollemia]]'', ''[[Agathis]]''
** Family '''[[Podocarpaceae]]''': ''[[Phyllocladus]]'', ''[[Lepidothamnus]]'', ''[[Prumnopitys]]'', ''[[Sundacarpus]]'', ''[[Halocarpus]]'', ''[[Parasitaxus]]'', ''[[Lagarostrobos]]'', ''[[Manoao]]'', ''[[Saxegothaea]]'', ''[[Microcachrys]]'', ''[[Pherosphaera]]'', ''[[Acmopyle]]'', ''[[Dacrycarpus]]'', ''[[Dacrydium]]'', ''[[Falcatifolium]]'', ''[[Retrophyllum]]'', ''[[Nageia]]'', ''[[Afrocarpus]]'', ''[[Podocarpus]]''
* Order '''[[Cupressales]]'''
** Family '''[[Sciadopityaceae]]''': ''[[Sciadopitys]]''
** Family '''[[Cupressaceae]]''': ''[[Cunninghamia]]'', ''[[Taiwania]]'', ''[[Athrotaxis]]'', ''[[Metasequoia]]'', ''[[Sequoioideae|Sequoia]]'', ''[[Sequoiadendron]]'', ''[[Cryptomeria]]'', ''[[Glyptostrobus]]'', ''[[Taxodium]]'', ''[[Papuacedrus]]'', ''[[Austrocedrus]]'', ''[[Libocedrus]]'', ''[[Pilgerodendron]]'', ''[[Widdringtonia]]'', ''[[Diselma]]'', ''[[Fitzroya]]'', ''[[Callitris]]'', ''[[Actinostrobus]]'', ''[[Neocallitropsis]]'', ''[[Thujopsis]]'', ''[[Thuja]]'', ''[[Fokienia]]'', ''[[Chamaecyparis]]'', ''[[Cupressus]]'', ''[[Juniperus]]'', ''[[Calocedrus]]'', ''[[Tetraclinis]]'', ''[[Platycladus]]'', ''[[Microbiota (plant)|Microbiota]]''
** Family '''[[Taxaceae]]''': ''[[Austrotaxus]]'', ''[[Pseudotaxus]]'', ''[[Taxus]]'', ''[[Cephalotaxus]]'', ''[[Amentotaxus]]'', ''[[Torreya]]''

=== Extinct groupings ===
* Order [[Cordaitales]]
* Order [[Calamopityales]]
* Order [[Callistophytales]]
* Order [[Caytoniales]]
* Order [[Gigantopteridales]]
* Order [[Glossopteridales]]
* Order [[Lyginopteridales]]
* Order [[Medullosales]]
* Order [[Peltaspermales]]
* Order [[Corystospermales]] (also known as Umkomasiales)
* Order [[Czekanowskiales]]
* Order [[Bennettitales]] (cycadeoids)
* Order [[Erdtmanithecales]]
* Order [[Pentoxylales]]
* Order [[Petriellales]]

==Life cycle==
[[File:Gymnosperm life cycle (en).png|thumb|Example of gymnosperm lifecycle]]
Gymnosperms, like all [[vascular plants]], have a sporophyte-dominant life cycle, which means they spend most of their life cycle with diploid cells, while the [[gametophyte]] (gamete-bearing phase) is relatively short-lived. Like all [[seed plants]], they are [[heterospory|heterosporous]], having two spore types, [[microspore]]s (male) produced in [[microsporangium]] and [[megaspore]]s (female) produced in [[Sporangium|megasporangium]] that are typically present in pollen cones or ovulate cones respectively. The microsporangium is carried by [[Sporophyll|microsporophyll]] (modified leaf) and seeds are carried by ovuliferous scales in the male and female cones respectively.<ref name="DK Pub.-2012" /><ref name="Concepts of Biology">{{Cite book |last1=Samantha |first1=Fowler |url=https://openstax.org/books/concepts-biology/pages/14-3-seed-plants-gymnosperms |title=Concepts of Biology |last2=Rebecca |first2=Roush |last3=James |first3=Wise |date=2013 |publisher=OpenStax |location=Houston, Texas |chapter=14.3 Seed Plants: Gymnosperms |access-date=31 March 2023}}</ref> The exception is the females in the cycad genus ''[[Cycas]]'', which form a loose structure called megasporophylls instead of cones.<ref>{{Cite journal |last1=Liu |first1=Yang |last2=Wang |first2=Sibo |last3=Li |first3=Linzhou |last4=Yang |first4=Ting |last5=Dong |first5=Shanshan |last6=Wei |first6=Tong |last7=Wu |first7=Shengdan |last8=Liu |first8=Yongbo |last9=Gong |first9=Yiqing |last10=Feng |first10=Xiuyan |last11=Ma |first11=Jianchao |last12=Chang |first12=Guanxiao |last13=Huang |first13=Jinling |last14=Yang |first14=Yong |last15=Wang |first15=Hongli |date=April 2022 |title=The Cycas genome and the early evolution of seed plants |journal=Nature Plants |language=en |volume=8 |issue=4 |pages=389–401 |bibcode=2022NatPl...8..389L |doi=10.1038/s41477-022-01129-7 |issn=2055-0278 |pmc=9023351 |pmid=35437001}}</ref> As with all heterosporous plants, the gametophytes develop within the spore wall. Pollen grains (microgametophytes) mature from microspores, and ultimately produce sperm cells.<ref name="Concepts of Biology" /> Megagametophytes develop from megaspores and are retained within the ovule. Gymnosperms produce multiple [[Archegonium|archegonia]], which produce the female gametes.{{cn|date=June 2024}}

During pollination, pollen grains are physically transferred between plants from the pollen cone to the ovule. Pollen is usually moved by wind or insects. Whole grains enter each ovule through a microscopic gap in the ovule coat ([[integument]]) called the micropyle. The pollen grains mature further inside the ovule and produce sperm cells. Two main modes of fertilization are found in gymnosperms. Cycads and ''[[Ginkgo]]'' have [[flagellum|flagellated]] motile sperm<ref name="AJB">{{Cite journal |last1=Southworth |first1=Darlene |last2=Cresti |first2=Mauro |date=September 1997 |title=Comparison of flagellated and nonflagellated sperm in plants |journal=American Journal of Botany |volume=84 |issue=9 |pages=1301–1311 |doi=10.2307/2446056 |jstor=2446056 |pmid=21708687 |doi-access=free}}</ref> that swim directly to the egg inside the ovule, whereas conifers and [[gnetophyte]]s have sperm with no flagella that are moved along a [[pollen tube]] to the egg. After [[Fertilisation|syngamy]] (joining of the sperm and egg cell), the zygote develops into an embryo (young sporophyte). More than one embryo is usually initiated in each gymnosperm seed. The mature seed comprises the embryo and the remains of the female [[Gametophyte#Seed plants|gametophyte]], which serves as a food supply, and the [[Seed-coat|seed coat]].<ref name="Walters1996">{{Cite book |last=Walters |first=Dirk R Walters Bonnie By |url=https://archive.org/details/vascularplanttax00walt |title=Vascular plant taxonomy |publisher=Kendall/Hunt Pub. Co. |year=1996 |isbn=978-0-7872-2108-9 |location=Dubuque, Iowa |page=[https://archive.org/details/vascularplanttax00walt/page/124 124] |quote=Gymnosperm seeds. |url-access=registration}}</ref>

Gymnosperms ordinarily reproduce by [[sexual reproduction]], and only rarely express parthenogenesis.<ref name="Horandl2024">Hörandl E. Apomixis and the paradox of sex in plants. Ann Bot. 2024 Mar 18:mcae044. doi: 10.1093/aob/mcae044. Epub ahead of print. PMID 38497809</ref> Sexual reproduction in gymnosperms appears to be required for maintaining long-term [[genome|genomic]] integrity.<ref name = Horandl2024/> [[Meiosis]] in sexual land plants provides a direct mechanism for [[DNA repair|repairing DNA]] in reproductive tissues.<ref name = Horandl2024/> The likely primary benefit of cross-pollination in gymnosperms, as in other eukaryotes, is that it allows the avoidance of inbreeding depression caused by the presence of recessive deleterious mutations.<ref>Charlesworth D, Willis JH. The genetics of inbreeding depression. Nat Rev Genet. 2009 Nov;10(11):783-96. doi: 10.1038/nrg2664. PMID 19834483</ref>

==Genetics==
The first published sequenced genome for any gymnosperm was the genome of ''[[Picea abies]]'' in 2013.<ref>{{Cite journal |last1=Nystedt |first1=B |last2=Street |first2=NR |last3=Wetterbom |first3=A |display-authors=etal |date=May 2013 |title=The Norway spruce genome sequence and conifer genome evolution |journal=Nature |volume=497 |issue=7451 |pages=579–584 |bibcode=2013Natur.497..579N |doi=10.1038/nature12211 |hdl=1854/LU-4110028 |pmid=23698360 |doi-access=free |hdl-access=free}}</ref>

==Uses==
Gymnosperms have major economic uses. Some, such as pine, fir, spruce, and cedar, are used for [[lumber]], paper production, and resin. Some other common uses for gymnosperms are [[soap]], [[varnish]], [[nail polish]], food, gum, and [[perfume]]s.<ref>{{Cite book |last1=Biswas |first1=C. |url=https://link.springer.com/content/pdf/10.1007%2F978-3-662-13164-0.pdf |title=The Gymnosperms |last2=Johri |first2=B.M. |publisher=Springer, Berlin, Heidelberg |year=1997 |isbn=978-3-662-13166-4 |pages=440–456 |chapter=Economic Importance |doi=10.1007/978-3-662-13164-0_23}}</ref>

== References ==
{{Reflist|30em|refs=

<ref name="Phylogenomics of gymnosperms">{{Cite journal |vauthors=Yang Y, Ferguson DK, Liu B, Mao KS, Gao LM, Zhang SZ, Wan T, Rushforth K, Zhang ZX |date=2020 |title=Recent advances on phylogenomics of gymnosperms and a new classification |journal=Plant Diversity |volume=44 |issue=4 |pages=340–350 |bibcode=2022PlDiv..44..340Y |doi=10.1016/j.pld.2022.05.003 |issn=2468-2659 |pmc=9363647 |pmid=35967253 |s2cid=249117306}}</ref>

}}

== General bibliography ==
{{refbegin}}
* {{Cite journal |last1=Cantino |first1=Philip D. |last2=Doyle |first2=James A. |last3=Graham |first3=Sean W. |last4=Judd |first4=Walter S. |last5=Olmstead |first5=Richard G. |last6=Soltis |first6=Douglas E. |last7=Soltis |first7=Pamela S. |last8=Donoghue |first8=Michael J. |date=August 2007 |title=Towards a phylogenetic nomenclature of Tracheophyta |journal=[[Taxon (journal)|Taxon]] |volume=56 |issue=3 |pages=822–846 |bibcode=2007Taxon..56..822C |doi=10.2307/25065864 |jstor=25065864 |ref={{harvid|Cantino|2007}}}}
{{refend}}

==External links==
{{Commonscat|Gymnosperms}}
{{wiktionary|gymnosperm}}
* [http://www.conifers.org/ Gymnosperm Database]
* [https://web.archive.org/web/20080409044237/http://www.huh.harvard.edu/research/mathews-lab/atolHtmlSite/ Gymnosperms on the Tree of Life]
* {{Cite EB1911|wstitle=Gymnosperms|author=Albert Seward|author-link=Albert Seward|short=x}}
* [[Christopher Nigel Page]] (2023).The Evolution of Arborescent Gymnosperms. [[Cambridge University Press]]. {{ISBN|978-1-009-26309-2}}. 2 volumes (volume one on the conifers of the Northern hemisphere and volume 2 on the conifers of the southern hemisphere).
{{Clear}}
{{Botany}}
{{Plant classification}}
{{Acrogymnospermae classification}}
{{Taxonbar|from=Q133712}}
{{Authority control}}
 [[Category:Gymnosperms|01]]
[[Category:Extant Late Devonian first appearances]]