View source for Biology:Gymnosperm

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

The '''gymnosperms''' ({{IPAc-en|'|dZ|I|m|n|ə|s|p|ɜːr|m|z|,_|-|n|oʊ|-}} {{respell|JIM|nə|spurmz|,_|-noh-}} {{audio|en-us-gymnosperm.ogg|pronunciation}} [[Social:Greek language|lit]]. revealed seeds) are a group of [[Biology:Spermatophyte|seed-producing plants]] that includes [[Biology:Pinophyta|conifers]], [[Biology:Cycad|cycad]]s, ''[[Biology:Ginkgo|Ginkgo]]'', and [[Biology:Gnetophyta|gnetophyte]]s, forming the clade '''Gymnospermae'''. The term ''gymnosperm'' comes from the composite word in {{lang-el|γυμνόσπερμος}} ({{lang-el|γυμνός|translit=gymnos|lit=naked|label=none}} and {{lang-el|σπέρμα|translit=sperma|lit=seed|label=none}}), literally meaning 'naked seeds'. The name is based on the unenclosed condition of their seeds (called [[Biology:Ovule|ovule]]s in their unfertilized state). The non-encased condition of their seeds contrasts with the seeds and ovules of flowering plants ([[Biology:Angiosperms|angiosperms]]), which are enclosed within an [[Biology:Ovary (botany)|ovary]]. Gymnosperm seeds develop either on the surface of scales or [[Biology:Leaf|leaves]], which are often modified to form [[Conifer cone|cones]], or on their own as in [[Biology:Taxus|yew]], ''[[Biology:Torreya|Torreya]]'', ''[[Biology:Ginkgo|Ginkgo]]''.<ref name="TPL">{{cite web|url=http://www.theplantlist.org/browse/G/|title=Gymnosperms on The Plant List|publisher=Theplantlist.org|access-date=2013-07-24|archive-date=2013-08-24|archive-url=https://web.archive.org/web/20130824024109/http://www.theplantlist.org/browse/G/|url-status=dead}}</ref> Gymnosperm lifecycles involve [[Biology:Alternation of generations|alternation of generations]]. They have a dominant diploid [[Biology:Sporophyte|sporophyte]] phase and a reduced [[Biology:Haploid|haploid]] [[Biology:Gametophyte|gametophyte]] phase which is dependent on the sporophytic phase. The term "gymnosperm" is often used in [[Biology:Paleobotany|paleobotany]] to refer to (the paraphyletic group of) all non-angiosperm seed plants. In that case, to specify the modern [[Biology:Monophyletic|monophyletic]] group of gymnosperms, the term '''Acrogymnospermae''' is sometimes used.

The gymnosperms and [[Biology:Angiosperms|angiosperms]] together comprise the [[Biology:Spermatophyte|spermatophyte]]s or seed plants. The spermatophytes are subdivided into five divisions, the angiosperms and four divisions of gymnosperms: the [[Biology:Cycadophyta|Cycadophyta]], [[Biology:Ginkgophyta|Ginkgophyta]], [[Biology:Gnetophyta|Gnetophyta]], and [[Biology:Pinophyta|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 |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 [[Biology:Pteridospermatophyta|pteridosperms/seed ferns]], as well other groups like the [[Biology:Bennettitales|Bennettitales]].<ref name="Raven-2013">{{Cite book|title=Biology of Plants|last=Raven|first=P.H.|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 (''[[Biology:Gnetum|Gnetum]]'', ''Ephedra'' and ''[[Biology:Welwitschia|Welwitschia]]''), and ''[[Biology:Ginkgo biloba|Ginkgo biloba]]'' (a single living species). About 65% of gymnosperms are [[Biology:Dioecy|dioecious]],<ref>{{Cite journal|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|doi=10.1016/j.baae.2018.05.009|issn=1439-1791|last1=Walas|first1=Łukasz|last2=Mandryk|first2=Wojciech|last3=Thomas|first3=Peter A.|last4=Tyrała-Wierucka|first4=Żanna|last5=Iszkuło|first5=Grzegorz|s2cid=90740232}}</ref> but conifers are almost all [[Biology: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|journal=Basic and Applied Ecology|volume=31|pages=1–9|doi=10.1016/j.baae.2018.05.009 |s2cid=90740232|url=http://eprints.keele.ac.uk/4961/1/29052018_1-s2.0-S1439179117304498-main.pdf}}</ref>

Some genera have [[Biology:Mycorrhiza|mycorrhiza]], fungal associations with roots (''Pinus''), while in some others (''Cycas'') small specialised roots called coralloid roots are associated with nitrogen-fixing [[Biology:Cyanobacteria|cyanobacteria]].

==Diversity and origin==
[[File:Encephalartos sclavoi reproductive cone.jpg|thumb|upright|''[[Biology:Encephalartos sclavoi|Encephalartos sclavoi]]'' cone, about 30 cm long]]
Over 1,000 living species of gymnosperm exist.<ref name="TPL" /> It was previously widely accepted that the gymnosperms originated in the Late Carboniferous period, replacing the [[Biology:Lycopodiophyta|lycopsid]] rainforests of the tropical region, but more recent phylogenetic evidence indicates that they diverged from the ancestors of [[Biology: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 |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 |doi=10.1073/pnas.1719588115 |pmc=5877938 |pmid=29463716|bibcode=2018PNAS..115E2274M |doi-access=free }}</ref> The radiation of gymnosperms during the late Carboniferous appears to have resulted from a whole [[Biology: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. |last11=Soltis |first11=Douglas E. |date=2011-04-10 |title=Ancestral polyploidy in seed plants and angiosperms |journal=Nature |volume=473 |issue=7345 |pages=97–100 |doi=10.1038/nature09916 |pmid=21478875 |bibcode=2011Natur.473...97J |s2cid=4313258 |author-link11=Douglas E. Soltis}}</ref> Early characteristics of seed plants are evident in fossil progymnosperms of the late [[Earth:Devonian|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 [[Biology:Mecoptera|scorpionflies]] that had specialized [[Biology:Proboscis|proboscis]] 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 | doi = 10.1126/science.1181154 | pmid = 19892970 | bibcode = 2009Sci...326..808O | 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 | doi = 10.1126/science.1178338 | pmid = 19892981 | pmc = 2944650 | bibcode = 2009Sci...326..840R }}</ref> Evidence has also been found that mid-Mesozoic gymnosperms were pollinated by [[Biology:Kalligrammatidae|Kalligrammatid lacewings]], a now-extinct family with members which (in an example of [[Biology:Convergent evolution|convergent evolution]]) resembled the modern butterflies that arose far later.<ref>{{cite journal | title=The evolutionary convergence of mid-Mesozoic lacewings and Cenozoic butterflies | journal = Proceedings of the Royal Society B: Biological Sciences |  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 | volume = 283 | issue = 1824| doi=10.1098/rspb.2015.2893 | pages=20152893 | pmid=26842570 | pmc=4760178| year = 2016 }}</ref>
[[File:Zamia integrifolia02.jpg|thumb|''Zamia integrifolia,'' a cycad native to Florida]]

All gymnosperms are [[Biology:Perennial|perennial]] [[Biology:Woody plant|woody plant]]s,<ref>{{Cite journal
| title= The tortoise and the hare: ecology of angiosperm dominance and gymnosperm persistence
| first= W. J.| last= Bond 
| journal=Biological Journal of the Linnean Society|volume= 36|issue= 3|date= March 1989 |pages =227–249 |doi=10.1111/j.1095-8312.1989.tb00492.x
}}</ref>  
Unlike in other extant gymnosperms the soft and highly [[Biology: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|language=en-US|volume=44|issue=3|pages=848–851|doi=10.21273/HORTSCI.44.3.848|s2cid=83644706 |issn=0018-5345|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 [[Biology:Herbaceous plant|herbaceous]] gymnosperms and compared to angiosperms they occupy fewer [[Biology:Ecological niche|ecological niche]]s, but have evolved both parasites (''[[Biology:Parasitaxus|Parasitaxus]]''), [[Earth:Epiphyte|epiphyte]]s (''[[Biology:Zamia pseudoparasitica|Zamia pseudoparasitica]]'') and [[Biology:Rheophyte|rheophyte]]s (''[[Biology:Retrophyllum minus|Retrophyllum minus]]'').<ref>{{Cite journal|last=Mill|first=R. R.|title=A Monographic Revision of Retrophyllum (Podocarpaceae) |date=2016-06-22|url=https://www.cambridge.org/core/journals/edinburgh-journal-of-botany/article/monographic-revision-of-retrophyllum-podocarpaceae/029F1D48E9AFB1134D527CFF639B7F2E|journal=Edinburgh Journal of Botany|language=en|volume=73|issue=2|pages=171–261|doi=10.1017/S0960428616000081|issn=1474-0036|doi-access=free}}</ref>

[[Biology:Conifer|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|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|url-status=dead|work=Catalogue of Life: 2008 Annual checklist |title= Conifer database|editor= A. Farjon | date=2006|archive-date=January 15, 2009}}</ref> Most conifers are [[Biology:Evergreen|evergreens]].<ref>Campbell, Reece, "Phylum Coniferophyta."Biology. 7th. 2005. Print. P.595</ref> The [[Biology:Leaf|leaves]] of many conifers are long, thin and needle-like, while other species, including most [[Biology:Cupressaceae|Cupressaceae]] and some [[Biology:Podocarpaceae|Podocarpaceae]], have flat, triangular scale-like leaves. ''[[Biology:Agathis|Agathis]]'' in Araucariaceae and ''[[Biology:Nageia|Nageia]]'' in Podocarpaceae have broad, flat strap-shaped leaves.

[[Biology:Cycad|Cycad]]s 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 [[Biology:Gnetum|Gnetale]]s and one species of ''[[Biology:Ginkgo|Ginkgo]]''.<ref name="Raven-2013" />

Today 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 |language=en |doi=10.1038/news.2010.499 |issn=1476-4687}}</ref>

==Classification==

{{cladogram|title=Phylogeny of Acrogymnospermae<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 |url=https://doi.org/10.1002/ajb2.1143 |volume=105 |issue=9 |pages=1531–1544 | doi=10.1002/ajb2.1143 |pmid= 30157290|pmc= |bibcode= |s2cid=52120430 |doi-access=}}</ref><ref>{{cite journal |last1=Leslie |first1=Andrew B. |display-authors=et al. |year=2018 |title=ajb21143-sup-0004-AppendixS4 |journal=American Journal of Botany |volume=105 |issue=9 |pages=1531–1544 |doi=10.1002/ajb2.1143 |pmid=30157290 |s2cid=52120430 |url=https://bsapubs.onlinelibrary.wiley.com/action/downloadSupplement?doi=10.1002%2Fajb2.1143&file=ajb21143-sup-0004-AppendixS4.pdf |doi-access=}}</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 |journal=Nature Plants |url=https://www.nature.com/articles/s41477-021-00964-4 |volume=7 |issue= 8|pages=1015–1025 |doi=10.1038/s41477-021-00964-4|biorxiv=10.1101/2021.03.13.435279 |pmid= 34282286|pmc= |bibcode= |s2cid=232282918 |doi-access=}}</ref><ref>{{cite report |last1=Stull |first1=Gregory W. |display-authors=et al. |year=2021 |title=main.dated.supermatrix.tree.T9.tre |publisher=Figshare |doi=10.6084/m9.figshare.14547354.v1 |url=https://figshare.com/articles/dataset/Gene_duplications_and_genomic_conflict_underlie_major_pulses_of_phenotypic_evolution_in_gymnosperms/14547354 |doi-access=}}</ref>|
{{clade|style=font-size:90%;line-height:80%;width:400px
|1={{Clade
  |1={{Clade
    |label1='''[[Biology:Ginkgoidae|Ginkgoidae]]'''
    |1={{Clade
      |label1='''[[Biology:Ginkgoales|Ginkgoales]]'''
      |1={{Clade
        |label1=[[Biology:Ginkgoaceae|Ginkgoaceae]]
        |1=''[[Biology:Ginkgo|Ginkgo]]''
         }}
       }}
    |label2='''[[Biology:Cycadidae|Cycadidae]]'''
    |2={{Clade
      |label1='''[[Biology:Cycadales|Cycadales]]'''
      |1={{Clade
        |label1=[[Biology:Cycadaceae|Cycadaceae]]
        |1=''[[Biology:Cycas|Cycas]]''
        |label2=[[Biology:Zamiaceae|Zamiaceae]]
        |2={{Clade
          |label1=Diooideae
          |1=''[[Biology:Dioon|Dioon]]''
          |label2=Zamioideae
          |2={{Clade
            |1={{Clade
              |1=''[[Biology:Bowenia|Bowenia]]''
              |2={{Clade
                |1=''[[Biology:Macrozamia|Macrozamia]]''
                |2={{Clade
                  |1=''[[Biology:Lepidozamia|Lepidozamia]]''
                  |2=''[[Biology:Encephalartos|Encephalartos]]''
                   }}
                 }}
               }}
            |2={{Clade
              |1={{Clade
                |1=''[[Biology:Stangeria|Stangeria]]''
                |2=''[[Biology:Ceratozamia|Ceratozamia]]''
                }}
              |2={{Clade
                |1=''[[Biology:Microcycas|Microcycas]]''
                |2=''[[Biology:Zamia|Zamia]]''
                 }}
               }}
             }}
           }}
         }}
       }}
     }}
  |2={{Clade
  |label1='''[[Biology:Pinidae|Pinidae]]'''
  |1={{Clade
    |1={{Clade
      |label1='''[[Biology:Gnetales|Gnetales]]'''
      |1={{Clade
        |1={{Clade
          |label1=[[Biology:Welwitschiaceae|Welwitschiaceae]]
          |1=''[[Biology:Welwitschia|Welwitschia]]''
          |label2=[[Biology:Gnetaceae|Gnetaceae]]
          |2=''[[Biology:Gnetum|Gnetum]]''
           }}
        |2={{Clade
          |label1=[[Biology:Ephedraceae|Ephedraceae]]
          |1=''[[Biology:Ephedra (plant)|Ephedra]]''
           }}
         }}
      |label2='''[[Biology:Pinales|Pinales]]'''
      |2={{Clade
        |label1=[[Biology:Pinaceae|Pinaceae]]
        |1={{Clade
          |label1=Abietoideae
          |1={{Clade
            |1=''[[Biology:Cedrus|Cedrus]]''
            |2={{Clade
              |1={{Clade
                |1=''[[Biology:Pseudolarix|Pseudolarix]]''
                |2={{Clade
                  |1=''[[Biology:Nothotsuga|Nothotsuga]]''
                  |2=''[[Biology:Tsuga|Tsuga]]''
                   }}
                 }}
              |2={{Clade
                |1=''[[Biology:Keteleeria|Keteleeria]]''
                |2=''[[Biology:Abies|Abies]]''
                 }}
               }}
             }}
          |label2=Pinoideae
          |2={{Clade
            |1={{Clade
              |1=''[[Biology:Pseudotsuga|Pseudotsuga]]''
              |2=''[[Biology:Larix|Larix]]''
               }}
            |2={{Clade
              |1={{Clade
                |1=''[[Biology:Cathaya|Cathaya]]''
                |2=''[[Biology:Picea|Picea]]''
                 }}
              |2=''[[Biology:Pinus|Pinus]]''
               }}
             }}
           }}
         }}
       }}
    |label2=
    |2={{Clade
      |label1='''Araucariales'''
      |1={{Clade
        |label1=[[Biology:Araucariaceae|Araucariaceae]]
        |1={{Clade
          |label1=Agathideae
          |1={{Clade
            |1=''[[Biology:Wollemia|Wollemia]]''
            |2=''[[Biology:Agathis|Agathis]]''
             }}
          |label2=Araucarieae
          |2=''[[Biology:Araucaria|Araucaria]]''
           }}
        |label2=[[Biology:Podocarpaceae|Podocarpaceae]]
        |2={{Clade
          |1={"Phyllocladoideae"}
          |2={"Podocarpoideae"}
           }}
         }}
      |label2='''Cupressales'''
      |2={{Clade
        |1={{Clade
          |label1=[[Biology:Sciadopityaceae|Sciadopityaceae]]
          |1=''[[Biology:Sciadopitys|Sciadopitys]]''
           }}
        |2={{Clade
          |label1=[[Biology:Taxaceae|Taxaceae]]
          |1={{Clade
            |label1=Cephalotaxeae
            |1=''[[Biology:Cephalotaxus|Cephalotaxus]]''
            |label2=Taxoideae
            |2={{Clade
              |1={{Clade
                |1=''[[Biology:Amentotaxus|Amentotaxus]]''
                |2=''[[Biology:Torreya|Torreya]]''
                 }}
              |2={{Clade
                |1=''[[Biology:Austrotaxus|Austrotaxus]]''
                |2={{Clade
                  |1=''[[Biology:Pseudotaxus|Pseudotaxus]]''
                  |2=''[[Biology:Taxus|Taxus]]''
                   }}
                 }}
               }}
             }}
          |label2=[[Biology:Cupressaceae|Cupressaceae]]
          |2={{Clade
            |1={{Clade
              |label1=Cunninghamioideae
              |1=''[[Biology:Cunninghamia|Cunninghamia]]''
               }}
            |2={{Clade
              |1={{Clade
                |label1=Taiwanioideae
                |1=''[[Biology:Taiwania|Taiwania]]''
                 }}
              |2={{Clade
                |1={{Clade
                  |label1=Athrotaxidoideae
                  |1=''[[Biology:Athrotaxis|Athrotaxis]]''
                   }}
                |2={{Clade
                  |1={{Clade
                    |label1=Sequoioideae
                    |1={{Clade
                      |1=''[[Biology:Metasequoia|Metasequoia]]''
                      |2={{Clade
                        |1=''[[Biology:Sequoiadendron|Sequoiadendron]]''
                        |2=''[[Biology:Sequoia (genus)|Sequoia]]''
                         }}
                       }}
                     }}
                  |2={{Clade
                    |1={{Clade
                      |label1=Taxodioideae
                      |1={{Clade
                        |1=''[[Biology:Cryptomeria|Cryptomeria]]''
                        |2={{Clade
                          |1=''[[Biology:Glyptostrobus|Glyptostrobus]]''
                          |2=''[[Biology:Taxodium|Taxodium]]''
                           }}
                         }}
                       }}
                    |2={{Clade
                      |1={"Actinostroboideae"}
                      |2={"Cupressoideae"}
                       }}
                     }}
                   }}
                 }}
               }}
             }}
           }}
         }}
       }}
     }}
   }}
   }}
|targetA={"Phyllocladoideae"}
|subcladeA={{Clade
|label1 =Phyllocladoideae
|1={{Clade hidden |id=1 |mode=left
  |1=''[[Biology:Halocarpus|Halocarpus]]''
  |2={{Clade
    |1={{Clade
      |1=''[[Biology:Pectinopitys|Pectinopitys]]''
      |2={{Clade
        |1=''[[Biology:Sundacarpus|Sundacarpus]]''
        |2=''[[Biology:Prumnopitys|Prumnopitys]]''
         }}
       }}
    |2={{Clade
      |1={{Clade
        |1=''[[Biology:Parasitaxus|Parasitaxus]]''
        |2={{Clade
          |1=''[[Biology:Lagarostrobos|Lagarostrobos]]''
          |2=''[[Biology:Manoao|Manoao]]''
           }}
         }}
      |2={{Clade
        |1=''[[Biology:Lepidothamnus|Lepidothamnus]]''
        |2=''[[Biology:Phyllocladus|Phyllocladus]]''
         }}
       }}
     }}
   }}
           }}
|targetB={"Podocarpoideae"}
|subcladeB={{Clade
|label1 =Podocarpoideae
|1={{Clade hidden |id=2 |mode=left
  |1=''[[Biology:Saxegothaea|Saxegothaea]]''
  |2={{Clade
    |1=''[[Biology:Microcachrys|Microcachrys]]''
    |2={{Clade
      |1=''[[Biology:Pherosphaera|Pherosphaera]]''
      |2={{Clade
        |1=''[[Biology:Acmopyle|Acmopyle]]''
        |2={{Clade
          |1={{Clade
            |1=''[[Biology:Dacrycarpus|Dacrycarpus]]''
            |2={{Clade
              |1=''[[Biology:Falcatifolium|Falcatifolium]]''
              |2=''[[Biology:Dacrydium|Dacrydium]]''
               }}
             }}
          |2={{Clade
            |1={{Clade
              |1=''[[Biology:Retrophyllum|Retrophyllum]]''
              |2={{Clade
                |1=''[[Biology:Afrocarpus|Afrocarpus]]''
                |2=''[[Biology:Nageia|Nageia]]''
                 }}
               }}
            |2=''[[Biology:Podocarpus|Podocarpus]]''
             }}
           }}
         }}
       }}
     }}
   }}
           }}
|targetC={"Actinostroboideae"}
|subcladeC={{Clade
|label1 =Actinostroboideae
|1={{Clade hidden |id=3 |mode=left
  |1=''[[Biology:Papuacedrus|Papuacedrus]]''
  |2={{Clade
    |1=''[[Biology:Austrocedrus|Austrocedrus]]''
    |2={{Clade
      |1={{Clade
        |1=''[[Biology:Pilgerodendron|Pilgerodendron]]''
        |2=''[[Biology:Libocedrus|Libocedrus]]''
         }}
      |2={{Clade
        |1={{Clade
          |1={{Clade
            |1=''[[Biology:Diselma|Diselma]]''
            |2=''[[Biology:Fitzroya|Fitzroya]]''
             }}
          |2=''[[Biology:Widdringtonia|Widdringtonia]]''
           }}
        |2={{Clade
          |1=''Neocallitropsis''
          |2=''[[Biology:Callitris|Callitris]]''
           }}
         }}
       }}
     }}
   }}
           }}
|targetD={"Cupressoideae"}
|subcladeD={{Clade
|label1 =Cupressoideae
|1={{Clade hidden |id=4 |mode=left
  |1={{Clade
    |1=''[[Biology:Thujopsis|Thujopsis]]''
    |2=''[[Biology:Thuja|Thuja]]''
     }}
  |2={{Clade
    |1=''[[Biology:Chamaecyparis|Chamaecyparis]]''
    |2={{Clade
      |1={{Clade
        |1={{Clade
          |1=''[[Biology:Tetraclinis|Tetraclinis]]''
          |2={{Clade
            |1=''[[Biology:Microbiota decussata|Microbiota]]''
            |2=''[[Biology:Platycladus|Platycladus]]''
             }}
           }}
        |2=''[[Biology:Calocedrus|Calocedrus]]''
         }}
      |2={{Clade
        |1={{Clade
          |1=''[[Biology:Cupressus|Cupressus]]''
          |2={{Clade
            |1=''[[Biology:Xanthocyparis|Xanthocyparis]]''
            |2=''[[Biology:Hesperocyparis|Hesperocyparis]]''
             }}
           }}
        |2=''[[Biology:Juniperus|Juniperus]]''
         }}
       }}
     }}
   }}
           }}
}}
}}

{{further|Biology:Spermatophyte}}
A formal classification of the living gymnosperms is the "Acrogymnospermae", which form a monophyletic group within the [[Biology:Spermatophyte|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|pages=55–70|doi=10.11646/phytotaxa.19.1.3|s2cid=86797396 }}</ref> The wider "Gymnospermae" group includes extinct gymnosperms and is thought to be [[Biology: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 [[Biology:Fern|fern]]-like vegetative morphology (the so-called "seed ferns" or pteridosperms).<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 [[Biology:Bennettitales|Bennettitales]], glossopterids, and ''[[Biology:Caytonia|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 |author1=Christenhusz, M. J. M. |author2=Byng, J. W. | year = 2016 | title = The number of known plants species in the world and its annual increase | journal = Phytotaxa | volume = 261 | pages = 201–217 | url = http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 | doi = 10.11646/phytotaxa.261.3.1 | issue = 3 | doi-access = free }}</ref>

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

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

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

'''Subclass [[Biology:Pinidae|Pinidae]]'''
*Order '''[[Biology:Pinales|Pinales]]'''
**Family '''[[Biology:Pinaceae|Pinaceae]]''': ''[[Biology:Cedrus|Cedrus]]'', ''[[Biology:Pinus|Pinus]]'', ''[[Biology:Cathaya|Cathaya]]'', ''[[Biology:Picea|Picea]]'', ''[[Biology:Pseudotsuga|Pseudotsuga]]'', ''[[Biology:Larix|Larix]]'', ''[[Biology:Pseudolarix|Pseudolarix]]'', ''[[Biology:Tsuga|Tsuga]]'', ''[[Biology:Nothotsuga|Nothotsuga]]'', ''[[Biology:Keteleeria|Keteleeria]]'', ''[[Biology:Abies|Abies]]''

*Order '''Araucariales'''
**Family '''[[Biology:Araucariaceae|Araucariaceae]]''': ''[[Biology:Araucaria|Araucaria]]'', ''[[Biology:Wollemia|Wollemia]]'', ''[[Biology:Agathis|Agathis]]''
**Family '''[[Biology:Podocarpaceae|Podocarpaceae]]''': ''[[Biology:Phyllocladus|Phyllocladus]]'', ''[[Biology:Lepidothamnus|Lepidothamnus]]'', ''[[Biology:Prumnopitys|Prumnopitys]]'', ''[[Biology:Sundacarpus|Sundacarpus]]'', ''[[Biology:Halocarpus|Halocarpus]]'', ''[[Biology:Parasitaxus|Parasitaxus]]'', ''[[Biology:Lagarostrobos|Lagarostrobos]]'', ''[[Biology:Manoao|Manoao]]'', ''[[Biology:Saxegothaea|Saxegothaea]]'', ''[[Biology:Microcachrys|Microcachrys]]'', ''[[Biology:Pherosphaera|Pherosphaera]]'', ''[[Biology:Acmopyle|Acmopyle]]'', ''[[Biology:Dacrycarpus|Dacrycarpus]]'', ''[[Biology:Dacrydium|Dacrydium]]'', ''[[Biology:Falcatifolium|Falcatifolium]]'', ''[[Biology:Retrophyllum|Retrophyllum]]'', ''[[Biology:Nageia|Nageia]]'', ''[[Biology:Afrocarpus|Afrocarpus]]'', ''[[Biology:Podocarpus|Podocarpus]]''

*Order '''Cupressales'''
**Family '''[[Biology:Sciadopityaceae|Sciadopityaceae]]''': ''[[Biology:Sciadopitys|Sciadopitys]]''
**Family '''[[Biology:Cupressaceae|Cupressaceae]]''': ''[[Biology:Cunninghamia|Cunninghamia]]'', ''[[Biology:Taiwania|Taiwania]]'', ''[[Biology:Athrotaxis|Athrotaxis]]'', ''[[Biology:Metasequoia|Metasequoia]]'', ''[[Biology:Sequoioideae|Sequoia]]'', ''[[Biology:Sequoiadendron|Sequoiadendron]]'', ''[[Biology:Cryptomeria|Cryptomeria]]'', ''[[Biology:Glyptostrobus|Glyptostrobus]]'', ''[[Biology:Taxodium|Taxodium]]'', ''[[Biology:Papuacedrus|Papuacedrus]]'', ''[[Biology:Austrocedrus|Austrocedrus]]'', ''[[Biology:Libocedrus|Libocedrus]]'', ''[[Biology:Pilgerodendron|Pilgerodendron]]'', ''[[Biology:Widdringtonia|Widdringtonia]]'', ''[[Biology:Diselma|Diselma]]'', ''[[Biology:Fitzroya|Fitzroya]]'', ''[[Biology:Callitris|Callitris]]'', ''[[Biology:Actinostrobus|Actinostrobus]]'', ''Neocallitropsis'', ''[[Biology:Thujopsis|Thujopsis]]'', ''[[Biology:Thuja|Thuja]]'', ''[[Biology:Fokienia|Fokienia]]'', ''[[Biology:Chamaecyparis|Chamaecyparis]]'',  ''[[Biology:Cupressus|Cupressus]]'', ''[[Biology:Juniperus|Juniperus]]'', ''[[Biology:Calocedrus|Calocedrus]]'', ''[[Biology:Tetraclinis|Tetraclinis]]'', ''[[Biology:Platycladus|Platycladus]]'', ''[[Biology:Microbiota (plant)|Microbiota]]''
**Family '''[[Biology:Taxaceae|Taxaceae]]''': ''[[Biology:Austrotaxus|Austrotaxus]]'', ''[[Biology:Pseudotaxus|Pseudotaxus]]'', ''[[Biology:Taxus|Taxus]]'', ''[[Biology:Cephalotaxus|Cephalotaxus]]'', ''[[Biology:Amentotaxus|Amentotaxus]]'', ''[[Biology:Torreya|Torreya]]''

=== Extinct groupings ===
*Order [[Biology:Cordaitales|Cordaitales]]
*Order [[Biology:Calamopityales|Calamopityales]]
* Order [[Biology:Callistophytales|Callistophytales]]
* Order [[Biology:Caytoniales|Caytoniales]]
* Order [[Biology:Gigantopteridales|Gigantopteridales]]
* Order [[Biology:Glossopteridales|Glossopteridales]]
* Order [[Biology:Lyginopteridales|Lyginopteridales]]
* Order [[Biology:Medullosales|Medullosales]]
* Order [[Biology:Peltaspermales|Peltaspermales]]
* Order [[Biology:Corystospermales|Corystospermales]] (also known as Umkomasiales)
* Order [[Biology:Czekanowskiales|Czekanowskiales]]
* Order [[Biology:Bennettitales|Bennettitales]] (cycadeoids''')'''
* Order [[Biology:Erdtmanithecales|Erdtmanithecales]]
*Order [[Biology:Pentoxylales|Pentoxylales]]
*Order [[Biology:Czekanowskiales|Czekanowskiales]]
*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 [[Biology:Gametophyte|gametophyte]] (gamete-bearing phase) is relatively short-lived. Like all seed plants, they are [[Biology:Heterospory|heterosporous]], having two spore types, [[Biology:Microspore|microspores]] (male) and [[Biology:Megaspore|megaspores]] (female) that are typically produced in pollen cones or ovulate cones, respectively.<ref name="Concepts of Biology">{{cite book |last1=Samantha |first1=Fowler |last2=Rebecca |first2=Roush |last3=James |first3=Wise |title=Concepts of Biology |date=2013 |publisher=OpenStax |location=Houston, Texas |url=https://openstax.org/books/concepts-biology/pages/14-3-seed-plants-gymnosperms |access-date=31 March 2023 |chapter=14.3 Seed Plants: Gymnosperms}}</ref> The exception is the females in the cycad genus ''[[Biology:Cycas|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|doi=10.1038/s41477-022-01129-7|pmid=35437001 |pmc=9023351 |issn=2055-0278}}</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 [[Biology:Archegonium|archegonia]], which produce the female gamete. 

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 ([[Biology:Integument|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 ''[[Biology:Ginkgo|Ginkgo]]'' have [[Biology:Flagellum|flagellated]] motile sperm<ref name="AJB">{{cite journal |last1=Southworth |first1=Darlene |last2=Cresti |first2=Mauro |title=Comparison of flagellated and nonflagellated sperm in plants |url=https://bsapubs.onlinelibrary.wiley.com/doi/pdf/10.2307/2446056 |journal=American Journal of Botany |access-date=26 March 2022 |pages=1301–1311 |doi=10.2307/2446056 |date=September 1997|volume=84 |issue=9 |jstor=2446056 |pmid=21708687 |doi-access=free }}</ref> that swim directly to the egg inside the ovule, whereas conifers and gnetophytes have sperm with no flagella that are moved along a [[Biology:Pollen tube|pollen tube]] to the egg. After [[Biology: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 [[Biology:Gametophyte#Seed plants|gametophyte]], which serves as a food supply, and the seed coat.<ref name="Walters1996">{{Cite book| last = Walters | first =  Dirk R Walters Bonnie By| year = 1996| title = Vascular plant taxonomy| page = [https://archive.org/details/vascularplanttax00walt/page/124 124]| url = https://archive.org/details/vascularplanttax00walt| url-access = registration | quote = Gymnosperm seeds. | isbn = 978-0-7872-2108-9| publisher = Kendall/Hunt Pub. Co.| location = Dubuque, Iowa}}</ref>

==Genetics==

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

==Uses==

Gymnosperms have major economic uses.  Pine, fir, spruce, and cedar are all examples of conifers that are used for [[Engineering:Lumber|lumber]], paper production, and resin. Some other common uses for gymnosperms are [[Chemistry:Soap|soap]], varnish, nail polish, food, gum, and perfumes.<ref>{{cite book |last1=Biswas |first1=C. |last2=Johri |first2=B.M. |year=1997 |chapter=Economic Importance |title=The Gymnosperms |pages=440–456 |publisher=Springer, Berlin, Heidelberg |doi=10.1007/978-3-662-13164-0_23 |isbn=978-3-662-13166-4 |url=https://link.springer.com/content/pdf/10.1007%2F978-3-662-13164-0.pdf}}</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|title=Recent advances on phylogenomics of gymnosperms and a new classification|journal=Plant Diversity|date=2020|volume=44|issue=4|pages=340–350|doi=10.1016/j.pld.2022.05.003|pmid=35967253 |pmc=9363647 |s2cid=249117306 |issn=2468-2659}}</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. |title=Towards a phylogenetic nomenclature of Tracheophyta |journal=Taxon |date=August 2007 |volume=56 |issue=3 |pages=822–846 |doi=10.2307/25065864|jstor=25065864 |ref={{harvid|Cantino|2007}}}}
{{refend}}

==External links==
*[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]
{{Clear}}
{{Botany}}
{{Plant classification}}
{{Taxonbar|from=Q133712}}
[[Category:Gymnosperms| 01]]

{{Sourceattribution|Gymnosperm}}