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=== Floral composition === ==== Conifers ==== [[File:Petrified Araucaria cone.jpg|thumb|Petrified ''[[Araucaria mirabilis]]'' cone from the Middle Jurassic of Argentina|alt=Two images of a round conifer cone, the left one is in cross-section]] [[Conifer]]s formed a dominant component of Jurassic floras. The Late Triassic and Jurassic was a major time of diversification of conifers, with most modern conifer groups appearing in the fossil record by the end of the Jurassic, having evolved from [[Voltziales|voltzialean]] ancestors.<ref>{{Cite journal|last1=Atkinson|first1=Brian A.|last2=Serbet|first2=Rudolph|last3=Hieger|first3=Timothy J.|last4=Taylor|first4=Edith L.|date=October 2018|title=Additional evidence for the Mesozoic diversification of conifers: Pollen cone of Chimaerostrobus minutus gen. et sp. nov. (Coniferales), from the Lower Jurassic of Antarctica|journal=Review of Palaeobotany and Palynology|language=en|volume=257|pages=77–84|doi=10.1016/j.revpalbo.2018.06.013|bibcode=2018RPaPa.257...77A |s2cid=133732087|doi-access=free}}</ref><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|url=http://doi.wiley.com/10.1002/ajb2.1143|journal=American Journal of Botany|language=en|volume=105|issue=9|pages=1531–1544|doi=10.1002/ajb2.1143|pmid=30157290|s2cid=52120430 }}</ref> [[Araucariaceae|Araucarian]] conifers have their first unambiguous records during the Early Jurassic, and members of the modern genus ''[[Araucaria]]'' were widespread across both hemispheres by the Middle Jurassic.<ref name="Leslie-2018" /><ref>{{Cite journal|last1=Stockey|first1=Ruth A.|last2=Rothwell|first2=Gar W.|date=July 2020|title=Diversification of crown group Araucaria : the role of Araucaria famii sp. nov. in the Late Cretaceous (Campanian) radiation of Araucariaceae in the Northern Hemisphere|journal=American Journal of Botany|language=en|volume=107|issue=7|pages=1072–1093|doi=10.1002/ajb2.1505|pmid=32705687|s2cid=225568264|issn=0002-9122|doi-access=free}}</ref><ref>{{Cite journal|last1=Escapa|first1=Ignacio H.|last2=Catalano|first2=Santiago A.|date=October 2013|title=Phylogenetic Analysis of Araucariaceae: Integrating Molecules, Morphology, and Fossils|url=https://www.journals.uchicago.edu/doi/10.1086/672369|journal=International Journal of Plant Sciences|language=en|volume=174|issue=8|pages=1153–1170|doi=10.1086/672369|bibcode=2013IJPlS.174.1153E |hdl=11336/3583|s2cid=56238574|issn=1058-5893|hdl-access=free}}</ref> Also abundant during the Jurassic is the extinct family [[Cheirolepidiaceae]], often recognised through their highly distinctive ''Classopolis'' pollen. Jurassic representatives include the pollen cone ''Classostrobus'' and the seed cone ''[[Pararaucaria]]''. Araucarian and Cheirolepidiaceae conifers often occur in association.<ref>{{Cite journal|last1=Stockey|first1=Ruth A.|last2=Rothwell|first2=Gar W.|date=March 2013|title=Pararaucaria carrii sp. nov., Anatomically Preserved Evidence for the Conifer Family Cheirolepidiaceae in the Northern Hemisphere|url=https://www.journals.uchicago.edu/doi/10.1086/668614|journal=International Journal of Plant Sciences|language=en|volume=174|issue=3|pages=445–457|doi=10.1086/668614|bibcode=2013IJPlS.174..445S |s2cid=59269291|issn=1058-5893}}</ref> The oldest definitive record of the cypress family ([[Cupressaceae]]) is ''[[Austrohamia|Austrohamia minuta]]'' from the Early Jurassic (Pliensbachian) of Patagonia, known from many parts of the plant.<ref>{{Cite journal|last1=Escapa|first1=Ignacio|last2=Cúneo|first2=Rubén|last3=Axsmith|first3=Brian|date=September 2008|title=A new genus of the Cupressaceae (sensu lato) from the Jurassic of Patagonia: Implications for conifer megasporangiate cone homologies|url=https://linkinghub.elsevier.com/retrieve/pii/S0034666708000481|journal=Review of Palaeobotany and Palynology|language=en|volume=151|issue=3–4|pages=110–122|doi=10.1016/j.revpalbo.2008.03.002|bibcode=2008RPaPa.151..110E }}</ref> The reproductive structures of ''Austrohamia'' have strong similarities to those of the primitive living cypress genera ''[[Taiwania]]'' and ''[[Cunninghamia]].'' By the Middle to Late Jurassic Cupressaceae were abundant in warm temperate–tropical regions of the Northern Hemisphere, most abundantly represented by the genus ''[[Elatides]]''.<ref>{{Cite journal|last1=Contreras|first1=Dori L.|last2=Escapa|first2=Ignacio H.|last3=Iribarren|first3=Rocio C.|last4=Cúneo|first4=N. Rubén|date=October 2019|title=Reconstructing the Early Evolution of the Cupressaceae: A Whole-Plant Description of a New Austrohamia Species from the Cañadón Asfalto Formation (Early Jurassic), Argentina|url=https://www.journals.uchicago.edu/doi/10.1086/704831|journal=International Journal of Plant Sciences|language=en|volume=180|issue=8|pages=834–868|doi=10.1086/704831|bibcode=2019IJPlS.180..834C |s2cid=202862782|issn=1058-5893}}</ref> The Jurassic also saw the first appearances of some modern genera of cypresses, such as ''[[Sequoia (genus)|Sequoia]]''.<ref name="Jehol">{{cite journal |last1=Ma |first1=Qing-Wen |last2=K. Ferguson |first2=David |last3=Liu |first3=Hai-Ming |last4=Xu |first4=Jing-Xian |date=2020 |title=Compressions of Sequoia (Cupressaceae sensu lato) from the Middle Jurassic of Daohugou, Ningcheng, Inner Mongolia, China |url=https://link.springer.com/article/10.1007/s12549-020-00454-z |journal=Palaeobiodiversity and Palaeoenvironments |volume=1 |issue=9 |page=1 |doi=10.1007/s12549-020-00454-z |s2cid=227180592 |access-date=9 March 2021 |ref=Jehol}}</ref> Members of the extinct genus ''[[Schizolepidopsis]]'' which likely represent a [[stem-group]] to the pine family ([[Pinaceae]]), were widely distributed across Eurasia during the Jurassic.<ref>{{Cite journal|last1=Domogatskaya|first1=Ksenia V.|last2=Herman|first2=Alexei B.|date=May 2019|title=New species of the genus Schizolepidopsis (conifers) from the Albian of the Russian high Arctic and geological history of the genus|url=https://linkinghub.elsevier.com/retrieve/pii/S0195667118304257|journal=Cretaceous Research|language=en|volume=97|pages=73–93|doi=10.1016/j.cretres.2019.01.012|bibcode=2019CrRes..97...73D |s2cid=134849082}}</ref><ref name="Matsunaga-2021">{{Cite journal|last1=Matsunaga|first1=Kelly K. S.|last2=Herendeen|first2=Patrick S.|last3=Herrera|first3=Fabiany|last4=Ichinnorov|first4=Niiden|last5=Crane|first5=Peter R.|last6=Shi|first6=Gongle|date=2021-05-10|title=Ovulate Cones of Schizolepidopsis ediae sp. nov. Provide Insights into the Evolution of Pinaceae|journal=International Journal of Plant Sciences|volume=182|issue=6|pages=490–507|doi=10.1086/714281|s2cid=235426888|issn=1058-5893|doi-access=free|bibcode=2021IJPlS.182..490M }}</ref> The oldest unambiguous record of Pinaceae is the [[Conifer cone|pine cone]] ''[[Eathiestrobus]]'', known from the Late Jurassic (Kimmeridgian) of Scotland, which remains the only known unequivocal fossil of the group before the Cretaceous.<ref>{{Cite journal|last1=Rothwell|first1=Gar W.|last2=Mapes|first2=Gene|last3=Stockey|first3=Ruth A.|last4=Hilton|first4=Jason|date=April 2012|title=The seed cone Eathiestrobus gen. nov.: Fossil evidence for a Jurassic origin of Pinaceae|journal=American Journal of Botany|language=en|volume=99|issue=4|pages=708–720|doi=10.3732/ajb.1100595|pmid=22491001|bibcode=2012AmJB...99..708R }}</ref> Despite being the earliest known member of the Pinaceae, ''Eathiestrobus'' appears to be a member of the [[Pinoideae|pinoid]] [[clade]] of the family, suggesting that the initial diversification of Pinaceae occurred earlier than has been found in the fossil record.<ref>{{Cite journal|last1=Smith|first1=Selena Y.|last2=Stockey|first2=Ruth A.|last3=Rothwell|first3=Gar W.|last4=Little|first4=Stefan A.|date=2017-01-02|title=A new species of Pityostrobus (Pinaceae) from the Cretaceous of California: moving towards understanding the Cretaceous radiation of Pinaceae|url=https://www.tandfonline.com/doi/full/10.1080/14772019.2016.1143885|journal=Journal of Systematic Palaeontology|language=en|volume=15|issue=1|pages=69–81|doi=10.1080/14772019.2016.1143885|bibcode=2017JSPal..15...69S |s2cid=88292891|issn=1477-2019}}</ref><ref name="Leslie-2018" /> The earliest record of the yew family ([[Taxaceae]]) is ''[[Palaeotaxus|Palaeotaxus rediviva]]'', from the Hettangian of Sweden, suggested to be closely related to the living ''[[Austrotaxus]]'', while ''[[Marskea|Marskea jurassica]]'' from the Middle Jurassic of Yorkshire, England and material from the Callovian–Oxfordian [[Daohugou Bed]] in China are thought to be closely related to ''[[Amentotaxus]],'' with the latter material assigned to the modern genus, indicating that Taxaceae had substantially diversified by the end of the Jurassic.<ref>{{Cite journal|last1=Dong|first1=Chong|last2=Shi|first2=Gongle|last3=Herrera|first3=Fabiany|last4=Wang|first4=Yongdong|last5=Herendeen|first5=Patrick S|last6=Crane|first6=Peter R|date=2020-06-18|title=Middle–Late Jurassic fossils from northeastern China reveal morphological stasis in the catkin-yew|journal=National Science Review|volume=7|issue=11|language=en|pages=1765–1767|doi=10.1093/nsr/nwaa138|pmid=34691509|pmc=8288717|issn=2095-5138|doi-access=free}}</ref> The oldest unambiguous members of [[Podocarpaceae]] are known from the Jurassic, found across both hemispheres, including ''[[Scarburgia]]'' and ''[[Harrisiocarpus]]'' from the Middle Jurassic of England, as well as unnamed species from the Middle-Late Jurassic of Patagonia.<ref>{{Cite journal |last1=Andruchow-Colombo |first1=Ana |last2=Escapa |first2=Ignacio H |last3=Aagesen |first3=Lone |last4=Matsunaga |first4=Kelly K S |date=2023-08-04 |title=In search of lost time: tracing the fossil diversity of Podocarpaceae through the ages |url=https://academic.oup.com/botlinnean/advance-article/doi/10.1093/botlinnean/boad027/7237351 |journal=Botanical Journal of the Linnean Society |volume=203 |issue=4 |pages=315–336 |language=en |doi=10.1093/botlinnean/boad027 |issn=0024-4074|hdl=11336/227952 |hdl-access=free }}</ref> During the Early Jurassic, the flora of the mid-latitudes of Eastern Asia were dominated by the extinct deciduous broad leafed conifer ''[[Podozamites]],'' which appears to not be closely related to any living family of conifer. Its range extended northwards into polar latitudes of Siberia and then contracted northward in the Middle to Late Jurassic, corresponding to the increasing aridity of the region.<ref>{{Cite journal |last1=Pole |first1=Mike |last2=Wang |first2=Yongdong |last3=Bugdaeva |first3=Eugenia V. |last4=Dong |first4=Chong |last5=Tian |first5=Ning |last6=Li |first6=Liqin |last7=Zhou |first7=Ning |date=2016-12-15 |title=The rise and demise of Podozamites in east Asia—An extinct conifer life style |url=http://www.sciencedirect.com/science/article/pii/S0031018216001188 |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |series=Mesozoic ecosystems – Climate and Biota |language=en |volume=464 |pages=97–109 |bibcode=2016PPP...464...97P |doi=10.1016/j.palaeo.2016.02.037 |issn=0031-0182}}</ref> ==== Ginkgoales ==== [[File:Ginkgo huttoni 1.jpg|thumb|upright|Leaves of ''[[Ginkgo huttonii]]'' from the Middle Jurassic of England|alt=Black fossils of leaves on yellow coloured rock|left]] [[Ginkgoales]], of which the sole living species is ''[[Ginkgo biloba]]'', were more diverse during the Jurassic: they were among the most important components of Eurasian Jurassic floras and were adapted to a wide variety of climatic conditions.<ref name="Zhou-2009" /> The earliest representatives of the genus ''[[Ginkgo]],'' represented by [[Ovule|ovulate]] and pollen organs similar to those of the modern species, are known from the Middle Jurassic in the Northern Hemisphere.<ref name="Zhou-2009" /> Several other lineages of ginkgoaleans are known from Jurassic rocks, including ''[[Yimaia]]'', ''[[Grenana]]'', ''[[Nagrenia]]'' and [[Karkeniaceae|''Karkenia'']]. These lineages are associated with ''Ginkgo-''like leaves, but are distinguished from living and fossil representatives of ''Ginkgo'' by having differently arranged reproductive structures.<ref name="Zhou-2009">{{Cite journal|last=Zhou|first=Zhi-Yan|date=March 2009|title=An overview of fossil Ginkgoales|url=https://linkinghub.elsevier.com/retrieve/pii/S1871174X0900002X|journal=Palaeoworld|language=en|volume=18|issue=1|pages=1–22|doi=10.1016/j.palwor.2009.01.001|quote=}}</ref><ref>{{Cite journal|last=Nosova|first=Natalya|date=October 2013|title=Revision of the genus Grenana Samylina from the Middle Jurassic of Angren, Uzbekistan|url=https://linkinghub.elsevier.com/retrieve/pii/S0034666713001115|journal=Review of Palaeobotany and Palynology|language=en|volume=197|pages=226–252|doi=10.1016/j.revpalbo.2013.06.005|bibcode=2013RPaPa.197..226N }}</ref> ''[[Umaltolepis]]'' from the Jurassic of Asia has strap-shaped ginkgo-like leaves with highly distinct reproductive structures with similarities to those of peltasperm and corystosperm seed ferns, has been suggested to be a member of Ginkgoales ''sensu lato.<ref name="Dong-2022">{{Cite journal |last1=Dong |first1=Chong |last2=Shi |first2=Gongle |last3=Zhang |first3=Xiaoqing |last4=Wang |first4=Zixi |last5=Wang |first5=Yongdong |date=November 2022 |title=Middle-Late Jurassic fossils from Northeast China confirm the affiliation of Umaltolepis seed-bearing structures and Pseudotorellia leaves |url=https://linkinghub.elsevier.com/retrieve/pii/S0034666722001610 |journal=Review of Palaeobotany and Palynology |language=en |volume=306 |pages=104763 |doi=10.1016/j.revpalbo.2022.104763|bibcode=2022RPaPa.30604763D |s2cid=251917169 }}</ref>'' ==== Bennettitales ==== [[File:Williamsoniaceae Thérèse Ekblom.webp|thumb|upright=1.15|Restoration of a member of [[Bennettitales]] belonging to Williamsoniaceae.]] [[Bennettitales]], having first become widespread during the preceding Triassic, were diverse and abundant members of Jurassic floras across both hemispheres.<ref name="Popa-2014">{{Cite journal|last=Popa|first=Mihai E.|date=June 2014|title=Early Jurassic bennettitalean reproductive structures of Romania|url=http://link.springer.com/10.1007/s12549-014-0165-9|journal=Palaeobiodiversity and Palaeoenvironments |language=en |volume=94|issue=2|pages=327–362|doi=10.1007/s12549-014-0165-9|bibcode=2014PdPe...94..327P |s2cid=128411467|issn=1867-1594}}</ref> The foliage of Bennettitales bears strong similarities to those of cycads, to such a degree that they cannot be reliably distinguished on the basis of morphology alone. Leaves of Bennettitales can be distinguished from those of cycads their different arrangement of [[stoma]]ta, and the two groups are not thought to be closely related.<ref>{{Citation|last=Taylor|first=T|title=Cycadophytes|date=2009 |url=https://linkinghub.elsevier.com/retrieve/pii/B9780123739728000176|work=Biology and Evolution of Fossil Plants|pages=703–741 |publisher=Elsevier|language=en|doi=10.1016/b978-0-12-373972-8.00017-6|isbn=978-0-12-373972-8|access-date=2020-12-12}}</ref> Jurassic Bennettitales predominantly belong to the group [[Williamsoniaceae]],<ref name="Popa-2014" /> which grew as shrubs and small trees. The Williamsoniaceae are thought to have had a [[divaricate]] branching habit, similar to that of living ''[[Banksia]]'', and adapted to growing in open habitats with poor soil nutrient conditions.<ref name="Pott-2014">{{Cite journal|last1=Pott|first1=Christian|last2=McLoughlin|first2=Stephen |date=2014-06-01|title=Divaricate growth habit in Williamsoniaceae (Bennettitales): unravelling the ecology of a key Mesozoic plant group |url=https://doi.org/10.1007/s12549-014-0157-9|journal=Palaeobiodiversity and Palaeoenvironments|language=en|volume=94|issue=2|pages=307–325 |doi=10.1007/s12549-014-0157-9|bibcode=2014PdPe...94..307P |s2cid=84440045|issn=1867-1608}}</ref> Bennettitales exhibit complex, [[flower]]-like reproductive structures some of which are thought to have been pollinated by insects. Several groups of insects that bear long proboscis, including extinct families such as [[Kalligrammatidae|kalligrammatid]] lacewings<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. |date=2016-02-10|title=The evolutionary convergence of mid-Mesozoic lacewings and Cenozoic butterflies|url= |journal=Proceedings of the Royal Society B: Biological Sciences|language=en|volume=283|issue=1824|pages=20152893|doi=10.1098/rspb.2015.2893|issn=0962-8452 |pmc=4760178 |pmid=26842570}}</ref> and extant ones such as [[Acroceridae|acrocerid]] flies,<ref>{{Cite journal|last1=Khramov|first1=Alexander V.|author-link1=Alexander V. Khramov|last2=Lukashevich|first2=Elena D.|date=July 2019|title=A Jurassic dipteran pollinator with an extremely long proboscis |url=https://linkinghub.elsevier.com/retrieve/pii/S1342937X1930070X|journal=Gondwana Research|language=en|volume=71|pages=210–215 |doi=10.1016/j.gr.2019.02.004|bibcode=2019GondR..71..210K|s2cid=134847380}}</ref> are suggested to have been pollinators of bennettitales, feeding on [[nectar]] produced by bennettitalean cones. ==== Cycads ==== [[Cycad]]s reached their apex of diversity during the Jurassic and Cretaceous Periods.<ref name="Cai-2018">{{Cite journal |last1=Cai |first1=Chenyang |last2=Escalona |first2=Hermes E. |last3=Li |first3=Liqin |last4=Yin |first4=Ziwei |last5=Huang |first5=Diying |last6=Engel |first6=Michael S. |date=September 2018 |title=Beetle Pollination of Cycads in the Mesozoic |journal=Current Biology |language=en |volume=28 |issue=17 |pages=2806–2812.e1 |doi=10.1016/j.cub.2018.06.036 |pmid=30122529|s2cid=52038878|doi-access=free |bibcode=2018CBio...28E2806C }}</ref> Despite the Mesozoic sometimes being called the "Age of Cycads", cycads are thought to have been a relatively minor component of mid-Mesozoic floras, with the Bennettitales and [[Nilssoniales]], which have cycad-like foliage, being dominant.<ref name="Coiro-2017">{{Cite journal |last1=Coiro |first1=Mario |last2=Pott |first2=Christian |date=December 2017|title=Eobowenia gen. nov. from the Early Cretaceous of Patagonia: indication for an early divergence of Bowenia? |journal=BMC Evolutionary Biology |language=en |volume=17 |issue=1 |pages=97 |doi=10.1186/s12862-017-0943-x|issn=1471-2148 |pmc=5383990 |pmid=28388891 |doi-access=free |bibcode=2017BMCEE..17...97C }}</ref> The Nilssoniales have often been considered cycads or cycad relatives, but have been found to be distinct on chemical grounds, and perhaps more closely allied with Bennettitales.''<ref name="Vajda-2017">{{Cite journal |last1=Vajda |first1=Vivi |last2=Pucetaite |first2=Milda |last3=McLoughlin |first3=Stephen |last4=Engdahl |first4=Anders |last5=Heimdal |first5=Jimmy |last6=Uvdal |first6=Per |date=August 2017 |title=Molecular signatures of fossil leaves provide unexpected new evidence for extinct plant relationships |url=https://www.nature.com/articles/s41559-017-0224-5 |journal=Nature Ecology & Evolution |language=en |volume=1 |issue=8 |pages=1093–1099 |doi=10.1038/s41559-017-0224-5 |issn=2397-334X |pmid=29046567 |bibcode=2017NatEE...1.1093V |s2cid=3604369}}</ref>'' The relationships of most Mesozoic cycads to living groups are ambiguous,<ref name="Coiro-2017" /> with no Jurassic cycads belonging to either of the two modern groups of cycads, though some Jurassic cycads possibly represent [[stem-group]] relatives of modern [[Cycadaceae]], like the leaf genus ''[[Paracycas]]'' known Europe, and [[Zamiaceae]], like some European species of the leaf genus ''[[Pseudoctenis]]''. Also widespread during the Jurassic was the extinct ''[[Ctenis]]'' lineage, which appears to be distantly related to modern cycads.<ref>{{Cite journal |last1=Coiro |first1=Mario |last2=Allio |first2=Rémi |last3=Mazet |first3=Nathan |last4=Seyfullah |first4=Leyla J. |last5=Condamine |first5=Fabien L. |date=2023-06-11 |title=Reconciling fossils with phylogenies reveals the origin and macroevolutionary processes explaining the global cycad biodiversity |journal=New Phytologist |volume=240 |issue=4 |pages=1616–1635 |language=en |doi=10.1111/nph.19010 |pmid=37302411 |s2cid=259137975 |issn=0028-646X|doi-access=free |pmc=10953041 |bibcode=2023NewPh.240.1616C }}</ref> Modern cycads are pollinated by beetles, and such an association is thought to have formed by the Early Jurassic.<ref name="Cai-2018" /> ==== Other seed plants ==== Although there have been several claimed records, there are no widely accepted Jurassic fossil records of [[flowering plant]]s, which make up 90% of living plant species, and fossil evidence suggests that the group diversified during the following Cretaceous.<ref>{{Cite journal|last=Bateman|first=Richard M|date=2020-01-01|editor-last=Ort|editor-first=Donald|title=Hunting the Snark: the flawed search for mythical Jurassic angiosperms|url=https://academic.oup.com/jxb/article/71/1/22/5571867|journal=Journal of Experimental Botany|language=en|volume=71|issue=1|pages=22–35|doi=10.1093/jxb/erz411|issn=0022-0957|pmid=31538196}}</ref> The earliest known [[gnetophytes]], one of the four main living groups of [[gymnosperm]]s, appeared by the end of the Jurassic, with the oldest unequivocal gnetophyte being the seed ''[[Dayvaultia]]'' from the Late Jurassic of North America.<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=2022-12-14 |title=Cutting the long branches: Consilience as a path to unearth the evolutionary history of Gnetales |journal=Frontiers in Ecology and Evolution |volume=10 |pages=1082639 |doi=10.3389/fevo.2022.1082639 |issn=2296-701X |doi-access=free |bibcode=2022FrEEv..1082639C }}</ref>[[File:Sagenopteris phillipsi Natural History Museum v18596 Retallack 1980.jpg|thumb|''[[Sagenopteris phillipsii|Sagenopteris phillipsi]]'' ([[Caytoniales]]) from the Middle Jurassic of Yorkshire, England|alt=A fossil leaf, four elongate leaves branch off from the same point on the stem]] "Seed ferns" ([[Pteridospermatophyta]]) is a collective term to refer to disparate lineages of fern like plants that produce seeds but have uncertain affinities to living [[Spermatophyte|seed plant]] groups. A prominent group of Jurassic seed ferns is the [[Caytoniales]], which reached their zenith during the Jurassic, with widespread records in the Northern Hemisphere, though records in the Southern Hemisphere remain rare. Due to their [[berry]]-like seed-bearing capsules, they have often been suggested to have been closely related or perhaps ancestral to flowering plants, but the evidence for this is inconclusive.<ref>{{Cite journal|last1=Elgorriaga|first1=Andrés|last2=Escapa|first2=Ignacio H.|last3=Cúneo|first3=N. Rubén|date=2019-09-02|title=Southern Hemisphere Caytoniales: vegetative and reproductive remains from the Lonco Trapial Formation (Lower Jurassic), Patagonia|url=https://www.tandfonline.com/doi/full/10.1080/14772019.2018.1535456|journal=Journal of Systematic Palaeontology|language=en|volume=17|issue=17|pages=1477–1495|doi=10.1080/14772019.2018.1535456|bibcode=2019JSPal..17.1477E |s2cid=92287804|issn=1477-2019}}</ref> [[Corystospermaceae|Corystosperm]]-aligned seed ferns, such as ''[[Pachypteris]]'' and ''[[Komlopteris]]'' were widespread across both hemispheres during the Jurassic.<ref>{{Cite journal |last1=Slodownik |first1=Miriam |last2=Hill |first2=Robert S. |last3=McLoughlin |first3=Stephen |date=October 2023 |title=Komlopteris: A persistent lineage of post-Triassic corystosperms in Gondwana |journal=Review of Palaeobotany and Palynology |language=en |volume=317 |pages=104950 |doi=10.1016/j.revpalbo.2023.104950|bibcode=2023RPaPa.31704950S |s2cid=260015702 |doi-access=free }}</ref> [[Czekanowskiales]], also known as Leptostrobales, are a group of seed plants uncertain affinities with persistent heavily dissected leaves borne on deciduous short shoots, subtended by scale-like leaves, known from the Late Triassic (possibly Late Permian<ref name="Kustatscher-2019">{{Cite journal|last1=Kustatscher|first1=Evelyn|last2=Visscher|first2=Henk|last3=van Konijnenburg-van Cittert|first3=Johanna H. A.|date=2019-09-01|title=Did the Czekanowskiales already exist in the late Permian?|journal=PalZ|language=en|volume=93|issue=3|pages=465–477|doi=10.1007/s12542-019-00468-9|s2cid=199473893|issn=1867-6812|doi-access=free|bibcode=2019PalZ...93..465K }}</ref>) to Cretaceous.<ref name="Taylor-2009">{{Citation|last=Taylor|first=T|title=Gymnosperms with obscure affinities|date=2009|url=https://linkinghub.elsevier.com/retrieve/pii/B978012373972800019X|work=Biology and Evolution of Fossil Plants|pages=757–785|publisher=Elsevier|language=en|doi=10.1016/b978-0-12-373972-8.00019-x|isbn=978-0-12-373972-8|access-date=2020-12-13}}</ref> They are thought to have had a tree- or shrub-like habit and formed a conspicuous component of Northern Hemisphere Mesozoic temperate and warm-temperate floras.<ref name="Kustatscher-2019" /> The genus ''[[Phoenicopsis]]'' was widespread in Early-Middle Jurassic floras of Eastern Asia and Siberia.<ref>{{Cite journal|last1=Sun|first1=Chunlin|last2=Li|first2=Yunfeng|last3=Dilcher|first3=David L.|last4=Wang|first4=Hongshan|last5=Li|first5=Tao|last6=Na|first6=Yuling|last7=Wang|first7=Anping|date=November 2015|title=An introductory report on the biodiversity of Middle Jurassic Phoenicopsis (Czekanowskiales) from the Ordos Basin, China|url=https://linkinghub.elsevier.com/retrieve/pii/S2095927316302869|journal=Science Bulletin|language=en|volume=60|issue=21|pages=1858–1865|doi=10.1007/s11434-015-0904-y|bibcode=2015SciBu..60.1858S|s2cid=140617907}}</ref> The [[Pentoxylales]], a small but clearly distinct group of [[liana]]-like seed plants of obscure affinities, first appeared during the Jurassic. Their distribution appears to have been confined to Eastern Gondwana.<ref>Pattemore, G.A., Rigby, J.F. and Playford, G., 2015. Triassic-Jurassic pteridosperms of Australasia: speciation, diversity and decline. Boletín Geológico y Minero, 126 (4): 689–722</ref> ==== Ferns and allies ==== Living families of ferns widespread during the Jurassic include [[Dipteridaceae]], [[Matoniaceae]], [[Gleicheniaceae]], [[Osmundaceae]] and [[Marattiaceae]].<ref>{{Cite journal|last=Skog|first=Judith E.|date=April 2001|title=Biogeography of Mesozoic leptosporangiate ferns related to extant ferns|url=http://dx.doi.org/10.1007/bf02812701|journal=Brittonia|volume=53|issue=2|pages=236–269|doi=10.1007/bf02812701|bibcode=2001Britt..53..236S |s2cid=42781830|issn=0007-196X}}</ref><ref>{{Cite journal|last1=Tian|first1=Ning|last2=Wang|first2=Yong-Dong|last3=Zhang|first3=Wu|last4=Zheng|first4=Shao-Lin|last5=Zhu|first5=Zhi-Peng|last6=Liu|first6=Zhong-Jian|date=2018-03-01|title=Permineralized osmundaceous and gleicheniaceous ferns from the Jurassic of Inner Mongolia, NE China|url=https://doi.org/10.1007/s12549-017-0313-0|journal=Palaeobiodiversity and Palaeoenvironments|language=en|volume=98|issue=1|pages=165–176|doi=10.1007/s12549-017-0313-0|bibcode=2018PdPe...98..165T |issn=1867-1608|s2cid=134149095}}</ref> [[Polypodiales]], which make up 80% of living fern diversity, have no record from the Jurassic and are thought to have diversified in the Cretaceous,<ref>{{Cite journal|last1=Regalado|first1=Ledis|last2=Schmidt|first2=Alexander R.|last3=Müller|first3=Patrick|last4=Niedermeier|first4=Lisa|last5=Krings|first5=Michael|last6=Schneider|first6=Harald|date=July 2019|title=Heinrichsia cheilanthoides gen. et sp. nov., a fossil fern in the family Pteridaceae (Polypodiales) from the Cretaceous amber forests of Myanmar|journal=Journal of Systematics and Evolution|language=en|volume=57|issue=4|pages=329–338|doi=10.1111/jse.12514|issn=1674-4918|doi-access=free|bibcode=2019JSyEv..57..329R |s2cid=182754946}}</ref> though the widespread Jurassic [[Herbaceous plant|herbaceous]] fern genus ''[[Coniopteris]],'' historically interpreted as a close relative of [[tree fern]]s of the family [[Dicksoniaceae]], has recently been reinterpreted as an early relative of the group.<ref name="Coniopteris">{{Cite journal|last1=Li|first1=Chunxiang|last2=Miao|first2=Xinyuan|last3=Zhang|first3=Li-Bing|last4=Ma|first4=Junye|last5=Hao|first5=Jiasheng|date=January 2020|title=Re-evaluation of the systematic position of the Jurassic–Early Cretaceous fern genus Coniopteris|url=https://linkinghub.elsevier.com/retrieve/pii/S0195667118303033|journal=Cretaceous Research|language=en|volume=105|pages=104136|doi=10.1016/j.cretres.2019.04.007|bibcode=2020CrRes.10504136L |s2cid=146355798}}</ref> The [[Cyatheales]], the group containing most modern tree ferns, appeared during the Late Jurassic, represented by members of the genus ''[[Cyathocaulis]]'', which are suggested to be early members of [[Cyatheaceae]] on the basis of cladistic analysis.<ref>{{Cite journal |last1=Korall |first1=Petra |last2=Pryer |first2=Kathleen M. |date=February 2014 |editor-last=Parmakelis |editor-first=Aristeidis |title=Global biogeography of scaly tree ferns (Cyatheaceae): evidence for Gondwanan vicariance and limited transoceanic dispersal |journal=Journal of Biogeography |language=en |volume=41 |issue=2 |pages=402–413 |doi=10.1111/jbi.12222 |issn=0305-0270 |pmc=4238398 |pmid=25435648|bibcode=2014JBiog..41..402K }}</ref> Only a handful of possible records exist of the [[Hymenophyllaceae]] from the Jurassic, including ''Hymenophyllites macrosporangiatus'' from the Russian Jurassic.<ref>{{Cite journal|last1=Axsmith|first1=Brian J.|last2=Krings|first2=Michael|last3=Taylor|first3=Thomas N.|date=September 2001|title=A filmy fern from the Upper Triassic of North Carolina (USA)|journal=American Journal of Botany|volume=88|issue=9|pages=1558–1567|doi=10.2307/3558399|issn=0002-9122|jstor=3558399|pmid=21669688|doi-access=free|bibcode=2001AmJB...88.1558A }}</ref> The oldest remains of modern [[Equisetaceae|horsetails]] of the genus ''[[Equisetum]]'' first appear in the Early Jurassic, represented by ''[[Equisetum dimorphum]]'' from the Early Jurassic of Patagonia<ref>{{Cite journal|last1=Elgorriaga|first1=Andrés|last2=Escapa|first2=Ignacio H.|last3=Bomfleur|first3=Benjamin|last4=Cúneo|first4=Rubén|last5=Ottone|first5=Eduardo G.|date=February 2015|title=Reconstruction and Phylogenetic Significance of a New Equisetum Linnaeus Species from the Lower Jurassic of Cerro Bayo (Chubut Province, Argentina)|url=http://www.bioone.org/doi/10.5710/AMGH.15.09.2014.2758|journal=Ameghiniana|language=en|volume=52|issue=1|pages=135–152|doi=10.5710/AMGH.15.09.2014.2758|bibcode=2015Amegh..52..135E |issn=0002-7014|s2cid=6134534|hdl=11336/66623|hdl-access=free}}</ref> and ''[[Equisetum laterale]]'' from the Early to Middle Jurassic of Australia.<ref>Gould, R. E. 1968. Morphology of ''Equisetum laterale'' Phillips, 1829, and ''E. bryanii'' sp. nov. from the Mesozoic of south‐eastern Queensland. Australian Journal of Botany 16: 153–176.</ref><ref name="Elgorriaga-2018">{{Cite journal|last1=Elgorriaga|first1=Andrés|last2=Escapa|first2=Ignacio H.|last3=Rothwell|first3=Gar W.|last4=Tomescu|first4=Alexandru M. F.|last5=Rubén Cúneo|first5=N.|date=August 2018|title=Origin of Equisetum : Evolution of horsetails (Equisetales) within the major euphyllophyte clade Sphenopsida|journal=American Journal of Botany|language=en|volume=105|issue=8|pages=1286–1303|doi=10.1002/ajb2.1125|pmid=30025163|doi-access=free}}</ref> [[Petrifaction|Silicified]] remains of ''[[Equisetum thermale]]'' from the Late Jurassic of Argentina exhibit all the morphological characters of modern members of the genus.<ref>{{Cite journal|last1=Channing|first1=Alan|last2=Zamuner|first2=Alba|last3=Edwards|first3=Dianne|last4=Guido|first4=Diego|date=2011|title=Equisetum thermale sp. nov. (Equisetales) from the Jurassic San Agustín hot spring deposit, Patagonia: Anatomy, paleoecology, and inferred paleoecophysiology|journal=American Journal of Botany|language=en|volume=98|issue=4|pages=680–697|doi=10.3732/ajb.1000211|issn=1537-2197|pmid=21613167|doi-access=free|bibcode=2011AmJB...98..680C |hdl=11336/95234|hdl-access=free}}</ref> The estimated split between ''[[Equisetum bogotense]]'' and all other living ''Equisetum'' is estimated to have occurred no later than the Early Jurassic.<ref name="Elgorriaga-2018" /> ==== Lower plants ==== [[Isoetes|Quillworts]] virtually identical to modern species are known from the Jurassic onwards. ''[[Isoetites rolandii]]'' from the Middle Jurassic of Oregon is the earliest known species to represent all major morphological features of modern ''Isoetes''. More primitive forms such as ''[[Nathorstiana]],'' which retain an elongated stem, persisted into the Early Cretaceous.<ref>{{Cite journal|last1=Wood|first1=Daniel|last2=Besnard|first2=Guillaume|last3=Beerling|first3=David J.|last4=Osborne|first4=Colin P.|last5=Christin|first5=Pascal-Antoine|date=2020-06-18|title=Phylogenomics indicates the "living fossil" Isoetes diversified in the Cenozoic|journal=PLOS ONE|language=en|volume=15|issue=6|pages=e0227525|doi=10.1371/journal.pone.0227525|issn=1932-6203|pmc=7302493|pmid=32555586|bibcode=2020PLoSO..1527525W|doi-access=free}}</ref> The moss ''[[Kulindobryum]]'' from the Middle Jurassic of Russia, which was found associated with dinosaur bones, is thought to be related to the [[Splachnaceae]], which grow on animal caracasses.<ref name="Mamontov-2019" /> ''[[Bryokhutuliinia]]'' from the same region is thought to be related to [[Dicranales]].<ref name="Mamontov-2019">{{Cite journal|last1=Mamontov|first1=Yuriy S.|last2=Ignatov|first2=Michael S.|date=July 2019|title=How to rely on the unreliable: Examples from Mesozoic bryophytes of Transbaikalia|journal=Journal of Systematics and Evolution|language=en|volume=57|issue=4|pages=339–360|doi=10.1111/jse.12483|s2cid=92268163|issn=1674-4918|doi-access=free|bibcode=2019JSyEv..57..339M }}</ref> ''[[Heinrichsiella]]'' from the Jurassic of Patagonia is thought to belong to either [[Polytrichaceae]] or [[Timmiellaceae]].<ref>{{Cite journal|last1=Bippus|first1=Alexander C.|last2=Savoretti|first2=Adolfina|last3=Escapa|first3=Ignacio H.|last4=Garcia-Massini|first4=Juan|last5=Guido|first5=Diego|date=October 2019|title=Heinrichsiella patagonica gen. et sp. nov.: A Permineralized Acrocarpous Moss from the Jurassic of Patagonia|url=http://dx.doi.org/10.1086/704832|journal=International Journal of Plant Sciences|volume=180|issue=8|pages=882–891|doi=10.1086/704832|bibcode=2019IJPlS.180..882B |s2cid=202859471|issn=1058-5893}}</ref> The liverwort ''[[Pellites|Pellites hamiensis]]'' from the Middle Jurassic [[Xishanyao Formation]] of China is the oldest record of the family [[Pelliaceae]].<ref>{{Cite journal|last1=Li|first1=Ruiyun|last2=Li|first2=Xiaoqiang|last3=Deng|first3=Shenghui|last4=Sun|first4=Bainian|date=August 2020|title=Morphology and microstructure of Pellites hamiensis nov. sp., a Middle Jurassic liverwort from northwestern China and its evolutionary significance|url=https://linkinghub.elsevier.com/retrieve/pii/S001669952030070X|journal=Geobios|volume=62|language=en|pages=23–29|doi=10.1016/j.geobios.2020.07.003|bibcode=2020Geobi..62...23L |s2cid=225500594}}</ref> [[Pallaviciniites|''Pallaviciniites sandaolingensis'']] from the same deposit is thought to belong to the subclass Pallaviciniineae within the [[Pallaviciniales]].<ref>{{Cite journal|last1=Li|first1=Rui-Yun|last2=Wang|first2=Xue-lian|last3=Chen|first3=Jing-Wei|last4=Deng|first4=Sheng-Hui|last5=Wang|first5=Zi-Xi|last6=Dong|first6=Jun-Ling|last7=Sun|first7=Bai-Nian|date=June 2016|title=A new thalloid liverwort: Pallaviciniites sandaolingensis sp. nov. from the Middle Jurassic of Turpan–Hami Basin, NW China|url=http://link.springer.com/10.1007/s12542-016-0299-3|journal=PalZ|language=en|volume=90|issue=2|pages=389–397|doi=10.1007/s12542-016-0299-3|bibcode=2016PalZ...90..389L |s2cid=131295547|issn=0031-0220}}</ref> ''[[Ricciopsis|Ricciopsis sandaolingensis]]'', also from the same deposit, is the only Jurassic record of [[Ricciaceae]].<ref>{{Cite journal|last1=Li|first1=Ruiyun|last2=Li|first2=Xiaoqiang|last3=Wang|first3=Hongshan|last4=Sun|first4=Bainian|date=2019|title=Ricciopsis sandaolingensis sp. nov., a new fossil bryophyte from the Middle Jurassic Xishanyao Formation in the Turpan-Hami Basin, Xinjiang, Northwest China|journal=Palaeontologia Electronica|volume=22|issue=2|page=917 |doi=10.26879/917|issn=1094-8074|doi-access=free|bibcode=2019PalEl..22..917L }}</ref>
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