Maidenhair tree Ginkgo biloba, copyright Liné1.

Belongs within: Spermatophyta.
Contains: Ginkgoales, Umkomasiales, Peltaspermales, Cordaitales, Dicranophyllales, Miroviaceae, Doliostrobaceae, Voltziales, Pinaceae, Cheirolepidiaceae, Cupressophyta.

The Coniferophyta have been recognised as a clade of gymnosperms including conifers, the ginkgo Ginkgo biloba, and the Palaeozoic Cordaitales. Members of this clade have simple, dichotomously veined leaves, simple strobili in the axils of the leaves, and platyspermic seeds (Doyle 1998). Though represented by only a single species in the modern flora, the Ginkgoopsida have an extensive fossil record extending back to the Permian (Herrera et al. 2017).

The name Coniferopsida is used here for a clade uniting the living conifers with the Palaeozoic Voltziales (probably a paraphyletic grouping). Unique morphological synapomorphies have not yet been identified for this clade (Rothwell & Mapes 2001). Crown group Coniferae are characterised relative to other living plants by their needle-like leaves, tiered proembryos and loss or extreme reduction of one copy of the inverted repeat in the chloroplast genome (Cantino et al. 2007). Recent phylogenies have mostly placed the Pinaceae as the sister clade to the remaining living conifers.

    |  i. s.: Carpentieria frondosaRM01
    |         GinkgophyllumRM01
    |         Polyspermophyllum sergiiRM01
    |--Ginkgoopsida [Ginkgophyta]GT12
    |    |  i. s.: TrichopityaceaeC93
    |    |           |--Trichopitys heteromorpha Florin 1949RM01, C93
    |    |           |--Dichophyllum moorei Ellias ex Andrews 1941C93
    |    |           |--Biarmopteris pulchra Zalessky 1939C93
    |    |           `--Mauerites gracilis Zalessky 1939C93
    |    |         Karkenia incurvaHS17
    |    |         BaieraGT12
    |    |           |--B. digitataR87
    |    |           |--B. pulchellaG31
    |    |           `--B. tenuifolia Johnston 1886R87
    |    |         Yimaia recurvaHS17
    |    |         Toretzia angustifoliaHS17
    |    |         Nagrenia samylinae [incl. Sphenobaiera angrenica]HS17
    |    |         KannaskoppiaHS17
    |    |         Doylea tetrahedraspermaHS17
    |    |--GinkgoalesFC07
    |    |--UmkomasialesHS17
    |    |--PeltaspermalesHS17
    |    |--PetriellealesG10
    |    `--VladimarialesHS17
    |         |--Entylissa umaltensisHS17
    |         |--Vladimaria Gordenko 2010 [Vladimariaceae]G10
    |         |    `--*V. octopartita Gordenko 2010G10
    |         `--Umaltolepis [Umaltolepidaceae]HS17
    |              |--*U. vachrameevii [incl. Pseudotorellia angustifolia]HS17
    |              |--U. coleoptera [incl. Pseudotorellia roobana]HS17
    |              |--U. hebeiensisHS17
    |              |--U. mongoliensis Herrera, Shi et al. 2017 [incl. Pseudotorellia resinosa]HS17
    |              `--U. rarinervisHS17
         |  i. s.: VojnovskyalesRM01
         |           |  i. s.: Sergeia neuburgiiRM01
         |           `--VojnovskyaceaeC93
         |                |--Kuznetskia planiuscula Meyen 1982C93
         |                `--VojnovskyaRM01
         |                     |--V. paradoxaRM01
         |                     `--V. usjatensis Gorelova in Gorelova et al. 1973C93
            `--Coniferopsida [Coniferales, Conopsida, Pinales, Pinopsida]RMH-C05
                 |  i. s.: TylodendronH-CRM01
                 |         Palaeotaxites praecursorH-CRM01
                 |         ParanocladusH-CRM01
                 |         Szecladia multinerviaH-CRM01
                 |         DadoxylonH-CRM01
                 |           |--D. farleyensis Walkom 1928F71
                 |           `--D. owangondianumH-CRM01
                 |         Araucarioxylon arizonicum Knowlton 1889H-CRM01, CA04
                 |         Walchiopremnon valdajolenseH-CRM01
                 |         CulmitzschiaH-CRM01
                 |         Buriadia [Buriadiaceae]H-CRM01
                 |           `--B. heterophylla (Feistmantel) Seward & Sahni 1920C93
                 |         CurioniaH-CRM01
                 |         FeysiaH-CRM01
                 |         Gomphostrobus bifidusH-CRM01, RM01
                 |         WalkomiaH-CRM01
                 |         CephalotaxopsisZBH03
                 |         Samaropsis pincombei Walkom 1928Z02, F71
                 |         Bilsdalea duraZ02
                 |         HeidiphyllumZ02
                 |         Podozamites Braun 1843 [Podozamitaceae]BO02
                 |           |--P. gracilis [=Araucarites gracilis]F71
                 |           `--P. guttiformisBO02
                 |         PityophyllumGT12
                 |         RhipidiocladusGT12
                 |         SchizolepisGT12
                 |         Cyparissidium gracileGT02
                 |         SequoitesGT02
                 |         MiroviaceaeG02
                 |         SciadopityoxylonPM02
                 |         XenoxylonPM02
                 |         Podocarpoxylon Gothan 1905JD05
                 |           |--P. angustiporosumJD05
                 |           |--P. helmstedtianum Gottwald 1966K02
                 |           `--P. knowltoniiJD05
                 |         ProtopodocarpoxylonS03
                 |         DoliostrobaceaeK02
                 |         DammaraK02
                 |           |--D. borealisK02
                 |           |--D. intermedia Ettingshausen 1886F71
                 |           |--D. macrospermaK02
                 |           `--D. podozamioides Ettingshausen 1886F71
                 |         Chamaecyparites hardtiiK02
                 |         Embothrites cuneatus Ettinghausen 1868K02
                 |         Pityanthus jefferyiRM01
                 |         Eutactoides Pole 2000P00
                 |           `--*E. wintonensis Pole 2000P00
                 |         Circoporoxylon Kraeusel 1949JD05
                 |         PalissyaceaeC93
                 |           |--PalissyaC93
                 |           `--StachyotaxusC93
                 |                |--S. lipoldii (Stur) Kräusel 1952C93
                 |                `--S. sahnii Kräusel 1952C93
                 |         Pararaucaria [Pararaucariaceae]C93
                 |           `--P. patagonica Wieland 1929C93
                 |         Araucariopitys americana Jeffrey 1907CA04
                 |         Cupressospermum saxonicumP92
                 `--Coniferae [Pinoideae]CD07
                      |  i. s.: LepidothamnusD03
                      |           |--L. fonckiD03
                      |           `--L. laxifoliusW91

*Type species of generic name indicated


[BO02] Barale, G., & M. Ouaja. 2002. La biodiversité végétale des gisements d’âge Jurassique supérieur-Crétacé inférieur de Merbah El Asfer (Sud-Tunisien). Cretaceous Research 23: 707–737.

[CD07] Cantino, P. D., J. A. Doyle, S. W. Graham, W. S. Judd, R. G. Olmstead, D. E. Soltis, P. S. Soltis & M. J. Donoghue. 2007. Towards a phylogenetic nomenclature of Tracheophyta. Taxon 56 (3): E1–E44.

[C93] Cleal, C. J. 1993. Gymnospermophyta. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 795–808. Chapman & Hall: London.

[CA04] Creber, G. T., & S. R. Ash. 2004. The Late Triassic Schilderia adamanica and Woodworthia arizonica trees of the Petrified Forest National Park, Arizona, USA. Palaeontology 47: 21–38.

[D03] Dusén, P. 1903. The vegetation of western Patagonia. In: Scott, W. B. (ed.) Reports of the Princeton University Expeditions to Patagonia, 18961899 vol. 8. Botany pp. 1–34. The University: Princeton (New Jersey).

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[FC07] Friis, E. M., P. R. Crane, K. R. Pedersen, S. Bengtson, P. C. J. Donoghue, G. W. Grimm & M. Stampanoni. 2007. Phase-contrast X-ray microtomography links Cretaceous seed with Gnetales and Bennettitales. Nature 450: 549–552.

[GT12] Giribet, G., A. L. Tourinho, C. Shih & D. Ren. 2012. An exquisitely preserved harvestman (Arthropoda, Arachnida, Opiliones) from the Middle Jurassic of China. Organisms, Diversity & Evolution 12: 51-66.

[G02] Gomez, B. 2002. A new species of Mirovia (Coniferales, Miroviaceae) from the Lower Cretaceous of the Iberian Ranges (Spain). Cretaceous Research 23: 761–773.

[GT02] Gomez, B., F. Thévenard, M. Fantin & L. Guisberti. 2002. Late Cretaceous plants from the Bonarelli Level of the Venetian Alps, northeastern Italy. Cretaceous Research 23: 671–685.

[G10] Gordenko, N. V. 2010. Vladimariales ordo nov. (Gymnospermae) from the Middle Jurassic deposits of the Mikhailovskii rudnik locality (Kursk Region, European Russia). Paleontological Journal 44 (10): 1281–1307.

[G31] Grabau, A. W. 1931. The Permian of Mongolia: A report on the Permian fauna of the Jisu Honguer limestone of Mongolia and its relations to the Permian of other parts of the world. American Museum of Natural History: New York.

[H-CRM01] Hernandez-Castillo, G. R., G. W. Rothwell & G. Mapes. 2001. Thucydiaceae fam. nov., with a review and reevaluation of Paleozoic walchian conifers. International Journal of Plant Science 162 (5): 1155–1185.

[HS17] Herrera, F., G. Shi, N. Ichinnorov, M. Takahashi, E. V. Bugdaeva, P. S. Herendeen & P. R. Crane. 2017. The presumed ginkgophyte Umaltolepis has seed-bearing structures resembling those of Peltaspermales and Umkomasiales. Proceedings of the National Academy of Sciences of the USA: E2385–E2391.

[JD05] Junge, F. W., M. Dolezych, H. Walther, T. Böttger, A. Kühl, L. Kunzmann, P. Morgenstern, T. Steinberg & R. Stange. 2005. Ein Fenster in Landschaft und Vegetation vor 37 Millionen Jahren: Lithologische, sedimentgeochemische und paläobotanische Befunde aus einem Paläoflusssystem des Weißelsterbeckens. Mauritiana 19 (2): 185–273.

[K02] Kvaček, Z. 2002. Novelties on Doliostrobus (Doliostrobaceae), an extinct conifer genus of the European Palaeogene. Časopis Národního Muzea, Řada Přírodovědná 171 (1–4): 47–62.

[PM02] Philippe, M., H. Méon, G. Lambert, B. Erdei, F. Thevenard & B. Gomez. 2002. A palm-tree and Sciadopitys swamp-forest from the Neogene of Bresse (eastern France). Comptes Rendus Palevol 1: 221–225.

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

[P00] Pole, M. S. 2000. Mid-Cretaceous conifers from the Eromanga Basin, Australia. Australian Systematic Botany 13 (2): 153–197.

[R87] Ratte, F. 1887. Notes on Australian fossils. Proceedings of the Linnean Society of New South Wales, series 2, 1 (4): 1065–1084, pls 15–17.

[RM01] Rothwell, G. W., & G. Mapes. 2001. Barthelia furcata gen. et sp. nov., with a review of Paleozoic coniferophytes and a discussion of coniferophyte systematics. International Journal of Plant Sciences 162 (3): 637–667.

[RMH-C05] Rothwell, G. W., G. Mapes & G. R. Hernandez-Castillo. 2005. Hanskerpia gen. nov. and phylogenetic relationships among the most ancient conifers (Voltziales). Taxon 54 (3): 733–750.

[S03] Skog, J. E. 2003. Report of the Committee for Fossil Plants: 4. Taxon 52: 341.

[W91] Williams, P. A. 1991. Subalpine and alpine vegetation of granite ranges in western Nelson, New Zealand. New Zealand Journal of Botany 29: 317–330.

[Z02] Zherikhin, V. V. 2002. Insect trace fossils. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 303–324. Kluwer Academic Publishers: Dordrecht.

[ZBH03] Zhou, Z., P. M. Barrett & J. Hilton. 2003. An exceptionally preserved Lower Cretaceous ecosystem. Nature 421: 807–814.

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