Belongs within: Apo-Spermatophyta.
Contains: Cycadophyta, Coniferophyta, Pan-Gnetophyta, Pan-Angiospermae, Arberiaceae.
The name Spermatophyta was defined by Cantino et al. (2007) to represent the crown group of seed-bearing plants. Relationships within the spermatophytes remain uncertain: most morphological studies have supported a nested position for angiosperms among the remaining living taxa, but molecular analyses have supported a sister relationship between the angiosperms and living gymnosperms.
The clade Pan-Cycadophyta includes the cycads and all fossil taxa more closely related to them than other living seed plants. Known members of this clade are characterised by girdling leaf trances (Cantino et al. 2007), where leaf traces or veins leading to the leaf arise opposite the leaf attachment and circle the trunk within the storage tissue (Cycad Pages). The Nilssoniaceae were Mesozoic pan-cycadophytes that bore pinnate or simple leaves smaller than those of modern cycads. Semionogyna is a Lower Cretaceous genus named for female spikes similar to those of modern Cycas, bearing ovules subtended by bracts with distal bracts empty.
Fossil spermatophytes outside the major living lineages (cycads, conifers, gnetaleans and angiosperms) are mostly of the pteridosperm (‘seed fern’) grade, but some such as members of the Permian genus Glossopteris were large trees with entire leaves. The Pentoxylaceae are a group of plants (probably shrubs and small trees) known from the Jurassic, characterised by polystelic stems with a separate arrangement of vascular tissues for each stele. The Emplectopteridaceae were Permian pteridosperms with pinnate or entire leaves bearing flexuous or loosely anastomosed venation, and bilateral ovules attached to the underside of the leaves.
Possible synapomorphies (Cantino et al. 2007): Endarch primary xylem in the stem, meiospores and microgametophytes with distal aperture, linear tetrad of megaspores, platyspermic ovules.
Spermatophyta [Acrogymnospermae, Phanerogamia, Pinophyta, Spermatophytatinae, Spermatophytina, Spermatopsida, Taxinae]
|--Pan-Cycadophyta [Cycadopsida]CD07
| | i. s.: AntarcticycasCD07
| | MichelilloaCD07
| |--NilssoniaT13 [NilssoniaceaeZ02, Nilssoniales]
| | |--N. gracilisN57
| | |--N. medianaT13
| | `--N. orientalisT13
| `--+--SemionogynaZ02
| `--CycadophytaFC07
`--+--ConiferophytaFC07
`--+--+--Pan-GnetophytaFC07
| `--Pan-AngiospermaeFC07
`--+--ArberiaceaeFC07
`--Pentoxylaceae [Pentoxylales]RL09
| i. s.: ‘Taeniopteris’ spatulata McClelland 1850C93
|--Pentoxylon sahnii Srivastava 1944HS17, C93
|--Nympaniophyllum hobsoniRL09
|--Nipanioxylon guptai Srivastava 1944C93
|--Nipaniophyllum raoi Sahni 1948C93
|--CarnoconitesRL09
| |--C. compactum Srivastava 1944C93
| `--C. rajmahalensis (Wieland) Bose et al. 1984C93
`--SahniaRL09
|--S. laxiphoraPL12
`--S. nipaniensis Vishnu-Mittre 1953C93
Spermatophyta incertae sedis:
Emplectopteridaceae [Gigantonomiales, Gigantopteridaceae]Z02
|--Gigantotheca paradoxa Li & Yao 1983C93
|--Emplectopteris Halle 1927C93
| `--E. triangularis Halle 1927C93
|--Gigantonomia Li & Yao 1983C93
| `--G. fukiensis (Yabe & Oishi) Li & Yao 1983C93
`--Gigantopteris Schenk 1883C93
|--G. americanaD56
|--G. dentataG31
|--G. elongataG31
|--G. nicotinaefoliaG31
|--G. rarinervisG31
`--G. taiyuanensis (Asama) Meyen 1987C93
SanmigueliaPL12
Balmeiopsis limbatus (Balme) Archangelsky 1979I02
Elaterocolpites castelainii Jardiné & Magloire 1965I02
Elateroplicites africaensis Herngreen 1973I02
ElaterosporitesI02
|--E. klaszii (Jardiné & Magloire) Jardiné 1967I02
`--E. verrucatusI02
GaleacorneaI02
|--G. causea Stover 1963I02
`--G. clavis Stover 1963I02
Steevesipollenites sinuosus Azema & Boltenhagen 1974I02
Herodontia mikanioidesJ87
ScolosanthusJ87
|--S. densiflorusJ87
`--S. selleanusJ87
Hottea torbecianaJ87
CycadopitesPL12
|--C. carpentieri (Delcourt & Sprumont) Singh 1964I02
|--C. durhamensisPL12
|--C. follicularisPL12
`--C. fragilis Singh 1964I02
Scalesia pedunculataRJ11
Synangispadixis tidwelliiPL12
Cacabus miersiiRJ11
Leucosphaera Gilg 1897KC01
Monoceras Gothan 1909KC01
Preussiella Gilg 1897KC01
Rotularia Sternb. 1825KC01
Pometia pinnataH83
MeumH64
|--M. athamanticumHH03
`--M. foeniculumH64
Pyrethrum partheniumH64
SalisburiaR10
|--S. billardieriR10
|--S. ginkgoR10
`--S. palmata [=Jeanpaulia palmata Ratte 1886]F71
*Type species of generic name indicated
References
[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.
[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.
[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.
[H83] Hammen, L. van der. 1983. Contribution to the knowledge of the soil-fauna of New Guinea. Zoologische Verhandelingen 206: 1–36.
[HH03] Hernández, J. R., & J. F. Hennen. 2003. Rust fungi causing galls, witches’ brooms, and other abnormal plant growths in northwestern Argentina. Mycologia 95 (4): 728–755.
[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.
[H64] Hogg, J. 1864. List of rarer phanerogamous plants discovered in the south-east of Durham since 1829. Report of the British Association for the Advancement of Science 33 (Notices and Abstracts): 96.
[I02] Ibrahim, M. I. A. 2002. Late Albian-Middle Cenomanian palynofacies and palynostratigraphy, Abu Gharadig-5 well, Western Desert, Egypt. Cretaceous Research 23: 775–788.
[J87] Judd, W. S. 1987. Floristic study of Morne La Visite and Pic Macaya National Parks, Haiti. Bulletin of the Florida State Museum—Biological Sciences 32 (1): 1–136.
[KC01] Kirk, P. M., P. F. Cannon, J. C. David & J. A. Stalpers. 2001. Ainsworth & Bisby’s Dictionary of the Fungi 9th ed. CAB International: Wallingford (UK).
[N57] Novojilov, N. 1957. Un nouvel ordre d’arthropodes particuliers: Kazacharthra, du Lias des monts Ketmen (Kazakhstan SE, U.R.S.S.). Bulletin de la Société Géologique de France, ser. 6, 7: 171–185, pls 14–15.
[PL12] Peñalver, E., C. C. Labandeira, E. Barrón, X. Delclòs, P. Nel, A. Nel, P. Tafforeau & C. Soriano. 2012. Thrips pollination of Mesozoic gymnosperms. Proceedings of the National Academy of Sciences of the USA 109 (22): 8623–8628.
[RL09] Ren, D., C. C. Labandeira, J. A. Santiago-Blay, A. Rasnitsyn, C. Shih, A. Bashkuev, M. A. V. Logan, C. L. Hotton & D. Dilcher. 2009. A probable pollination mode before angiosperms: Eurasian, long-proboscid scorpionflies. Science 326: 840–847.
[R10] Richard, A. 1810. Note sur les plantes dites conifères. Annales du Muséum d’Histoire Naturelle 16: 296–299.
[RJ11] Rising, J. D., A. Jaramillo, J. L. Copete, P. G. Ryan & S. C. Madge. 2011. Family Emberizidae (buntings and New World sparrows). In: Hoyo, J. del, A. Elliott & D. A. Christie (eds) Handbook of the Birds of the World vol. 16. Tanagers to New World Blackbirds pp. 428–683. Lynx Edicions: Barcelona.
[T13] Thomas, H. H. 1913. The fossil flora of the Cleveland district of Yorkshire: I—The flora of the Marske quarry. With notes on the stratigraphy, by the Rev. George Johne Lane, F.G.S. Annals and Magazine of Natural History, series 8, 12: 150.
[Z02] Zherikhin, V. V. 2002. Ecological history of the terrestrial insects. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 331–388. Kluwer Academic Publishers: Dordrecht.