Fossil leaves of the Permian tree Glossopteris browniana, photographed at the British Natural History Museum by Martin Antoš.

Belongs within: Lignophyta.
Contains: Cycadophyta, Coniferophyta, Pan-Angiospermae, Erdtmanithecales, Bennettitales, Gnetales.

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. Fossil spermatophytes outside the major living lineages (cycads, conifers, gnetaleans and angiosperms) are mostly of the 'seed fern' grade, but some such as members of the Permian genus Glossopteris were large trees with entire leaves.

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 Peltaspermales are a group of seed ferns known from the Permian and Triassic, in which reproductive structures were borne in clusters radiating from a long central axis.

Possible synapomorphies (from Cantino et al. 2007): Endarch primary xylem in the stem, meiospores and microgametophytes with distal aperture, linear tetrad of megaspores, platyspermic ovules.

<==Spermatophyta [Acrogymnospermae, Pinophyta, Spermatophytatinae, Spermatophytina, Spermatopsida]
    |--Pan-Cycadophyta [Cycadopsida] CD07
    |    |  i. s.: Anomozamites RY03
    |    |         Antarcticycas CD07
    |    |         Michelilloa CD07
    |    |         Crossozamia NM11
    |    |--Trigonocarpus Brongniart 1828 Z02a [Trigonocarpales Z02b]
    |    |--Nilssoniaceae [Nilssoniales] Z02b
    |    |--Lagenostoma FC07 [Lagenostomales Z02b]
    |    |    `--L. lomaxi FC07
    |    `--+--Semionogyna Z02b
    |       `--Cycadophyta FC07
    `--+--Coniferophyta FC07
       |--Peltaspermales Z02b
       |    |--Permotheca Z02b
       |    |--Comia Z02b
       |    |--Tatarina Z02b
       |    |--Glossophyllum Z02b
       |    |--Pachypteris austropapillosa Z02b, M01
       |    |--Peltaspermum FC07
       |    `--Antevsia PL12
       |         |--A. extans PL12
       |         `--A. zeilleri PL12
       `--+--Pentoxylaceae [Pentoxylales] RL09
          |    |--Pentoxylon FC07
          |    |--Carnoconites compactus RL09
          |    |--Sahnia RL09
          |    |    |--S. laxiphora PL12
          |    |    `--S. nipaensis PL12
          |    `--Nympaniophyllum hobsoni RL09
          |--+--Pan-Angiospermae CD07
          |  `--Pan-Gnetophyta CD07
          |       |--Erdtmanithecales FC07
          |       |--Bennettitales FC07
          |       `--+--Gnetales FC07
          |          `--Piroconites D98
          `--Glossopteris [Arberiales, Glossopteridales] FC07
               |--G. ampla F71
               |--G. angustifolia F71
               |    |--G. a. var. angustifolia F71
               |    `--G. a. var. taeniopteroides F71
               |--G. browniana F71
               |    |--G. b. var. browniana F71
               |    `--G. b. var. praecursor Feistmantel 1878-1879 F71
               |--G. conspicua F71
               |--G. linearis F71
               |--G. mitchelli Walkom 1928 F71
               |--G. primaeva Feistmantel 1878-1879 F71
               `--G. spathulato-cordata F71

Spermatophyta incertae sedis:
  Gigantopteris [Gigantonomiales, Gigantopteridaceae] Z02b
  Sanmiguelia Z02b
  Iraniales Z02b
  Balmeiopsis limbatus (Balme) Archangelsky 1979 I02
  Elaterocolpites castelainii Jardiné & Magloire 1965 I02
  Elateroplicites africaensis Herngreen 1973 I02
  Elaterosporites I02
    |--E. klaszii (Jardiné & Magloire) Jardiné 1967 I02
    `--E. verrucatus I02
  Galeacornea I02
    |--G. causea Stover 1963 I02
    `--G. clavis Stover 1963 I02
  Steevesipollenites sinuosus Azema & Boltenhagen 1974 I02
  Callixene marginata D03
  Herodontia mikanioides J87
  Proustia stenophylla J87
  Scolosanthus J87
    |--S. densiflorus J87
    `--S. selleanus J87
  Graffenrieda chrysandra J87
  Hottea torbeciana J87
  Reedia spathacea JK08
  Spermolepis echinata PB83
  Endiandra palmerstoni H42
  Cardwellia sublimis H42
  Cycadopites RL09
    |--C. carpentieri (Delcourt & Sprumont) Singh 1964 I02
    |--C. durhamensis PL12
    |--C. follicularis PL12
    `--C. fragilis Singh 1964 I02
  Scalesia pedunculata HSS13
  Synangispadixis tidwellii PL12

*Type species of generic name indicated


[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.

[D98] Doyle, J. A. 1998. Phylogeny of vascular plants. Annual Review of Ecology and Systematics 29: 567-599.

[D03] Dusén, P. 1903. The vegetation of western Patagonia. In Reports of the Princeton University Expeditions to Patagonia, 1896-1899, vol. 8—Botany (W. B. Scott, ed.) 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.

[H42] Hill, G. F. 1942. Termites (Isoptera) from the Australian Region (including Australia, New Guinea and islands south of the Equator between 140°E. longitude and 170°W. longitude). Commonwealth of Australia Council for Scientific and Industrial Research: Melbourne.

[HSS13] Hirschfeld, E., A. Swash & R. Still. 2013. The World's Rarest Birds. Princeton University Press: Princeton (New Jersey).

[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.

[JK08] Johnstone, R. E., & T. Kirkby. 2008. Distribution, status, social organisation, movements and conservation of Baudin's cockatoo (Calyptorhynchus baudinii) in south-west Western Australia. Records of the Western Australian Museum 25 (1): 107-118.

[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.

[M01] McLoughlin, S. 2001. The breakup history of Gondwana and its impact on pre-Cenozoic floristic provincialism. Australian Journal of Botany 49: 271-300.

[NM11] Nagalingum, N. S., C. R. Marshall, T. B. Quental, H. S. Rai, D. P. Little & S. Mathews. 2011. Recent synchronous radiation of a living fossil. Science 334: 796-799.

[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.

[PB83] Price, M. V. & J. H. Brown. 1983. Patterns of morphology and resource use in North American desert rodent communities. Great Basin Naturalist Memoirs 7: 117-134.

[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.

[RY03] Ren, D., & J. Yin. 2003. New ‘osmylid-like’ fossil Neuroptera from the Middle Jurassic of Inner Mongolia, China. Journal of the New York Entomological Society 111 (1): 1-11.

[Z02a] Zherikhin, V. V. 2002a. Insect trace fossils. In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 303-324. Kluwer Academic Publishers: Dordrecht.

[Z02b] Zherikhin, V. V. 2002b. Ecological history of the terrestrial insects. In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 331-388. Kluwer Academic Publishers: Dordrecht.

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