Reconstruction of male reproductive structure of Sahnia nipaensis, from here.

Belongs within: Apo-Spermatophyta.
Contains: Cycadophyta, Coniferophyta, Arberiaceae, Pan-Gnetophyta, Pan-Angiospermae.

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).

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 (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, Phanerogamia, Pinophyta, Spermatophytatinae, Spermatophytina, Spermatopsida]
    |--Pan-Cycadophyta [Cycadopsida] CD07
    |    |  i. s.: Anomozamites RY03
    |    |         Antarcticycas CD07
    |    |         Michelilloa CD07
    |    |--Nilssoniaceae [Nilssoniales] Z02
    |    `--+--Semionogyna Z02
    |       `--Cycadophyta FC07
    `--+--Coniferophyta FC07
       `--+--Arberiaceae FC07
          |--+--Pan-Gnetophyta FC07
          |  `--Pan-Angiospermae FC07
          `--Pentoxylaceae [Pentoxylales] RL09
               |  i. s.: ‘Taeniopteris’ spatulata McClelland 1850 C93
               |--Pentoxylon sahnii Srivastava 1944 FC07, C93
               |--Nympaniophyllum hobsoni RL09
               |--Nipanioxylon guptai Srivastava 1944 C93
               |--Nipaniophyllum raoi Sahni 1948 C93
               |--Carnoconites RL09
               |    |--C. compactum Srivastava 1944 C93
               |    `--C. rajmahalensis (Wieland) Bose et al. 1984 C93
               `--Sahnia RL09
                    |--S. laxiphora PL12
                    `--S. nipaniensis Vishnu-Mittre 1953 C93

Spermatophyta incertae sedis:
  Emplectopteridaceae [Gigantonomiales, Gigantopteridaceae] Z02
    |--Gigantotheca paradoxa Li & Yao 1983 C93
    |--Emplectopteris Halle 1927 C93
    |    `--E. triangularis Halle 1927 C93
    |--Gigantonomia Li & Yao 1983 C93
    |    `--G. fukiensis (Yabe & Oishi) Li & Yao 1983 C93
    `--Gigantopteris Schenk 1883 C93
         |--G. americana D56
         `--G. taiyuanensis (Asama) Meyen 1987 C93
  Sanmiguelia PL12
  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
  Herodontia mikanioides J87
  Scolosanthus J87
    |--S. densiflorus J87
    `--S. selleanus J87
  Graffenrieda chrysandra J87
  Hottea torbeciana J87
  Reedia spathacea JK08
  Cycadopites PL12
    |--C. carpentieri (Delcourt & Sprumont) Singh 1964 I02
    |--C. durhamensis PL12
    |--C. follicularis PL12
    `--C. fragilis Singh 1964 I02
  Scalesia pedunculata HSS13
  Synangispadixis tidwellii PL12
  Cacabus miersii RJ11
  Leucosphaera Gilg 1897 KC01
  Monoceras Gothan 1909 KC01
  Preussiella Gilg 1897 KC01
  Rotularia Sternb. 1825 KC01

*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.

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

[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.

[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.

[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).

[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.

[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.

[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.

[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.

Last updated: 31 January 2020.

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