Reconstruction of Cooksonia, copyright Matteo De Stefano.

Belongs within: Embryophyta.
Contains: Rhyniaceae, Tracheophyta.

The Apo-Tracheophyta are a broad clade of land plants, defined by Cantino et al. (2007) as the most inclusive clade possessing tracheids (differentially thickened water conducting cells) homologous with those of Pinus sylvestris.

<==Apo-Tracheophyta [Psilophyta, Pteridophyta, Rhyniophyta, Rhyniophytina, Rhyniopsida, Tracheidatae]
    |--Lyonophyton Remy & Remy 1980SP12 [incl. Aglaophyton Edwards 1986D98, SP12]
    |    |--‘Rhynia’ majorD98 (see below for synonymy)
    |    `--‘Retusotriletes’ warringtoniiW97
       |    |  i. s.: Rowleya trifurcataC02
       |    |         Fannningopteris Clifford 2002C02
       |    |           `--*F. wyattii Clifford 2002 [=F. queenslandica Clifford 2002 (n. n.)]C02
       |    `--StauropteridaceaeC93
       |         |--Gillespiea randolphensis Erwin & Rothwell 1989C93
       |         `--StauropterisC02
       |              |--S. berwickensisC02
       |              |--S. biseriata Cichan & Taylor 1982C93
       |              |--S. burntislandicaC02
       |              `--S. oldhamiaC02
             `--Cooksonia Lang 1937 (see below for synonymy)SP12
                  |--*C. pertoni Lang 1937SP12
                  |    |--C. p. ssp. pertoniSP12
                  |    |--C. p. ssp. apiculispora (see below for synonymy)SP12
                  |    `--C. p. ssp. synorispora [incl. Synorisporites verrucatus Richardson & Lister 1969]SP12
                  |--‘Ambitisporites’ avitus (Hoffmeister 1959) Steemans et al. 1996SW04
                  |--C. banksii Habgood et al. 2002SP12
                  |--C. bohemicaEE86
                  |--C. cambrensis Edwards 1979C93
                  |--C. crassiparietilisEE86
                  |--‘Ambitisporites’ dilutusW97
                  |--C. hemisphaerica Lang 1937C93
                  |--‘Synorisporites’ libycusSP12
                  |--‘Synorisporites’ maculosus Dufka 1995SP12
                  |--‘Synorisporites’ papillensis McGregor 1973SP12
                  `--C. paranensis Gerrienne et al. 2001SP12
Apo-Tracheophyta incertae sedis:
  Steganotheca Edwards 1970EE86
    |--‘Cooksonia’ downtonensis Heard 1939EE86
    `--S. striata Edwards 1970EE86
  Uskiella spargens Shute & Edwards 1989EE86
  Tortilicaulis Edwards 1979G97
    |--T. offaeus Edwards et al. 1994G97
    `--T. transwalliensis Edwards 1979G97
    |--D. maraisiaEE86
    `--D. pulchraEE86
  Eogaspesiea gracilisEE86
    |--P. laevigatus Steemans 1989SP12
    |--P. magnus Hughes & Playford 1961P63
    |--P. perinatus Hughes & Playford 1961P63
    `--P. tessellatusPSC99
  Emphanisporites McGregor 1961EE86, SP12
    |--E. annulatus McGregor 1961SP12
    |--E. micrornatus Richardson & Lister 1969LT80
    |--E. neglectusLT80
    |--E. obscurus McGregor 1961LT80
    |--E. protophanus Richardson & Ioannides 1973SP12
    |--E. rotatus (McGregor) McGregor 1973LT80
    `--E. zavallatus Richardson et al. 1982SP12
         |--E. z. var. zavallatusSP12
         `--E. z. var. gedinniensis Steemans & Gerrienne 1984SP12
  Rebskia Schweitzer 2000 [Rebskiaceae]S00
    `--*R. musaeformis Schweitzer 2000S00
    |--A. bollandensisSP12
    |--A. minuta McGregor 1973LT80
    `--A. richardsonii Richardson et al. 2001SP12
    |--R. dubiusW97
    |--R. frivolus Chibrickova 1959LT80
    |--R. pychovii Naumova 1953LT80
    `--R. triangulatus (Streel) Streel 1967LT80
  Barinophytaceae [Barinophytales]C93
    |--Krithodeophyton croftii Edwards 1968C93
  Schizostege Hillebr. 1888KC01

Cooksonia Lang 1937 [incl. Ambitisporites Hoffmeister 1959, Streelispora Richardson & Lister 1969, Synorisporites Richardson & Lister 1969]SP12

Cooksonia pertoni ssp. apiculispora [incl. Streelispora newportensis (Chaloner & Streel) Richardson & Lister 1969]SP12

‘Rhynia’ majorD98 [=Aglaophyton major (Kidston & Gwynne-Vaughan) Edwards 1986D98, C93; incl. Lyonophyton rhyniensis Remy & Remy 1980SP12]

*Type species of generic name indicated


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. Pteridophyta. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 779–794. Chapman & Hall: London.

[C02] Clifford, H. T. 2002. A new Devonian fern, Fanningopteris wyattii, from Queensland. Memoirs of the Queensland Museum 48: 71–77.

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

[EE86] Edwards, D., & D. S. Edwards. 1986. A reconsideration of the Rhyniophytina Banks. In: Spicer, R. A., & B. A. Thomas (eds) Systematic and Taxonomic Approaches in Palaeobotany. Systematics Association Special Volume 31: 199–220. Clarendon Press: Oxford.

[G97] Gerrienne, P. 1997. The fossil plants from the Lower Devonian of Marchin (northern margin of Dinant Synclinorium, Belgium): V. Psilophyton genseliae sp. nov., with hypotheses on the origin of Trimerophytina. Review of Palaeobotany and Palynology 98: 303–324.

[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).

[LT80] Loeffler, E. J., & G. Thomas. 1980. A new pteraspidid ostracoderm from the Devonian Senni Beds Formation of south Wales and its stratigraphic significance. Palaeontology 23: 287–396.

[PSC99] Paton, R. L., T. R. Smithson & J. A. Clack. 1999. An amniote-like skeleton from the early Carboniferous of Scotland. Nature 398: 508–513.

[P63] Playford, G. 1963. Lower Carboniferous microfloras of Spitsbergen. Part two. Palaeontology 5 (4): 619–678.

[S00] Schweitzer, H. J. 2000. Neue Pflanzenfunde im Unterdevon der Eifel (Deutschland). Senckenbergiana Lethaea 80 (2): 371–395.

[SP12] Steemans, P., E. Petus, P. Breuer, P. Mauller-Mendlowicz & P. Gerrienne. 2012. Palaeozoic innovations in the micro- and megafossil plant record: from the earliest plant spores to the earliest seeds. In: Talent, J. A. (ed.) Earth and Life: Global biodiversity, extinction intervals and biogeographic perturbations through time pp. 437–477. Springer.

[SW04] Steemans, P., & C. H. Wellman. 2004. Miospores and the emergence of land plants. In: Webby, B. D., F. Paris, M. L. Droser & I. G. Percival (eds) The Great Ordovician Biodiversification Event pp. 361–366. Columbia University Press.

[W97] Wang N. 1997. Restudy of thelodont microfossils from the lower part of the Cuifengshan Group of Qujing, eastern Yunnan, China. Vertebrata PalAsiatica 35 (1): 1–17.

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