Specimens of Calceola sandalina from the University of Corunna, photographed by Porshunta.

Belongs within: Cnidaria.
Contains: Pachythecaliina, Lithostrotionidae, Cyathophyllidae.

The Rugosa are a group of corals known from the Palaeozoic. They are characterised by the presence of single mesenteries (represented in the fossils by septa) that were added in four sectors (Gudo 2002). Gudo (2002) suggested that rugose corals could be subdivided into two groups based whether new septa were added serially (Seriales) or symmetrically (Symmetricales) in each of the four sectors, but noted that it is as yet unclear which of these divisions many taxa would fall into. The relationships of rugose corals to modern scleractinian corals remains open to debate: many authors have presumed them to be an independent lineage that was not directly connected to the Scleractinia but some have interpreted the Rugosa as including the ancestors of the latter.

<==Rugosa [Tetracorallia]
    |--Calceola [Seriales] G02
    |    |--C. calceoloides [=Rhizophyllum calceoloides] TC71
    |    `--C. sandalina TC71
    `--Symmetricales G02
         |--Goniophyllum pyramidale G02
         |--Argutastraea quadrigemina G02
         |--Araeopoma G02
         `--Polycoelia G02

Rugosa incertae sedis:
  Enterolasma EB01
  Spineria SchouppĂ© & Stacul 1959 SR02
  Pachythecaliina SR02
  Heritschioides F79
  Craterophyllum F79
    |--C. eifeliense Fedorowski 1967 F79
    `--C. verticillatum F79
  Timania rainbowensis F79
  Neokoninckophyllum kansasense F79
  Heliophyllum F79
    |--H. chillagoensis (Etheridge 1911) [=Cyathophyllum chillagoensis] F71
    `--H. halli F79
  Tabulophyllum F79
    |--T. rotundum Spassky 1960 F79
    `--T. schlueteri F79
  Clisaxophyllum ava F79
    |--C. a. ava F79
    `--C. a. atetsuense Minato & Nakazawa 1957 F79
  Bothrophyllum dobroljubovae F79
  Spirophyllum geminum Fedorowski 1970 F79
  Entelophyllum F79
    |--E. articulatum F79
    |--E. latum Hill 1940 F71
    `--E. yassense (Etheridge 1892) [=Heliophyllum yassense, Xylodes yassense] F71
  Kodonophyllum truncatum F79
  Lithostrotionidae Nu79
  Iovaphyllum F79
  Brachyelasma irregulare He in Kong & Huang 1978 RZJ04
  Tongxinophyllum tongxinensis Gao 1987 RZJ04
  Ipciphyllum arnouldi RS01
  Plerophyllina RS01
    |  i. s.: Calophyllum RS01
    |           |--C. clavataseptatum Ding in Ding, Xia et al. 1984 DX84
    |           |--C. cylindricum DX84
    |           `--C. gemmatum Iljina 1984 RS01
    `--Plerophyllidae WSZ04
         |--Commutia Fedorowski 1973 WSZ04
         |    |--*C. szulczewskii Fedorowski 1973 WSZ04
         |    `--C. exoleta Wang, Sugiyama & Zhang 2004 WSZ04
         `--Plerophyllum DX84
              |--P. cainodon [=Zaphrentis (Plerophyllum) cainodon] F71
              |--P. clavatum DX84
              |--P. culleni (Etheridge 1891) [=Zaphrentis (Plerophyllum) culleni] F71
              |--P. crassoseptatum Ding in Ding, Xia et al. 1984 DX84
              |--P. giganteum Ding in Ding, Xia et al. 1984 DX84
              |--P. gregorianum [=Zaphrentis (Plerophyllum) gregoriana] F71
              |--P. multiseptatum Ding in Ding, Xia et al. 1984 DX84
              |--P. neimonggolense Ding in Ding, Xia et al. 1984 DX84
              |--P. regulare DX84
              `--P. robustum [=Zaphrentis (Plerophyllum) robusta] F71
  Pentaphyllum WSZ04
    |--P. hibernicum WSZ04
    |--P. hithis WSZ04
    `--P. sinense Ding in Ding, Xia et al. 1984 DX84
  Ufmia infracarbonica WSZ04
  Cyathaxonia WSZ04
    |--C. cornu WSZ04
    |--C. cynodon H04
    `--C. qinghaiensis CSZ03
  Craspedophyllum subcaespitosum T10
  Acinophyllum davisi No79
  Edaphophyllum sulcatum No79
  Omphyma turbinata H04
  Cyathophyllidae H04
  Pachyphyllum devoniense H04
  Menophyllum tenuimarginum H04
  Hadrophyllum multiradiatum H04
  Clisiophyllum turbinatum H04
  Acervularia ananas H04
  Syringophyllum organum H04
  Lophotichium S00
    |--L. bellum Ding in Ding, Xia et al. 1984 DX84
    |--L. convexotabulatum Ding in Ding, Xia et al. 1984 DX84
    `--L. dugoutense S00
  Protomacgeea S00

*Type species of generic name indicated


[CSZ03] Chen, Z.-Q., G. R. Shi & L.-P. Zhan. 2003. Early Carboniferous athyridid brachiopods from the Qaidam Basin, northwest China. Journal of Paleontology 77: 844–862.

[DX84] Ding Y., Xia G., Duan C., Li W., Liu X. & Liang Z. 1984. Study on the early Permian stratigraphy and fauna in Zhesi district, Nei Mongol Zizhiqu (Inner Mongolia). Bulletin Tianjin Institure Geol. Min. Res. 10.

[EB01] Eckert, J. D., & C. E. Brett. 2001. Early Silurian (Llandovery) crinoids from the Lower Clinton Group, western New York State. Bulletins of American Paleontology 360: 1–88.

[F79] Fedorowski, J. 1979. On some aspects of coloniality in Permian corals. In: Larwood, G., & B. R. Rosen (eds) Biology and Systematics of Colonial Organisms pp. 155–171. Academic Press: London.

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

[G02] Gudo, M. 2002. Soft body reconstructions of Palaeozoic corals: implications for the system of Anthozoa (Coelenterata). Lethaia 35: 328–344.

[H04] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

[No79] Norris, A. W. 1979. Devonian in the Western Hemisphere. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A218–A253. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[Nu79] Nudds, J. R. 1979. Coloniality in the Lithostrotionidae (Rugosa). In: Larwood, G., & B. R. Rosen (eds) Biology and Systematics of Colonial Organisms pp. 173–192. Academic Press: London.

[RS01] Roniewicz, E., & J. Stolarski. 2001. Triassic roots of the amphiastraeid scleractinian corals. Journal of Paleontology 75 (1): 34–45.

[RZJ04] Rong, J.-Y., R.-B. Zhan & J. Jin. 2004. The Late Odovician and Early Silurian pentameride brachiopod Holorhynchus Kiaer, 1902 from north China. Journal of Paleontology 78 (2): 287–299.

[S00] Stolarski, J. 2000. Origin and phylogeny of Guyniidae (Scleractinia) in the light of microstructural data. Lethaia 33: 13–38.

[SR02] Stolarski, J., & A. Russo. 2002. Microstructural diversity of the stylophyllid (Scleractinia) skeleton. Acta Palaeontologica Polonica 47 (4): 651–666.

[T10] Taylor, T. G. 1910. The Archaeocyathinae from the Cambrian of South Australia with an account of the morphology and affinities of the whole class. Memoirs of the Royal Society of South Australia 2 (2): 55–188.

[TC71] Thomson, K. S., & K. S. W. Campbell. 1971. The structure and relationships of the primitive Devonian lungfish—Dipnorhynchus sussmilchi (Etheridge). Yale University Peabody Museum of Natural History Bulletin 38: 1–109.

[WSZ04] Wang, X.-D., Sugiyama T. & F. Zhang. 2004. Intraspecific variation in a new solitary rugose coral, Commutia exoleta, from the Lower Carboniferous of the Baoshan Block, southwest China. Journal of Paleontology 78 (1): 77–83.

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