Belongs within: Hexacorallia.
Contains: Pachythecaliina, Streptelasmatida, Cystiphyllida, Lithostrotionidae, Disphyllidae, Phillipsastreidae, Verbeekiellidae, Waagenophyllidae, Plerophyllina.
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.
Recognised subgroups of the Rugosa include the Stauriida, in which the septa are laminar and composed of continuous trabeculae.
Rugose horns
Published 4 November 2024
Life in the Palaeozoic era was very different to today. Many of the organisms that lived in that time represented lineages that resemble nothing alive today. And then there were those that may have resembled modern forms at first glance but on closer inspection would have looked weirdly different, such as the rugose corals.
‘Coral’ is a general term for any cnidarian producing a hard, often mineralised skeleton, but the most familiar corals in the modern era are members of the order Scleractinia. Though dominant in many modern oceans, particularly in their formation of tropical reefs, scleractinians did not definitively appear in the fossil record until the Middle Triassic. During the preceding Palaeozoic, the role of ‘corals’ was taken by other cnidarian taxa whose relationship to the scleractinians remains the subject of much debate. The rugose corals or tetracorals were one such group that first definitely appeared in the Middle Ordovician (though some earlier taxa have been suggested as possible Rugosa going back to the Middle Cambrian). The earliest known rugose corals were solitary, typically with a more or less conical skeleton (giving many such forms the alternative name of ‘horn corals’), but colonial forms later became common. Despite their superficial similarity, rugose corals differed from scleractinian corals in having skeletons of calcite rather than aragonite. They also differed in their manner of growth. In both modern and rugose corals, the living polyp was supported in its mineralised cup by radiating septa. However, modern and rugose corals differ in the order in which new septa were added (Oliver 1996). In scleractinians, growth of septa is described as ‘cyclic’, with new septa being added in between older septa all around the cup. In rugose corals, growth of septa was ‘serial’, with septa being added one after the other around each quadrant of the cup. As a result of this difference in growth pattern, scleractinian corallites typically have a more radial appearance than rugose corallites which have a more bilateral appearance. Hence the alternative name for Rugosa of Tetracorallia, referring to the four-way symmetry of rugose corals instead of the six-way symmetry of modern Hexacorallia.
Though common in Palaeozoic deposits, rugose corals were rarely the dominant reef-builders in the manner of modern scleractinians. A major factor appears to have been their different way of life. Rugose corals did not typically cement themselves to their substrate but instead lay free on the sea floor. Solitary forms might be held upright by having the lower end of the cup embedded into the sediment, or they might lie on their side with the open end curved upwards. Colonial forms would have been held in place by the weight of the colony itself. But because they were not fixed in place, rugose corals were less able to tolerate high-energy environments. Most rugose corals inhabited the relatively calm waters of the fore-reef zone whereas the reef itself was occupied by more firmly attached organisms such as tabulate corals and stromatoporoids (Clarkson 1993).
Rugose corals thrived through most of the Palaeozoic; though their numbers crashed at the end of the Devonian, they rapidly recovered during the Carboniferous. Nevertheless, they disappeared entirely at the end of the Permian. A gap of some fifteen million years would ensue before the first scleractinians appeared during the Triassic. As noted above, there has been much disagreement about how the scleractinians related to their rugose fore-runners. Were they descendants of some remnant population of rugose corals that had weathered the end-Permian extinction, or were they an independent lineage descended from non-mineralised ancestors? Supporters of their relationship point to their similarities in structure and morphology, detractors to their differences in composition and development as well as that pesky post-Permian gap (which appears to be a genuine gap rather than a simple deficiency in the fossil record). It seems likely that this question will remain disputed for some time yet.
Systematics of Rugosa
<==Rugosa [Proteroseptata, Tetracorallia, Tetraseptata]
|--Calceola [Seriales]G02
| |--C. calceoloides [=Rhizophyllum calceoloides]TC71
| `--C. sandalina (Linnaeus 1771)NS93
`--SymmetricalesG02
| i. s.: Argutastraea quadrigeminaG02
|--CystiphyllidaG02
`--StauriidaJB12
| i. s.: LithostrotionidaeNu79
| CyathophyllidaeJB12
| |--RadiophyllumNS93
| |--Commutatophyllum cincinnatum Kaplan 1971NS93
| `--CyathophyllumH04
| |--C. articulatumH04
| |--C. expansumH04
| |--C. marminiH04
| |--C. patula [=Entelophyllum yassense var. patulum]F71
| |--C. retiforme Etheridge 1891F71
| `--C. zaphrentoides Etheridge 1891F71
| PycnostylidaeNS93
| |--ProtopilophyllumNS93
| |--Cyathopaedium paucitabulatum (Schluter 1883)NS93
| |--Depasophyllum adnetum Grabau 1936NS93
| |--Fletcherina simplex (Yabe & Hayasaka 1915)NS93
| `--PycnostylusF71
| |--P. congregationis (Etheridge 1907) [=Tryplasma congregationis]F71
| `--P. dendroides (Etheridge 1907) [=Tryplasma dendroides]F71
| Neocolumnaria [Neocolumnariidae]NS93
| `--N. vagranensis Soshkina 1949NS93
| Centristela [Centristelidae]NS93
| |--C. anavarensis (Goryanov in Bulvanker et al. 1968)NS93
| `--C. fasciculata Tsyganko 1967NS93
| AmplexidaeNS93
| |--Amplexoides severnensis (Parks 1915)NS93
| |--Gorskyella tschigariensis (Fomichev 1953)NS93
| `--AmplexusF71
| |--A. arundinaceus [=Zaphrentis (Plerophyllum) arundinaceus]F71
| `--A. coralloides [incl. A. coralloides var. permocarbonica]G31
| KiziliidaeNS93
| |--Kizilia concavitabulata Degtyarev 1965NS93
| `--Melanophyllidium lativesiculosum Kropacheva 1966NS93
| Ditoecholasma [Ditoecholasmatidae]NS93
| |--D. fanninganum (Safford 1869)NS93
| `--D. lawrencense Sutherland 1965NS93
| PaliphyllidaeNS93
| |--ProtocyathactisNS93
| |--SumsarophyllumNS93
| |--Neopaliphyllum soshkinae Zheltonogova 1961NS93
| `--PaliphyllumNS93
| |--P. lamniscatumG31
| `--P. mapingenseG31
| KodonophyllidaeNS93
| |--Kodonophyllum truncatumF79
| |--Schlotheimophyllum patellatus (Schlotheim 1820)NS93
| |--Sinochlamydophyllum crassiseptatum Guo 1976NS93
| `--Zelophyllia tabulatum (Soshkina 1937)NS93
| MucophyllidaeNS93
| |--Kungejophyllum ajagusense Sultanbekova 1971NS93
| |--Briantia repleta Barrois 1889NS93
| `--PseudamplexusNS93
| |--P. ligeriensis (Barrois 1889)NS93
| `--P. princeps (Etheridge 1907) [=Tryplasma princeps]F71
| AmsdenoididaeNS93
| |--Amsdenoides acutiannulatum (Amsden 1949)NS93
| `--Multicarinophyllum multicarinatum Spasskiy 1965NS93
| CyathaxoniidaeNS93
| |--Columnaxon angelae Scrutton 1971NS93
| `--CyathaxoniaWSZ04
| |--C. cornuWSZ04
| |--C. cynodonH04
| `--C. qinghaiensisCSZ03
| PetraiidaeNS93
| |--Protozaphrentis minor Yu 1957NS93
| |--Petraiella kielcensis Rozkowska 1969NS93
| `--PetraiaM87
| |--P. binaM87
| `--P. corniculumM87
| MetriophyllidaeNS93
| |--Duncanella borealis Nicholson 1874NS93
| |--Asserculinia prima Schouppe & Stacul 1959NS93
| |--Lophotichium Moor & Jeffords 1945DX84
| | |--*L. vescum Moore & Jeffords 1945DX84
| | |--L. bellum Ding in Ding, Xia et al. 1984DX84
| | |--L. convexotabulatum Ding in Ding, Xia et al. 1984DX84
| | `--L. dugoutenseS00
| `--Diphycarinophyllum Ding in Ding, Xia et al. 1984DX84
| `--*D. baotegense Ding in Ding, Xia et al. 1984DX84
| LaccophyllidaeNS93
| |--Laccophyllum acuminatum Simpson 1900NS93
| `--AmplexocariniaNS93
| KielcephyllidaeNS93
| |--Kielcephyllum cupulum Rozkowska 1969NS93
| |--Kozlowskinia flos Rozkowska 1969NS93
| `--Thecaxon rozkowskae Weyer 1978NS93
| Lindstroemia [Lindstroemiidae]NS93
| `--L. columnaris Nicholson & Thomson 1876NS93
| HadrophyllidaeNS93
| |--HadrophyllumNS93
| | |--H. multiradiatumH04
| | `--H. orbignyi Milne Edwards & Haime 1850NS93
| `--MicrocyclusNS93
| |--M. discus Meek & Worthen 1868NS93
| `--M. multiradiatusNo79
| Combophyllum [Combophyllidae]NS93
| |--C. ibericum Plusquellec in Marin & Plusquellec 1973NS93
| `--C. osismorum Milne Edwards & Haime 1850NS93
| EntelophyllidaeNS93
| |--Petrozium dewari Smith 1930NS93
| |--Kysylagatophyllum michnevitchi Kaplan 1971NS93
| |--ScyphophyllumNS93
| `--EntelophyllumF79
| |--E. articulatumF79
| |--E. latum Hill 1940F71
| `--E. yassense (Etheridge 1892) [=Heliophyllum yassense, Xylodes yassense]F71
| ExpressophyllidaeNS93
| |--Pseudopilophyllum moyerense Ivanovskiy 1963NS93
| |--Contortophyllum tchernovi Strelnikov 1968NS93
| `--MiculaNS93
| |--M. antiqua Sytova 1952NS93
| |--M. decussataVH02
| |--M. muraK79
| `--M. simplex (Strelnikov 1968)NS93
| ArachnophyllidaeNS93
| |--Arachnophyllum epistomoides Etheridge 1909F71
| |--Angullophyllum warrisi McLean 1974NS93
| `--CraterophyllumNS93
| |--C. eifeliense Fedorowski 1967F79
| `--C. verticillatumF79
| KetophyllidaeNS93
| |--Dentilasma honorabilis Ivanovskiy 1962NS93
| |--Heterolasma foerstei Ehlers 1919NS93
| |--Chavsakia chavsakiensis Lavrusevich 1959NS93
| `--Nataliella poslavskajae Sytova in Sytova & Ulitina 1966NS93
| KyphophyllidaeNS93
| |--DonacophyllumNS93
| |--BouvierphyllumNS93
| |--MikkwaphyllumNS93
| |--WapitiphyllumNS93
| |--ParasmithiphyllumNS93
| |--TarphyphyllumNS93
| `--KakisaphyllumNS93
| EndophyllidaeNS93
| |--Endophyllum schluteri Etheridge 1898F71
| |--Mictocystis endophylloides Etheridge 1908NS93
| `--SmithiphyllumNS93
| SpongophyllidaeNS93
| |--Heterospongophyllum simplex He in Kong & Huang 1978NS93
| |--Neovepresiphyllum immersum (Hill 1942)NS93
| `--Spongophyllum Edwards & Haime 1851E13
| |--S. bipartitum (Etheridge 1889)E13, F71 [=Lonsdaleia (Spongophyllum) bipartitaF71]
| |--S. enorme Etheridge 1913E13 [=Crinophyllum enormeF71, Yassia enormisF71]
| |--S. giganteum Etheridge 1899E13, F71 [=Australophyllum giganteumF71]
| |--S. halysitoides Etheridge 1918F71
| `--S. kunthi Schlüter 1881E13
| PtenophyllidaeNS93
| |--Cymatelasma corniculum Hill & Butler 1936NS93
| `--Hankaxis tinocystis (Frech 1885)NS93
| FasciphyllidaeNS93
| |--Fasciphyllum conglomeratum (Schluter 1881)NS93
| `--Crista compacta Tsyganko 1971NS93
| StringophyllidaeNS93
| |--RhegmaphyllumNS93
| |--StringophyllumNS93
| |--ParasociophyllumNS93
| `--SociophyllumNS93
| LykophyllidaeNS93
| |--HolophragmaNS93
| |--Pseudophaulactis lykophylloides Zaprudskaya in Ivanovskiy 1963NS93
| |--Rukhinia cuneata Strelnikov 1963NS93
| |--Zeravschania prima Lavrusevich 1964NS93
| `--Camurophyllum camurum Kravtsov 1966NS93
| HalliidaeNS93
| |--Kobeha walcotti Merriam 1974NS93
| |--AulacophyllumNS93
| `--AspasmophyllumNS93
| AcervulariidaeNS93
| |--Acervularia ananasH04
| `--Oliveria planotabulata Sutherland 1965NS93
| ColumnariidaeNS93
| |--Circumtextiphyllum annulatum Kaplan 1971NS93
| `--ColumnariaNS93
| |--C. pauciseptata Etheridge 1897 [=Columnopora (l. c.) pauciseptata]F71
| `--C. sulcata Goldfuss 1826NS93
| DisphyllidaeNS93
| PhillipsastreidaeNS93
| ZaphrentidaeNS93
| |--HeliophyllumNS93
| | |--H. chillagoensis (Etheridge 1911) [=Cyathophyllum chillagoensis]F71
| | `--H. halliF79
| `--Zaphrentis Rafinesque & Clifford 1820W77
| |--Z. corniculaH04
| |--Z. excentrica Meek 1872W77
| `--Z. phymatodes Etheridge 1891F71
| BethanyphyllidaeNS93
| |--Bethanyphyllum robustum (Hall 1876)NS93
| `--Tortophyllum cysticum (Winchell 1866)NS93
| Campophyllum [Campophyllidae]NS93
| |--C. columnare Etheridge 1891F71
| |--C. cylindricum (Onoprienko 1979)NS93
| |--C. flexuosum (Goldfuss 1826)NS93
| |--C. lindstromiF71
| |--C. nikitiniG31
| |--C. recessum Hill 1940F71
| `--C. schrenkiG31
| Implicophyllum [Ptychophyllidae]NS93
| `--I. vesiculosum Sytova in Sytova & Ulitina 1966NS93
| StereolasmatidaeNS93
| |--Amplexiphyllum hamiltoniae (Hall 1876)NS93
| |--LopholasmaNS93
| | |--L. carinatum Simpson 1990NS93
| | `--L. ilitschenseDX84
| |--Stewartophyllum intermittens (Hall 1876)NS93
| `--StereolasmaNS93
| |--S. minusG31
| `--S. rectum (Hall 1876)NS93
| AntiphyllidaeNS93
| |--PseudoclaviphyllumNS93
| |--Lytvolasma asymmetricum Soshkina 1925NS93
| `--FasciculophyllumNS93
| |--F. dobroljubovae Rozkowska 1969NS93
| `--F. rushianum Rozkowska 1969NS93
| HapsiphyllidaeNS93
| |--Adradosia barroisi Birenheide & Soto 1977NS93
| `--Duplophyllum zaphrentoides Koker 1924NS93
| ZaphrentoididaeNS93
| |--Zaphrentoides ecavatus Hill 1954NS93
| |--Amplexizaphrentis adyrensis (Soshkina 1960)NS93
| `--Basleophyllum indicum (Koker 1924)NS93
| Anisophyllum [Anisophyllidae]NS93
| `--A. agassizi Milne Edwards & Haime 1850NS93
| Endothecium [Endotheciidae]NS93
| `--E. apertum Koker 1924NS93
| AdamanophyllidaeNS93
| |--Tachyphyllum artyshtense Dobrolyubova in Soshkina et al. 1962NS93
| `--Adamanophyllum incertum Vasilyuk 1959NS93
| PentaphyllidaeNS93
| |--Oligophyllum quinqueseptatum Pocta 1902NS93
| |--PentamplexusNS93
| `--Pentaphyllum de Koninck 1872 [incl. Cryptophyllum Carruthers 1919]DX84
| |--*P. armatum de Koninck 1872DX84
| |--P. gracileF97
| |--P. hibernicumWSZ04
| |--P. hithisWSZ04
| |--P. pauperumDX84
| `--P. sinense Ding in Ding, Xia et al. 1984DX84
| LophophyllidaeNS93
| |--Lophophyllidium spinosumNS93, DX84
| |--Lophophyllum Edwards & Haime 1850W77
| | |--L. caninoides Gorskiy 1935NS93
| | |--L. corniculumF71
| | |--L. densumCSZ03
| | `--L. proliferum (McChesney 1860) (see below for synonymy)W77
| `--Lophocarinophyllum Grabau 1922DX84
| `--L. yakovlevi [incl. L. yakovlevi f. minuta Fomitschev 1941]DX84
| Timorphyllum [Timorphyllidae]NS93
| `--T. wanneri Gerth 1921NS93
| VerbeekiellidaeNS93
| CyathopsidaeNS93
| |--Arctophyllum intermedium (Toula 1875)NS93
| |--FomichevellaNS93
| |--Paracaninia Chi 1937NS93, DX84
| | |--P. sinensis Chi 1937NS93
| | `--P. tzuchiangensis (Huang 1932) [=Caninia tzuchiangensis]DX84
| `--CaniniaNS93
| |--C. cornucopiae Michelin 1840NS93
| |--C. dorlodotiZP86
| `--C. murataiG31
| BothrophyllidaeNS93
| |--Bothrophyllum dobroljubovaeF79
| |--Hornsundia lateseptata Fedorowski 1965NS93
| `--CaninophyllumNS93
| |--C. adapertum (Onoprienko 1979)NS93
| `--C. sumphuens (Etheridge 1891) [=Zaphrentis sumphuens]F71
| UraliniidaeNS93
| |--Uralinia megacystosa Gorskiy 1935NS93
| |--Enygmophyllum dubium Gorskiy 1935NS93
| |--CystophentisNS93
| |--Bifossularia ussowi (Gabuniya 1919)NS93
| |--Keyserlingophyllum keyserlingophylloidesNS93, ZP86
| `--Liardiphyllum hagei Sutherland 1954NS93
| EndamplexidaeNS93
| |--Endamplexus dentatus Koker 1924NS93
| `--Spineria Schouppé & Stacul 1959NS93, SR02
| `--S. diplochone (Koker 1924)NS93
| AulophyllidaeNS93
| |--Heintzella multiseptata Fedorowski 1967NS93
| `--AulophyllumF71
| |--A. davidis Etheridge 1891F71
| `--A. permicumG31
| EkvasophyllidaeNS93
| |--ZaphriphyllumNS93
| |--Faberophyllum occultum Parks 1951NS93
| |--Turbophyllum multiconum Parks 1951NS93
| `--EkvasophyllumCSZ03
| |--E. heijianshanensisCSZ03
| `--E. inclinatum Parks 1951NS93
| PalaeosmiliidaeNS93
| |--Palaeosmilia planumCSZ03
| `--PalastraeaNS93
| AphrophyllidaeNS93
| |--Aphrophyllum hallense Smith 1920F71
| |--Naoides rangariensis Pickett 1967NS93
| |--AphrophylloidesNS93
| |--CoenaphrodiaNS93
| |--MerlewoodiaNS93
| |--NothaphrophyllumNS93
| `--Symplectophyllum mutatumNS93, F71
| DurhaminidaeNS93
| |--Protolonsdaleiastraea atbassarica Gorskiy 1932NS93
| `--Tanbaella izuruhense (Sakaguchi & Yamagiwa 1958)NS93
| AxophyllidaeNS93
| |--AxophylumNS93
| `--LonsdaleiaNS93
| |--L. chihsiaensisG31
| |--L. chinensisG31
| |--L. floriformis [incl. L. floriformis mut. carnica]G31
| |--L. kiangsiensisG31
| `--L. stelckiCSZ03
| PetalaxidaeNS93
| |--Paralithostrotion podboriense (Dobrolyubova 1958)NS93
| |--Lytvophyllum tschernowi (Soshkina in Soshkina et al. 1941)NS93
| `--Lithostrotionella unica (Yabe & Hayasaka 1915)NS93
| GeyerophyllidaeNS93
| |--Amygdalophylloides Dobrolyubova & Kabakovich 1948NS93, DX84
| | |--*A. ivanovi (Dobrolyubova 1937) [=Amygdalophyllum ivanovi]DX84
| | | |--A. i. ivanoviDX84
| | | `--A. i. kovroviDX84
| | `--A. asteroides Ding in Ding, Xia et al. 1984DX84
| |--DarwasophyllumNS93
| |--Carinthiaphyllum kahleri Heritsch 1936NS93
| `--Lonsdaleoides boswelli Heritsch 1936NS93
| Koninckocarinia [Koninckocariniidae]NS93
| `--K. flexuosa (Dobrolyubova 1937)NS93
| WaagenophyllidaeNS93
| PseudopavonidaeNS93
| |--AmygdalophyllidiumNS93
| |--HiroshimaphyllumNS93
| |--OmniphyllumNS93
| |--OzakiphyllumNS93
| `--PseudopavonaNS93
|--PlerophyllinaG02
|--StauriinaWE04
| |--CyathophylloidesWE04
| |--CrenulitesWE04
| `--Favistina [Stauriidae]NS93
| `--F. undulata (Bassler 1950)NS93
`--CalostylinaWE04
|--CalostylidaeNS93
| |--CalostylisNS93
| |--NingnanophyllumNS93
| `--YohophyllumNS93
`--LambelasmatidaeNS93
|--Lambeophyllum profundum (Conrad 1843)NS93
|--Prototryplasma oroniana Ivanovskiy 1963NS93
`--LambelasmaWE04
Rugosa incertae sedis:
EnterolasmaEB01
PachythecaliinaSR02
HeritschioidesF79
Timania rainbowensisF79
Neokoninckophyllum kansasenseF79
TabulophyllumF79
|--T. rotundum Spassky 1960F79
`--T. schlueteriF79
Clisaxophyllum avaF79
|--C. a. avaF79
`--C. a. atetsuense Minato & Nakazawa 1957F79
Spirophyllum geminum Fedorowski 1970F79
IovaphyllumF79
Brachyelasma irregulare He in Kong & Huang 1978RZJ04
Tongxinophyllum tongxinensis Gao 1987RZJ04
Craspedophyllum subcaespitosumT10
Acinophyllum davisiNo79
Edaphophyllum sulcatumNo79
Omphyma turbinataH04
Pachyphyllum devonienseH04
Menophyllum tenuimarginumH04
Clisiophyllum turbinatumH04
ProtomacgeeaS00
DiffingiidaeNS93
|--Diffingia valida Fedorowski 1985NS93
`--TurgidiffiaNS93
Plerodiffia [Plerodiffiidae]NS93
`--P. eaglebuttensis Fedorowski 1985NS93
StreptelasmatidaWE04
Sochkineophyllum japonicumN00
|--S. j. japonicumN00
`--S. j. pauciseptatumN00
Lophophyllum proliferum (McChesney 1860) [=Cyathaxonia prolifera; incl. L. proliferum var. sauridens White 1877]W77
*Type species of generic name indicated
References
[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.
Clarkson, E. N. K. 1993. Invertebrate Palaeontology and Evolution 3rd ed. Chapman and Hall: London.
[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.
[E13] Etheridge, R., Jr. 1913. A very remarkable species of Spongophyllum from the Upper Silurian rocks of New South Wales. Records of the Australian Museum 10 (3): 35–37, pls 4–7.
[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.
[F97] Fedorowski, J. 1997. Rugosa and Scleractinia—a commentary on some methods of phylogenetic reconstruction. Acta Palaeontologica Polonica 42 (3): 446–456.
[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.
[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.
[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.
[JB12] Johnson, M. E., & B. G. Baarli. 2012. Development of intertidal biotas through Phanerozoic time. In: Talent, J. A. (ed.) Earth and Life: Global biodiversity, extinction intervals and biogeographic perturbations through time pp. 63–128. Springer.
[K79] Kauffman, E. G. 1979. Cretaceous. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A418–A487. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).
[M87] Mitchell, J. 1887. Notes on the geology of Bowning, N.S.W. Proceedings of the Linnean Society of New South Wales, series 2, 1 (4): 1193–1204.
[N00] Niko, S. 2000. Youngest record of tentaculitoids: Hidagaienites new genus from near the Carboniferous–Permian boundary in central Japan. Journal of Paleontology 74 (3): 381–385.
[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.
[NS93] Nudds, J. R., & J. J. Sepkoski Jr. 1993. Coelenterata. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 101–124. Chapman & Hall: London.
Oliver, W. A., Jr. 1996. Origins and relationships of Paleozoic coral groups and the origin of the Scleractinia. Paleontological Society Papers 1: 107–134.
[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.
[VH02] Vallejo, C., P. A. Hochuli, W. Winkler & K. von Salis. 2002. Palynological and sequence stratigraphic analysis of the Napo Group in the Pungarayacu 30 well, sub-Andean zone, Ecuador. Cretaceous Research 23: 845–859.
[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.
[W77] White, C. A. 1877. Report upon the invertebrate fossils collected in portions of Nevada, Utah, Colorado, New Mexico, and Arizona, by parties of the expeditions of 1871, 1872, 1873, and 1874. U.S. Geographical Surveys West of the One Hundredth Meridian 4 (1): 1–219, pls 1–21.
[ZP86] Zhang, R., & J. Pojeta, Jr. 1986. New bivalves from the Datang Stage, Lower Carboniferous, Guangdong Province, China. Journal of Paleontology 60 (3): 669–679.