Holotype of Onchotrochus hatifnatus, from Dr H. Eliasova.

Belongs within: Hexacorallia.
Contains: Volzeioidea, Stylophyllidae, Faviina, Turbinoliidae, Caryophyllia, Agariciidae, Siderastreidae, Astrocoeniina, Poritidae, Guyniidae, Flabellum, Goniopora.

The Caryophylliina are a group of corals characterised by the possession of laminar septa that are composed of one fan of numerous, simple trabeculae (Stolarski 2000). The monophyly of this group is uncertain. Among the taxa included in the Caryophylliina are the Guyniidae, represented in the modern fauna by a single living species, Guynia annulata, distinguished from other living coralls by smooth-faced septa that are deeply recessed relative to the epithecal rim, with undifferentiated calcification centres, and an aseptate basal section of the initial ontogenetic stage (Stolarski 2000).

The Dendrophylliidae are a group of corals characterised by irregularly porous thecal walls. Most species lack zooxanthellae, and include both solitary and colonial forms.

The Fungiina are a group of corals including both solitary and colonial forms, with fenestrate septa with beaded or dentate margins (Cairns 2002).

Coral love
Published 17 November 2008
The solitary coral Javania erhardti, a member of the ‘caryophylliine’ family Flabellidae, from here.

Cnidarian classification can be a terrible thing. Like many other groups of soft-bodied animals, useful characters for distinguishing taxa can be few and far between, and those few characters that are available may be difficult to identify and readily subject to evolutionary change. Corals are no exception. The current generally-used classification of living corals divides them between seven suborders, distinguished from each other primarily by the structure of their septa, the ribs of skeletal material within each polyp. The Caryophylliina, for instance, have simple septa, with little in the way of internal ornamentation. Members of this suborder have been found from as long ago as the early Jurassic, not too long after the modern Scleractinia corals originated in the mid-Triassic. Caryophylliines remain a successful group—nearly five hundred species have been described, and they include both shallow-water and deep-sea forms. The majority of caryophylliines do not contain zooxanthellae (symbiotic dinoflagellates) and while there are some colonial forms, the majority are solitary.

Which sounds all very fine and dandy, but is not anywhere near as informative as one might think. It doesn’t take much reading between the lines to note that the ‘Caryophylliina’ with their ‘simple septa’ are essentially united by the absence of the features characterising other suborders. As such, it is hardly surprising that they should be nearly as old as the Scleractinia as a whole, because they are quite possibly phylogenetically equivalent to the Scleractinia as a whole. A caryophylliine is simply a coral that doesn’t put on airs. This possibility is bourne out by molecular analysis (Le Goff-Vitry et al. 2004), which divides scleractinians between two major clades that have been dubbed the ‘robust’ and ‘complex’ clades (or ‘Robusta’ and ‘Complexa’ by Kerr 2005). Though not corresponding to earlier morphological divisions, the two clades are not without morphological support. ‘Robust’ corals have solid, heavily calcified skeletons, while ‘complex’ corals have lighter, more porous skeletons. Caryophylliines, it turns out, are distributed between both clades, and multiple subclades within those clades. Even the type family of the ‘suborder’, the Caryophylliidae, is not monophyletic, with a suggested division between no less than five clades scattered among the Robusta and Complexa (Kerr 2005).

Skeleton of the ‘caryophylliine’ coral Deltocyathus rotulus, showing the fairly plain septa. Photo by Stephen Cairns.

The necessary changes to scleractinian classification could yet be even more radical. Medina et al. (2006) found that the Complexa were more closely related to the soft-bodied, skeletonless Corallimorpharia than they were to the Robusta. It remains an open question whether the calcified skeleton evolved independently in the two clades, or whether the corallimorpharians represent a secondary loss of the skeleton, but my one suspicions lean towards the latter, especially since it has been demonstrated that the loss of the ability to secrete a skeleton does not constitute a death sentence for a coral (Fine & Chernov 2007). Skeletal construction may yet play a role in coral taxonomy, as researchers identify more reliable ultrastructural characters (Stolarski & Roniewicz 2001), but I think we can safely say that the ‘Caryophylliina’ as we have hitherto known it is, well, dead in the water.

Systematics of Scleractinia
<==Scleractinia (see below for synonymy)RB14
    |  |  |    |--StylophyllidaeSR02
    |  |  |    `--Gigantostylis [Gigantostyliidae]SR02
    |  |  |         `--G. epigonus Frech 1890SR02
    |  |  `--+--ThecocyathidaeSR07
    |  |     |    |  i. s.: ‘Microsmilia’ matheyi Koby 1888S00
    |  |     |    `--Thecocyathus tintinabulum Goldfuss 1826SR02
    |  |     `--Anthemiphyllia [Anthemiphylliidae]DFC03
    |  |          |--A. catinata Wells 1907SR02
    |  |          |--A. dentata Alcock 1902SR02
    |  |          |--A. patera De Pourtalès 1878SR02
    |  |          `--A. spiniferaSR02
    |  `--+--VaughanellaDFC03
    |     `--+--FaviinaDFC03
    |        `--+--TurbinoliidaeS00
    |           `--Caryophylliidae [Caryophylliicae]S00
    |                |  i. s.: ‘Thecocyathus’ laevigatus Pourtalès 1871C77
    |                |         ‘Microsmilia’ delemontana (Thurmann 1851)S00
    |                |--Peponocyathus stimpsonii (Pourtalès 1871)C77
    |                |--Anomocora fecunda (Pourtalès 1871)C77
    |                |--Coenosmilia arbuscula Pourtalès 1874C77
    |                |--Asterosmilia prolifera (Pourtalès 1871)C77
    |                |--Coenocyathus cylindricus Milne-Edwards & Haime 1848C77
    |                |--HeterocyathusRSM14
    |                |    |--H. aequicostatus Milne Edwards & Haime 1848RSM14
    |                |    `--H. sulcatus (Verrill 1866)MG-H11
    |                |--CaryophylliaDFC03
    |                |--Tethocyathus microphyllus (Reuss 1871)S00
    |                |--CyathocerasJB12
    |                |--CrispatotrochusDFC03
    |                |--Eusmilia [Eusmiliidae, Eusmiliinae]PR72
    |                |    `--E. fastigiataPR72
    |                |--LopheliaC77
    |                |    |--L. pertusa (Linnaeus 1758)S00
    |                |    `--L. prolifera (Pallas 1766)C77
    |                |--DesmophyllumMG-H11
    |                |    |--D. cristagalliPP64
    |                |    `--D. dianthus (Esper 1794)MG-H11
    |                |--Dasmosmilia Pourtalès 1880C77
    |                |    |--D. lymani (Pourtalès 1871) [=Parasmilia lymani]C77
    |                |    `--D. variegata (Pourtalès 1871)C77
    |                |--TrochocyathusC77
    |                |    |--T. cooperi (Gardiner 1905)MG-H11
    |                |    |--T. planusF91
    |                |    `--T. rawsonii Pourtalès 1874C77
    |                |--StephanocyathusMG-H11
    |                |    |  i. s.: S. coronatus (Pourtalès 1867)MG-H11
    |                |    |--S. (Acinocyathus)MG-H11
    |                |    |    |--S. (A.) explanans (Marenzeller 1904)MG-H11
    |                |    |    `--S. (A.) spiniger (Marenzeller 1888)MG-H11
    |                |    `--S. (Odontocyathus) weberianus (Alcock 1902)MG-H11
    |                `--DeltocyathusC77
    |                     |--D. calcar Pourtalès 1874C77
    |                     |--D. complanatusS98
    |                     |--D. italicus (Michelotti 1838)C77
    |                     |--D. magnificus Moseley 1876MG-H11
    |                     |--D. rotulus (Alcock 1898)MG-H11
    |                     |--D. sarsi (Gardiner & Waugh 1938)MG-H11
    |                     |--D. suluensis Alcock 1902MG-H11
    |                     `--D. violaF71
    `--+--+--Thalamophyllia riiseiDFC03, RB14
       |  `--AgariciodaePR72
       |       |  i. s.: ThamnastraeaQ72 [ThamnastreidaePR72]
       |       |         |--T. arachnoidesH04
       |       |         `--T. seraQ72
       |       |--AgariciidaeDFC03
       |       `--+--SiderastreidaeS99
       |          `--CalamophylliidaeS99
       |               |--Isastrea haueri Cuif 1975NS93
       |               `--Calamophyllia sandbergeri Felix 1891NS93
          `--+--+--+--Leptopsammia pruvotiDFC03, PP64
             |  |  `--TubastraeaDFC03
             |  |       |--T. aureaPR72
             |  |       |--T. coccinea Lesson 1829RSM14
             |  |       |--T. micranthus (Ehrenberg 1834)RSM14
             |  |       `--T. tenuilamellosa (Milne Edwards & Haime 1848)BC01
             |  `--+--PlacotrochusDFC03
             |     |    |--P. deltoideusQ72
             |     |    |--P. elegans Tenison-Woods 1877F71
             |     |    `--P. elongatusQ72
             |     `--PoriticaeS99
             |          |--MicrosolenidaeS99
             |          |    |--MicrosolenaHS02
             |          |    |--ChomatoserisNS93
             |          |    `--GosaviaraeaNS93
             |          `--+--PoritidaeDFC03
             |             `--ActinacididaeS99
             |                  |--ActinacisJB12
             |                  |--AllocoeniopsisJB12
             |                  `--ActinaraeaNS93
                |  |    |--Truncatoguynia Cairns 1989S00
                |  |    |    `--*T. irregularis Cairns 1989S00
                |  |    `--Stenocyathus Pourtalès 1871 [Stenocyathidae]S00
                |  |         |--*S. vermiformis (Pourtalès 1868) [=Coenocyathus vermiformis]S00
                |  |         |--S. alabamensis Wells 1947S00
                |  |         `--S. hoffmeisteri Wells 1977S00
                |  `--FlabellicaeS99
                |       |--GuyniidaeS99
                |       `--Flabellidae [Flabellicae]S00
                |            |--Javania cailleti (Duchassaing & Michelotti 1864)C77
                |            |--Rhizotrochus tuberculatus (Tenison Woods 1879)MG-H11
                |            |--FlabellumDFC03
                |            |--PlacotrochidesS00
                |            |--FalcatoflabellumS00
                |            |--Adkinsella edwardsensis Wells 1933NS93
                |            |--Polymyces Cairns 1979 non Battarra ex Earle 1909 (ICBN)S00
                |            |    `--*P. fragilis (Pourtalès 1871) [=Rhizotrochus fragilis]S00
                |            `--TruncatoflabellumMG-H11
                |                 |--T. aculeatum (Milne Edwards & Haime 1848)RSM14
                |                 |--T. australiensis Cairns 1998MG-H11
                |                 |--T. candeanum (Milne Edwards & Haime 1848)MG-H11
                |                 `--T. macroeschara Cairns 1998MG-H11
                      |    |--R. manuelensis Chevalier 1966C77
                      |    |--R. minutaC-SC03
                      |    `--R. verrilli van der Horst 1922RSM14
                      `--Balanophyllia Wood 1844DFC03, C77
                           |--B. carinata (Semper 1872)MG-H11
                           |--B. cornu Moseley 1881MG-H11
                           |--B. cyathoides (Pourtalès 1871)C77
                           |--B. dentata Tenison Woods 1879MG-H11
                           |--B. desmophyllioides Vaughan 1907MG-H11
                           |--B. floridana Pourtalès 1868 [=B. florideana (l. c.)]C77
                           |--B. italicaPP64
                           `--B. palifera Pourtalès 1878C77
Scleractinia incertae sedis:
  Discocyathus Milne Edwards & Haime 1848SR02
  Schizophyllum fissilis Pourtalès 1874SR02
  Ceratotrochus magnaghiiSR02
  Astraeomorpha Reuss 1854CS04
  Pseudoastraeopora Russo 1979SR01
    `--P. hortensis (Oppenheim 1900)SR01
  Agathelia Reuss 1854SR01
  Stylohelia de Fromentel 1861SR01
  Bracthelia Beauvais & Beauvais 1975SR01
  Palaeohelia albiensis Beauvais 1982SR01
  Etallonasteria minima (Étallon 1864)SR01
  Solenosmilia variabilisMG-H11
  Hoplangia durotrix Gosse 1860S00
  Retiophyllia frechiS00
  Manacina mayoriSC62
  FungiacyathusMG-H11 [FungiacyathidaeDB07]
    |--F. crispus (Pourtalès 1871)C77
    |--F. fragilis Sars 1872MG-H11
    |--F. marenzelleriRB14
    `--F. stephanus (Alcock 1893)MG-H11
  TrachyphylliaRSM14 [TrachyphylliidaeDB07]
    `--T. geoffroyi (Audouin 1826)RSM14
  Anisopsammia rostrataPP64
  Dimorphastrea [Latomeandridae]JB12
    |--Gillastraea delicata Mel’nikova 1983NS93
         |--C. granulata Mel’nikova 1983NS93
         |--C. invurva Mel’nikova 1983NS93
         `--C. tenuiseptata (Mel’nikova 1967)NS93
  Distichophyllia [Distichophyllidae]NS93
  Pamiroseris [Pamiroseriidae]NS93
    |--P. rectilamellosa (Winkler 1861)NS93
    `--P. silesiaca (Beyrich 1852)NS93
    |  i. s.: Aulastraeopora deangelisi Prever 1909SR01
    |         Preverastraea diplothecata (Hackemesser 1936)SR01
    |         Bodeurina languedociensis Beauvais 1980SR01
         |--Acanthogyra micra Eliasova 1973NS93
         |--Kologyra aldingeri Geyer 1955NS93
         |--Pruvostastrea labyrinthiformis Alloiteau 1957NS93
         |--Aplosmilia coalescens Eliášová 1973NS93, SR01
  Bathypsammia tintinnabulum (Pourtalès 1868)C77
  Astroides calycularis (Pallas 1766)RL04
  Heteropsammia cochlea (Spengler 1781)MG-H11
    |--D. cornigeraPP64
    `--D. manniB79
  Duncanopsammia axifuga (Milne Edwards & Haime 1848)RSM14

Scleractinia [Agariciicae, Caryophyllida, Caryophylliina, Caryophyllioidea, Dendrophyllida, Dendrophylliidae, Dendrophylliina, Fungiida, Fungiina, Guynioidea, Madreporaria]RB14

*Type species of generic name indicated


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[B79] Buss, L. W. 1979. Habitat selection, directional growth and spatial refuges: why colonial animals have more hiding places. In: Larwood, G., & B. R. Rosen (eds) Biology and Systematics of Colonial Organisms pp. 459–497. Academic Press: London.

[C77] Cairns, S. D. 1977. Stony corals. I. Caryophylliina and Dendrophylliina (Anthozoa: Scleractinia). Memoirs of the Hourglass Cruises 3 (4): 1–27.

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[DB07] Daly, M., M. R. Brugler, P. Cartwright, A. G. Collins, M. N. Dawson, D. G. Fautin, S. C. France, C. S. McFadden, D. M. Opresko, E. Rodriguez, S. L. Romano & J. L. Stake. 2007. The phylum Cnidaria: a review of phylogenetic patterns and diversity 300 years after Linnaeus. Zootaxa 1668: 127–182.

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

[F91] Freess, W. B. 1991. Beiträge zur Kenntnis von Fauna und Flora des marinen Mitteloligozäns bei Leipzig. Altenburger Naturwissenschaftliche Forschungen 6: 3–74.

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

[HS02] Höfling, R., & R. W. Scott. 2002. Early and Mid-Cretaceous buildups. SEPM Special Publication 72: 521–548.

[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.

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[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

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[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.

[PP64] Peres, J. M., & J. Picard. 1964. Nouveau manuel de bionomie benthique de la mer Mediterranee. Recueil des Travaux de la Station Marine d’Endoume, Bulletin 31 (27): 5–137.

[PR72] Powers, D. A., & F. J. Rohlf. 1972. A numerical taxonomic study of Caribbean and Hawaiian reef corals. Systematic Zoology 21 (1): 53–64.

[Q72] Quilty, P. G. 1972. The biostratigraphy of the Tasmanian marine Tertiary. Papers and Proceedings of the Royal Society of Tasmania 106: 25–44.

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

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