The caryophylliines are known from the early Jurassic (180 million years ago) to the Recent, and occur in most marine environments to a depth of 3200 m. They are an extremely diverse group, consisting of 91 living genera and 457 living species (Cairns et al., 1999). Although some species form large colonies that may contribute to both shallow-water and deep-water reef/bank structure, most members of this suborder are small (less than 30 mm) and inconspicuous, occurring in cryptic shallow-water environments or in cold (as low as -1°C), dark, deep waters. Although several other scleractinian families occur in deep water (e.g., Dendrophylliidae, Micrabaciidae, Fungiacyathidae), the caryophylliines have been the most successful in exploiting the deep-sea realm. Because many occur below the euphotic zone or cryptically in shallow water, most species are azooxanthellate (do not contain symbiotic unicellular dinoflagellates) and therefore rarely attain a large size.
The higher classification of the suborder Caryophylliina is in a state of flux as more information is being discovered through microstructural and molecular analyses. Stolarski (1996, 2000) has best summarized the conflicting morphological characters, which resulted in two fairly different classification hypotheses for the superfamilies and families of the caryophylliines, which are both different from the classification presented here. In general, we agree with he work of Stolarski (2000), and, in general, adopt his "Hypothesis A", but with some modifications that reflect a more traditional view. Thus, we continue to recognize the superfamily Flabelloidea and place the Guyniidae in that superfamily, but we do accept his new family Stenocyathidae, which he places in the superfamily Caryophyllioidea, and his new family Schizocyathidae, which he places in the superfamily Volzeioidea.
Suborders within the Scleractinia are distinguished by the structure of their septa. The septa of the caryophylliines are lamellar, composed of one fan system of simple, very small trabeculae (minitrabeculate), resulting in a smooth or nearly smooth inner margin (Vaughan and Wells, 1943; Wells, 1956).
The superfamilies within this suborder are characterized by the structure of their walls (theca). The most primitive of the three superfamilies, the Volzeioidea, have an exclusively epithecate wall (see Stolarski, 1995, 1996); that of the Flabelloidea is marginothecate; and that of the Caryophyllioidea is septo- or parathecate. In the latter two cases a nontrabecular deposit of epitheca or textura may be added to the exterior of the corallum (Stolarski, 1995).
Evolution and Systematics
Discussion of Phylogenetic Relationships
- Volzeioidea †
No phylogenetic analysis has been performed on the suprageneric taxa of the Caryophylliina; however, Stolarski (1995, 1996), Romano and Palumbi (1996), and Romano and Cairns (2000) are currently investigating and reassessing the higher level relationships in the suborder by using characteristics of the corallum microstructure and DNA sequencing, respectively (see Scleractinia branch page). Roniewicz and Morycowa (1993) and Veron (1995: 110) should also be consulted for more recent, non-cladistic evolutionary trees of all scleractinian families.
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