Polyaxone monaxonids: revision of raspailiid sponges with polyactine megascleres (Cyamon and Trikentrion)

Abstract

Among the thousands of non-tetractinellid (monaxonid) Demospongiae species, less than twenty possess polyactine (usually three- or four-claded) megascleres. These are currently assigned to two closely related genera, viz. Cyamon Gray and Trikentrion Ehlers, both members of the raspailiid subfamily Cyamoninae. The two genera are considered valid on account of differences in the shape and the ornamentation of the polyaxone spicules. Cyamon predominantly has four-claded equiangular spicules with all cladi spined or rugose, whereas Trikentrion usually has a majority of three-claded spicules on which spines are found only on a single basal clade. Nevertheless, the differences between the two genera appear to overlap in several known and newly discovered species, necessitating a revision of the two groups. Two new species of Cyamon were found to occur on inshore sandstone platforms off the coast of Mauritania. One of the new species, Cyamon amphipolyactinum sp. n., possesses unique small 'double' polyactine spicules in addition to the usual calthrops-like polyactine megascleres characteristic for Cyamon. The second new species, Cyamon arguinense sp. n., possesses polyactine megascleres of which only one of the cladi is spined the remaining three or more cladi being smooth, a feature that is considered characteristic of sponges of the genus Trikentrion. The type species of Cyamon, C. vickersii (Bowerbank) appears to have been misinterpreted as a Caribbean species, because circumstantial evidence strongly indicates an Indian Ocean origin. This has the consequence that specimens recorded subsequently under the name C. vickersii from various Western Atlantic localities are reassigned to Cyamon agnani (Boury-Esnault), a species originally described from Brazil. A new species, reported as Cyamon vickersii sensu Burton & Rao from the east coast of India, and available to us only as a single thick section mounted on a glass slide, is named Cyamon hamatum sp. n. The Cyamon membership of the only deep-sea species, Cyamon spinispinosum (Topsent) is drawn in doubt due to considerable morphological deviation from mainstream Cyamon. The type species of Trikentrion, T. muricatum (Pallas), is extensively described and discussed, and a neotype is assigned. West African Trikentrion laeve (Carter) is for the first time since its original description properly redescribed from the type material. The specimen recorded by Burton as Trikentrion laeve from Congo turned out to be different from the original material of Carter and is assigned to a new species, Trikentrion africanum sp. n. All species of both genera considered valid are reviewed, mostly based on the examination of type or other original specimens. Our revision shows the existence of twelve species of Cyamon and six species of Trikentrion. A key to the species is provided and remarks on the geographic distribution of both genera are made. Based on our study, the differences between Cyamon and Trikentrion are re-evaluated. Only one character absolutely distinguishes the two genera, the presence (Trikentrion) or absence (Cyamon) of trichodragmata. A further discriminating character is the possession of short thick styles (most Cyamon species) versus thick oxeas (many Trikentrion), but this is complicated by absence of the oxeas in three Trikentrion species. Although spination of the polyactine spicules in itself cannot serve to distinguish the two genera with certainty, those of Trikentrion are usually recognizable by excessive hook-like spines against a finer spination in Cyamon. Possibly, the polyactine spicules of both groups are non-homologous, with Cyamon polyactines derived from styles and Trikentrion polyactines from oxeas, but this remains to be further investigated.