Trachyphyllia, MILNE EDWARDS & HAIME, 1848

Huang, Danwei, Benzoni, Francesca, Fukami, Hironobu, Knowlton, Nancy, Smith, Nathan D. & Budd, Ann F., 2014, Taxonomic classification of the reef coral families Merulinidae, Montastraeidae, and Diploastraeidae (Cnidaria: Anthozoa: Scleractinia), Zoological Journal of the Linnean Society 171 (2), pp. 277-355 : 340-342

GENUS TRACHYPHYLLIA MILNE EDWARDS & HAIME, 1848 A: 492 ( FIG. 26 )

Synonym

Wellsophyllia Pichon, 1980: 255 (type species: Wellsophyllia radiata Pichon, 1980: 257 , figs 1–4 = Callogyra formosa Bedot, 1907: 176 , pl. 15: figs 63–69, non Callogyra formosa Verrill, 1901: 86 , pl. 24: figs 1, 2; see Best & Hoeksema, 1987; original designation, Pichon, 1980: 255).

Type species

Manicina amarantum Dana, 1846: 189 , pl. 9: fig. 1 = Turbinolia geoffroyi Audouin, 1826: 233 , pl. 4: figs 1.1, 1.2; subsequent designation, Milne Edwards & Haime, 1849a, vol. 11: 275.

Original description

‘Diffère surtout du précédent [ Colpophyllia ] en ce que les séries restent libres par les côtés, que la columelle est spongieuse et bien marquée et que les cloisons présentent un lobe paliforme bien distinct.’ ( Milne Edwards & Haime, 1848a, vol. 27: 492).

Subsequent descriptions

Milne Edwards & Haime, 1849a, vol. 11: 275; Milne Edwards & Haime, 1857, vol. 2: 340, 341; Quenstedt, 1881: 1006; Duncan, 1884: 82; Quelch, 1886: 77; Saville Kent, 1893: 161; Delage & Hérouard, 1901: 626; Faustino, 1927: 146, 147; Matthai, 1928: 95; Yabe et al., 1936: 21; Vaughan & Wells, 1943: 170, 171; Alloiteau, 1952: 618; Wells, 1956: F407; Nemenzo, 1959: 105; Chevalier, 1975: 201; Veron et al., 1977: 207; Scheer & Pillai, 1983: 139; Wood, 1983: 174; Veron, 1986: 538; Chevalier & Beauvais, 1987: 719; Veron, 2000, vol. 3: 272.

Diagnosis (apomorphies in italics)

Colonial and free-living, with intracalicular budding only. Corallites monomorphic and uniserial; monticules absent. Phaceloid (flabello-meandroid). Calice width large (> 15 mm), with high relief (> 6 mm). Septa in ≥ four cycles (≥ 48 septa). Free septa present but irregular. Septa spaced <six septa per 5 mm. Costosepta unequal in relative thickness. Columellae trabecular and spongy (> three threads), <1/4 of calice width, and continu- ous amongst adjacent corallites. Septal (multiaxial) lobes well developed. Epitheca well developed and endotheca low−moderate (tabular) ( Fig. 26A, D ).

Tooth base at midcalice circular. Tooth tip at midcalice irregular; tip orientation perpendicular to septum. Tooth height low (<0.3 mm) and tooth spacing narrow (<0.3 mm), with> six teeth per septum. Granules aligned on septal face, perpendicular to septal margin; irregular in shape. Interarea palisade ( Fig. 26B, E ).

Walls formed by dominant paratheca and partial septotheca; trabeculothecal elements may be present; abortive septa absent. Thickening deposits fibrous. Costa centre clusters weak; 0.3–0.6 mm between clusters; medial lines strong. Septum centre clusters weak; 0.3– 0.5 mm between clusters; medial lines strong. Transverse crosses present. Columella centres clustered ( Fig. 26C, F ).

Species included

Trachyphyllia geoffroyi ( Audouin, 1826: 233, pl. 4: figs 1.1, 1.2); syntypes of Manicina amarantum: USNM 85, YPM IZ 1974 (two dry specimens; Fig. 26A ); type locality of Manicina amarantum : Singapore ( Verrill, 1864: 48); phylogenetic data: molecular and morphology.

Taxonomic remarks

Trachyphyllia was established by Milne Edwards & Haime (1848a, vol. 27: 492) initially without a type, and compared with the genus Colpophyllia , a meandroid Atlantic genus. Manicina amarantum Dana, 1846: 189 , pl. 9: fig. 1, was designated the type species shortly after, but this name had been used earlier on an Atlantic species Colpophyllia amaranthus ( Houttuyn, 1772: 128) ( Verrill, 1901: 81; Matthai, 1914: 97). The next available name that could be used was the second Trachyphyllia species studied by Milne Edwards & Haime (1849a, vol. 11: 276), Trachyphyllia geoffroyi . This incidentally was a young coral of Dana’s species collected from the Red Sea, figured in Audouin (1826: 233, pl. 4: fig. 1.1).

Trachyphyllia remains a monotypic genus, phylogenetically recovered unexpectedly in a clade along with Dipsastraea and Coelastrea . It may be nested amongst these genera ( Huang et al., 2011; Arrigoni et al., 2012), or as an outgroup to them ( Fukami et al., 2008). Regardless, the long branch subtending it suggests that it is genetically very distinct, and we maintain its present generic status until more samples have been analysed.

Trachyphyllia is widely distributed on reefs of the Indo-Pacific, and absent east of Fiji.

Morphological remarks

Many apomorphies define Trachyphyllia , and even more so on the molecular tree simply because it is separat- ed from Coelastrea , to which it is morphologically closest. Based on an integrated analysis of both data types, eight apomorphies of macro- and micromorphology are identified (see Diagnosis above), distinguishing this genus from Coelastrea , which in contrast has discrete corallites of medium width (4–15 mm) and relief (3–6 mm), limited or fused walls, evenly thick costosepta, medium tooth height (0.3–0.6 mm) and spacing (0.3– 1 mm), and scattered granules.

Trachyphyllia is the only free-living coral in Merulinidae as defined here, noting that Catalaphyllia , possibly also a merulinid (see remarks for Merulinidae above; Romano & Cairns, 2000; Barbeitos et al., 2010; Huang, 2012; Huang & Roy, 2013), can also be freeliving ( Wells, 1971; Veron et al., 1977). This represents an autapomorphy that is not phylogenetically informative within the family, but which is Trachyphyllia ’s most distinctive feature.

FAMILY MONTASTRAEIDAE YABE & SUGIYAMA, 1941: 72

Synonym

Montastreinae Vaughan & Wells, 1943: 171 (misspelling).

Figures

Figure 26. Trachyphyllia Milne Edwards & Haime, 1848a, is flabello-meandroid and free-living, with large (> 15 mm) and high-relief (> 6 mm) calices, septa in ≥ four cycles (≥ 48 septa) spaced <six septa per 5 mm, and well-developed septal (multiaxial) lobes. Septal teeth are low (<0.3 mm) and narrowly spaced (<0.3 mm); irregularly shaped granules aligned on septal face. Walls formed by dominant paratheca and partial septotheca, with strong costal and septal medial lines. A–F, Trachyphyllia geoffroyi (Audouin, 1826), the type and only living species of Trachyphyllia; macromorphology, syntype YPM 1974, unknown locality (A), and hypotype USNM 75192, the Philippines (D); micromorphology (scanning electron microscopy), hypotype USNM 91340, Negri Sembilan, western coast of Peninsula Malaysia (B, E); microstructure (transverse thin section), hypotype UF 1383 (FA1073), Palau (C, F).