Colpophyllia, MILNE EDWARDS & HAIME, 1848: 492

Budd, Ann F., Fukami, Hironobu, Smith, Nathan D. & Knowlton, Nancy, 2012, Taxonomic classification of the reef coral family Mussidae (Cnidaria: Anthozoa: Scleractinia), Zoological Journal of the Linnean Society 166 (3), pp. 465-529 : 518-520

GENUS COLPOPHYLLIA MILNE EDWARDS & HAIME, 1848: 492 ( FIGS 10E, F , 15E–J, 21A–C , 25A–C )

Synonyms: None.

Type species: Meandrina gyrosa de Lamarck, 1816: 247 [= Madrepora natans Houttuyn, 1772: 124 ]; by original designation. The holotype MNHN-scle105 is lost (A. Andouche, pers. comm., 2009). Also lost are (1) Milne Edwards & Haime’s specimen of ‘ Colpophyllia fragilis Dana’, which was described and illustrated by Matthai (1928: 103–104) (A. Andouche, pers. comm. 2009); and (2) Esper’s (1795) figured specimen of Madrepora natans , which Matthai (1928: 102, pl. 67, figs 1, 2) designated as the ‘type’ of Colpophyllia natans ( Müller, 1775) ( Scheer, 1990) . We herein designate specimen SUI 130588 (Carlon #828), collected by D. B. Carlon in 2007 at Crawl Cay, Bocas del Toro, Panama, as the neotype of Colpophyllia natans ( Houttuyn, 1772) ( Fig. 10E, F ).

Original type species locality: Unknown [Recent].

Original description: ‘Polypier composé, massif. Les séries de polypiérites étant intimement soudées entre elles par les côtes. Columelle rudimentaire ou nulle. Cloisons serrées, excessivement minces, à bord trèsfinement deniculé et faiblement échancré dans son milieu, de manière à simuler inférieurement un lobe peu marqué. Endothèque vésiculaire très-abondante’ ( Milne Edwards & Haime, 1848: 492).

Subsequent morphological descriptions ( Matthai, 1928 and later): Matthai (1928: 99–109); Vaughan & Wells (1943: 171); Wells (1956: F403); Walton Smith (1971: 82–83); Zlatarski & Estalella (1982: 83–85); Veron (2000: vol. 3: 210–211).

Diagnosis: Macromorphology: meandroid (uniserial), with large valleys (> 15 mm); limited costate coenosteum forming a distinctive ‘double wall’; discontinuous, compact trabecular columella with lamellar linkage between centres; reduced epitheca; abundant vesicular endotheca; septal lobes ( Figs 10E–F , 15E–J).

Micromorphology: regular fan-shaped septal teeth with elliptical bases orientated transverse to the septal plane; spiky septal granules organized in lines; interarea of teeth along a septum is distinctively horizontally layered (banded); teeth in different septal cycles and along individual septa similar in size and shape ( Fig. 21A–C ).

Microstructure: parathecal corallite wall; centres of calcification within costosepta and columella form a well-developed medial line crossed by carinae; limited thickening deposits ( Fig. 25A–C ).

Species included: Colpophyllia natans ( Houttuyn, 1772) [holotype is from an unknown locality, and is lost; neotype (herein designated) = SUI130588 ( Carlon #828), Fig. 10E, F , Bocos del Toro, Panama] .

Colpophyllia amaranthus ( Houttuyn, 1772) [holotype is from an unknown locality, and is lost ( Matthai, 1928); neotype (herein designated) = USNM100498 , Fig. 15G, H, Venezuela] .

Colpohyllia breviserialis Milne Edwards & Haime, 1849 [holotype = BM (NH)18.40.5.29.6, Fig. 15I, unknown locality] .

Remarks: As explained by Wells (1936), there are two different species that have been assigned the name ‘gyrosa’: (1) Madrepora natans Houttuyn, 1772 = Madrepora gyrosa Ellis & Solander, 1786 = Meandrina gyrosa de Lamarck, 1816 = Colpophyllia natans Matthai, 1928 ; (2) Manicina gyrosa Ehrenberg, 1834 = Manicina gyrosa Matthai, 1928 = Manicina mayori Wells, 1936 . The first is the type species of Colpophyllia , and its type specimen is lost. A neotype (SUI130588) has been therefore designated for Colpophyllia natans . Wells (1936: 105) proposed the name Manicina mayori to replace the second species, Manicina gyrosa Ehrenberg , because Ellis & Solander (1786) had already named the first species Madrepora gyrosa .

In the classification system of Vaughan & Wells (1943) and Wells (1956), the genus Colpophyllia is distinguished by having a meandroid colony form; a ‘double’ wall; small septal lobes; discontinuous series and collines; centres linked by lamellae; a spongy, parietal (= trabecular) columella. Septal margins are strongly dentate; trabeculae are usually simple, in one or two fan systems ( Vaughan & Wells, 1943: 153, 154, 163, 171). Our observations agree. In addition, the genus has a distinctive parathecal wall (dissepimental), and centres of calcification in the costosepta and columella form well-defined medial lines that are crossed by distinct carinae (or short transverse lines). Septal teeth are small and fan-shaped, orientated transverse to the plane of the septum. The genus Colpophyllia is distinguished from Mussismilia by its meandroid colony form, lamellar centre linkage, septal lobes, and smaller septal teeth. It is distinguished from Favia , Diploria , and Manicina by its lamellar linkage, reduced epitheca, abundant vesicular endotheca, and parathecal wall. Colpophyllia is distinguished from its meandroid Indo-Pacific counterpart, Oulophyllia Milne Edwards & Haime, 1848, on the basis of its double wall, lamellar linkage, and septal lobes; its parathecal walls (trabeculothecal in Oulophyllia ), its regular tricorne septal teeth with elliptical-perpendicular bases (irregular multiaxial in Oulophyllia ); and its strong, aligned granules.

Cairns, Hoeksema & Land (1999) recognized three species of Colpophyllia ( Colpophyllia natans , Colpophyllia amaranthus , Colpophyllia breviserialis ), which differ in valley length, depth, and numbers of septa per cm. Colpophyllia amaranthus ( Fig. 15G, H) is characterized by deep discontinuous valleys (up to 30 mm) and more numerous septa (ten to 12 centres per cm). Colpophyllia breviserialis ( Fig. 15I, J) is characterized by short valleys having fewer than five centres. Colpophyllia natans ( Figs 10E, F , 15E, F) is characterized by longer and more continuous valleys, a distinctive double wall, and eight to nine septa per cm. Both Zlatarski & Estalella (1982) and Veron (2000) synonymized the three species; Walton Smith (1971) recognized only Colpophyllia natans and Colpophyllia amaranthus .

Figures

Figure 10. Type specimens of type species of genera in the subfamily Faviinae. A–D, genus Favia Milne Edwards & Haime, 1857; Madrepora fragum Esper, 1795; neotype (C, D, designated herein) = MNHN-scle560, Haiti. E, F, genus Colpophyllia Milne Edwards & Haime, 1848; Meandrina gyrosa de Lamarck, 1816 [= Madrepora natans Houttuyn, 1772]; neotype (designated herein) = SUI130588 (Carlon #828), Crawl Cay, Bocas del Toro, Panama. G, H, genus Diploria Milne Edwards & Haime, 1848; Meandrina cerebriformis de Lamarck, 1816; holotype = MNHN-scle102, unknown locality.

Figure 21. Colpophyllia, Manicina, and Mussismilia micromorphology (scanning electron microscopy): left column, wall; middle column, mid-septum; right column, columella. Colpophyllia, Manicina, and Mussismilia have paddle-shaped to tricorne teeth with elliptical bases orientated perpendicular to the septal plane. Tooth height is low (<0.3 mm) in Colpophyllia and Manicina, and medium to high (> 0.3 mm) in Mussismilia. The interarea of teeth is smooth in Manicina but horizontally banded in Colpophyllia and Mussismilia. Granules are spiked and aligned. A–C, Colpophyllia natans (Houttuyn, 1772); figured specimens = SUI122804 (FA1100) Discovery Bay, Jamaica (A); SUI122802 (FA1071), Bocas del Toro, Panama (B, C). D–F, Manicina areolata (Linnaeus, 1758); figured specimen = SUI122824 (FA1107), Bocas del Toro, Panama. G–I, Mussismilia hartti (Verrill, 1868); figured specimen = YPM4516, Maria Farinha, Pernambuco, Brazil. J–L, Mussismilia braziliensis (Verrill, 1868); figured specimen = YPM9104, Santa Barbara Island, Abrolhos Archipelago, Bahia, Brazil. M–O, Mussismilia leptophylla (Verrill, 1868); figured specimen = SUI99645 (FA1029), Abrolhos Reef, Abrolhos Archipelago, Bahia, Brazil (M); YPM9087, Lixa Reef, Abrolhos Archipelago, Bahia, Brazil. (N, O).

Figure 25. Colpophyllia, Manicina, and Mussismilia microstructure (transverse thin section): left column, wall; middle column, corallite interior; right column, close-up of septa. Corallite walls (w) in Colpophyllia and Mussismilia are parathecal; whereas in Manicina they are septothecal with trabeculothecal elements. Trabeculothecal elements are also present in Mussismilia braziliensis and to a lesser extent in Mussismilia leptophylla. Moderately well-defined costoseptal medial lines (m) and well-developed carinae (cr) occur in all three genera. A–C, Colpophyllia natans (Houttuyn, 1772); figured specimen = SUI122802 (FA (FA1071), Bocas del Toro, Panama (A); SUI122804 (FA1100, J115), Discovery Bay, Jamaica (B, C). D–F, Manicina areolata (Linnaeus, 1758); figured specimen = SUI122822 (FA1067), Bocas del Toro, Panama. G–I, Mussismilia hartti (Verrill, 1868); figured specimen = YPM4516, Maria Farinha, Pernambuco, Brazil. J–L, Mussismilia braziliensis (Verrill, 1868); figured specimen = YPM9104, Santa Barbara Island, Abrolhos Archipelago, Bahia, Brazil. M–O, Mussismilia leptophylla (Verrill, 1868); holotype = YPM1517A, Abrolhos Reef, Abrolhos Archipelago, Bahia, Brazil.