Cyphastrea, 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 : 304-307

GENUS CYPHASTREA MILNE EDWARDS & HAIME, 1848 A: 494 ( FIG. 9 )

Type species

Astrea microphthalma Lamarck, 1816: 261 ; original designation, Milne Edwards & Haime, 1848a, vol. 27: 494.

Original description

‘Diffère des trois genres précédents [ Astrea , Plesiastrea , Solenastrea ] par la compacité du coenenchyme et par

la structure poutrellaire de la partie interne des cloisons.’ ( Milne Edwards & Haime, 1848a, vol. 27: 494).

Subsequent descriptions

Milne Edwards & Haime, 1849b, vol. 12: 114; Milne Edwards & Haime, 1857, vol. 2: 484–485; Tenison-Woods, 1878: 322; Klunzinger, 1879: 50–51; Duncan, 1884: 107; Quelch, 1886: 106; Saville Kent, 1893: 27; Gardiner, 1899: 761; Delage & Hérouard, 1901: 630; Gardiner, 1904: 778; Matthai, 1914: 38–39; Vaughan, 1918: 87; Hoffmeister, 1925: 19; Faustino, 1927: 114–115; Yabe et al., 1936: 23; Vaughan & Wells, 1943: 174; Alloiteau, 1952: 625; Crossland, 1952: 117; Wells, 1956: F406; Nemenzo, 1959: 112; Chevalier, 1975: 9; Veron et al., 1977: 167; Wijsman-Best, 1980: 239; Scheer & Pillai, 1983: 133; Wood, 1983: 167, 170; Veron, 1986: 520; Chevalier & Beauvais, 1987: 716; Sheppard, 1990: 12; Sheppard & Sheppard, 1991: 139; Veron, 2000, vol. 3: 240.

Diagnosis (apomorphies in italics)

Colonial, with extracalicular budding only. Corallites monomorphic and discrete (one to three centres); monticules absent. Coenosteum generally spinose (costate in Cyphastrea agassizi and apical corallites of Cyphastrea decadia ), moderate amount (<corallite diameter; extensive in Cyphastrea decadia ). Calice width small (<4 mm), with low relief (<3 mm). Costosepta not confluent. Septa in ≤ three cycles (≤ 36 septa). Free septa regular. Septa spaced> 11 septa per 5 mm. Costosepta unequal in relative thickness. Columellae trabecular but compact (one to three threads), <1/4 of calice width, and discontinuous amongst adjacent corallites. Paliform (uniaxial) lobes weak or moderate. Epitheca well developed and endotheca lowmoderate (tabular) ( Fig. 9A, D, G ).

Tooth base at midcalice circular. Tooth tip at midcalice irregular; tip orientation multiaxial. Tooth height low (<0.3 mm) and tooth spacing narrow (<0.3 mm), with> six teeth per septum. Granules scattered on septal face; strong (pointed). Interarea smooth ( Fig. 9B, E, H ).

Walls formed by dominant septotheca; abortive septa absent. Thickening deposits fibrous. Costa centre clusters weak; 0.3–0.6 mm between clusters; medial lines weak. Septum centre clusters weak; <0.3 mm between clusters; medial lines weak. Transverse crosses absent. Columella centres clustered ( Fig. 9C, F, I ).

Species included

1. Cyphastrea microphthalma ( Lamarck, 1816: 261) ; holotype: MNHN IK-2012–14002 (dry specimen; Fig. 9A ); type locality: ‘les mers de la Nouvelle- Hollande’ ( Lamarck, 1816: 261); phylogenetic data: molecular and morphology.

2. Cyphastrea agassizi ( Vaughan, 1907: 101, pl. 25: figs 2, 2a, 3, 3a); syntypes: USNM 21633 , 21634 (two dry specimens); type locality: O’ahu, Hawai’i; phylogenetic data: partial morphology .

3. Cyphastrea chalcidicum ( Forskål, 1775: 136) ; holotype: lost ( Matthai, 1914: 42); neotype: NHMUK 1978.1.1.4 ( Wijsman-Best, 1980: 242), designated by Veron et al. (1977: 173), status unknown; type locality: southwest Swain Reefs, Australia, 5–14 m in depth; phylogenetic data: molecular and morphology.

4. Cyphastrea decadia Moll & Best, 1984: 56 , figs 5, 6; holotype: RMNH 15271 (dry specimen); paratypes: RMNH 15272 , 15273 (two dry specimens); type locality: 111 m offshore of north Pajenekang , Spermonde Archipelago, Indonesia, 8 m depth; phylogenetic data: partial morphology .

5. Cyphastrea hexasepta Veron, Turak & DeVantier, 2000 ( Veron, 2000, vol. 3: 245, fig. 5; see also Veron, 2002: 171, figs 312–314; ICZN, 2011: 163); lectotype (designated herein): MTQ G55834 (dry specimen); type locality: northern Red Sea coast of Saudi Arabia, 10 m depth; phylogenetic data: none.

6. Cyphastrea japonica Yabe & Sugiyama, 1932: 161 (see also Yabe et al., 1936: 25, pl. 17: figs 4–6); holotype: TIU 40323 (dry specimen); type locality: Misaki, Shikoku, Japan; phylogenetic data: molecular only ( Chen et al., 2004).

7. Cyphastrea ocellina ( Dana, 1846: 218, plate 10: fig. 10); syntypes: YPM IZ 474, 4330 (two dry specimens); type locality: Hawai’i; phylogenetic data: molecular only ( Romano & Palumbi, 1996).

8. Cyphastrea serailia ( Forskål, 1775: 135) ; syntypes: ZMUC ANT-000367 ( Fig. 9G ) to ANT-000373, figured in Matthai (1914, pl. 11: figs 4–9; seven dry specimens); type locality: Red Sea; phylogenetic data: molecular and morphology.

Taxonomic remarks

Cyphastrea Milne Edwards & Haime, 1848a , vol. 27: 494, was established to accommodate species distinguished by their compact coenosteum – ‘compacité du coenenchyme’ ( Milne Edwards & Haime, 1848a, vol. 27: 494). Following which, only one species – Cyphastrea agassizi – has ever been placed in another genus, implying limited confusion with its taxonomy.

Molecular data indicate that Cyphastrea is very dissimilar from other taxa as it is subtended by a long branch from its sister genus, Orbicella . Yet, the Cyphastrea + Orbicella clade (subclade C) is a wellsupported relationship that has been recovered in several studies ( Fukami et al., 2004 a, 2008; Huang et al., 2011; Arrigoni et al., 2012).

Cyphastrea is widely distributed on reefs of the Indo- Pacific, present in French Polynesia and the Pitcairn Islands in the Southern Hemisphere ( Glynn et al., 2007), but absent in the eastern Pacific in the north.

Morphological remarks

Cyphastrea is a morphologically well-defined and moderately supported genus (bootstrap support of 69 and decay index of 2), but it is also exclusively associated with Echinopora , Orbicella , and Paramontastraea . Cyphastrea spp. share the plesiomorphic state of spinose coenosteum with Echinopora and Paramontastraea , amongst other characters with Orbicella , supporting them as a clade that is sister to the rest of Merulinidae .

Despite the recent emphasis that corals east and west of the Americas are genetically distinct from one another ( Fukami et al., 2004a), and whilst Cyphastrea and Orbicella are found solely in the Indo-Pacific and Atlantic realms, respectively, synapomorphies are present for the clade comprising them, namely small calice width (likelihood of 0.69 based on the Mk1 model) and trabecular but compact columellae (likelihood 0.86). They also have walls formed predominantly by septotheca, a plesiomorphic state shared only with Paramontastraea .

On its own, Cyphastrea is defined by the synapomorphy of strong pointed granules on the septal face (likelihood 0.99). Because its closest relative does not overlap geographically, it is easily identified with the apomorphies shared with Orbicella . The multiaxial tooth tips, although also present amongst Echinopora and Paramontastraea , are much more conspicuous in Cyphastrea because of their small corallites.

To date, phylogenetic data are only available for about half of the members of Cyphastrea (see also Romano & Palumbi, 1996; Chen et al., 2004).

Figures

Figure 9. Cyphastrea Milne Edwards & Haime, 1848a has discrete corallites that bud extracalicularly, small (<4 mm) and low-relief (<3 mm) calices, regular free septa, and compact columellae. Septal teeth are low (<0.3 mm) and narrowly spaced (<0.3 mm), with multiaxial tips; strong (pointed) granules scattered on septal face. Walls formed by dominant septotheca. A–C, Cyphastrea microphthalma (Lamarck, 1816), type species of Cyphastrea; macromorphology, holotype MNHN IK-2012–14002, unknown locality (A); micromorphology (scanning electron microscopy; B) and microstructure (transverse thin section; C), hypotype RMNH 15111 (FA1035), southwest Sulawesi, Indonesia. D–F, Cyphastrea chalcidicum (Forskål, 1775); macromorphology, hypotype MTQ G61902, Orpheus Island, Australia (D); micromorphology (E) and microstructure (F), hypotype RMNH 12787, Gulf of Aqaba, Israel. G–I, Cyphastrea serailia (Forskål, 1775); macromorphology, syntype ZMUC ANT-000367, Red Sea (G; photo by M. A. Krag); micromorphology (H) and microstructure (I), hypotype RMNH 15106, southwest Sulawesi, Indonesia.