Dipsastraea, DE BLAINVILLE, 1830: 338

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 : 307-313

GENUS DIPSASTRAEA DE BLAINVILLE, 1830: 338 ( FIG. 10 )

Synonyms

Barabattoia Yabe & Sugiyama, 1941: 72 (type species: Barabattoia mirabilis Yabe & Sugiyama, 1941: 72 , pl. 61: figs 1, 1a–e; original designation, Yabe & Sugiyama, 1941: 72); Bikiniastrea Wells, 1954: 456 (type species: Bikiniastr[e]a laddi Wells 1954: 456 , pl. 172; original designation, Wells, 1954: 456).

Type species

Madrepora favus Forskål, 1775: 132 ; subsequent designation, Wells, 1936: 109.

Original description

‘Plus ou moins globuleuses, formées de loges profondes, infundibuliformes, subpolygonales, à parois communes, à bords élevés, multisillonnés et échinulés.’ ( de Blainville, 1830: 338).

Subsequent descriptions

de Blainville, 1834: 373; Wells, 1936: 109.

Diagnosis

Colonial, with intracalicular budding only. Corallites monomorphic and discrete (one to three centres); monticules absent. Coenosteum costate, moderate amount (<corallite diameter), limited (includes double wall) in some species. Generally, calice width medium (4–15 mm), with medium relief (3–6 mm); few species with wider and/or deeper calice. Costosepta not confluent. Septa in three cycles (24–36 septa). Free septa present but generally irregular (regular in Dipsastraea helianthoides and Dipsastraea laxa ). Septa spaced six to 11 septa per 5 mm. Costosepta equal in relative thickness. Columellae trabecular and spongy (> three threads), <1/4 of calice width, and continuous amongst adjacent corallites. Paliform (uniaxial) lobes weak or moderate. Epitheca well developed and endotheca low−moderate (tabular) ( Fig. 10A, D, G, J ).

Tooth base at midcalice circular. Tooth tip at midcalice irregular; tip orientation perpendicular to septum. Tooth height medium (0.3–0.6 mm) and tooth spacing medium (0.3–1 mm), with> six teeth per septum. Granules scattered on septal face; irregular in shape. Interarea palisade ( Fig. 10B, E, H, K ).

Walls formed by dominant paratheca and partial septotheca; abortive septa absent. Thickening deposits fibrous. Costa centre clusters generally strong but highly variable; 0.3–0.6 mm between clusters; medial lines weak. Septum centre clusters weak; 0.3–0.5 mm between clusters; medial lines weak. Transverse crosses present. Columella centres clustered ( Fig. 10C, F, I, L ).

Species included

1. Dipsastraea favus ( Forskål, 1775: 132) ; lectotype: ZMUC ANT-000466 (dry specimen; Fig. 10A ); type locality: Red Sea; phylogenetic data: molecular and morphology .

2. Dipsastraea albida ( Veron, 2000, vol. 3: 112, figs 1, 2) (see also Veron, 2002: 140, figs 257–259; ICZN, 2011: 164); lectotype (designated herein): MTQ G55788 (dry specimen); type locality: Ras Mohammed National Park, Sharm al-Sheikh, Sinai Peninsula, Egypt, 17 m depth; phylogenetic data: none.

3. Dipsastraea amicorum ( Milne Edwards & Haime, 1849b, vol. 12: 171, vol. 10, pl. 9: fig. 9); holotype: MNHN IK-2010-470 (dry specimen; Fig. 10D ); type locality: Tongatapu, Tonga; phylogenetic data: none .

4. Dipsastraea camranensis ( Latypov, 2013: 223) ; holotype: FEBRAS 24193 (dry specimen); paratype: FEBRAS 24194 (dry specimen); type locality: Hon Nai Island , Cam Ranh Bay, Vietnam, 3 m depth; phylogenetic data: none .

5. Dipsastraea danai ( Milne Edwards & Haime, 1857, vol. 2: 442); holotype: USNM 32 (dry specimen); paratype: USNM 31 (dry specimen); type locality: Tongatapu, Tonga; phylogenetic data: molecular and partial morphology .

6. Dipsastraea helianthoides ( Wells, 1954: 458, pl. 174: figs 3–6); holotype: USNM 44980 (dry specimen); type locality: Bikini Island , Bikini Atoll, Marshall Islands; phylogenetic data: morphology only .

7. Dipsastraea lacuna (Veron, Turak & DeVantier, 2000, vol. 3: 111, fig. 6) (see also Veron, 2002: 139, figs 254–256; ICZN, 2011: 164); lectotype (designated herein): MTQ G55836 (dry specimen); type locality: northern Red Sea coast of Saudi Arabia; phylogenetic data: none.

8. Dipsastraea laddi ( Wells, 1954: 456, pl. 172: figs 1–4); holotype: USNM 44942 (dry specimen; Fig. 10G–I ); type locality: lagoon of Bikini Atoll , Marshall Islands, about 4 m depth; phylogenetic data: morphology only .

9. Dipsastraea laxa ( Klunzinger, 1879: 49, plate 5, fig. 3, plate 10, figs 9a, b); holotype: ZMB Cni 2193 (dry specimen); type locality: Koseir, Egypt; phylogenetic data: morphology only .

10. Dipsastraea lizardensis (Veron, Pichon & Wijsman- Best, 1977: 45, figs 74–78, 428–430); holotype: NHMUK 1977.1 .1.2 (dry specimen) ; paratype: MTQ G59707 (dry specimen) ; paratype: RMNH 10733 (dry specimen); type locality: McGillivray Reef , Lizard Island, Australia, 7 m depth; phylogenetic data: molecular and morphology .

11. Dipsastraea maritima ( Nemenzo, 1971: 169, pl. 9: figs 1, 2); holotype: UP C-859 (dry specimen); type locality: Puerto Princesa Bay , Palawan, the Philippines; phylogenetic data: none .

12. Dipsastraea marshae ( Veron, 2000, vol. 3: 122, figs 1, 2) (see also Veron, 2002: 145, figs 269, 270; ICZN, 2011: 164); lectotype (designated herein): WAM Z12910 (dry specimen); type locality: Ashmore Reef, northwest Australia, 9 m depth; phylogenetic data: none.

13. Dipsastraea matthaii ( Vaughan, 1918: 109, pl. 39: figs 2, 2a, b); holotype: USNM 38381 (dry specimen); type locality: Seychelles, Aldabra Atoll , Assumption Island, or Glorioso Islands; phylogenetic data: molecular and morphology .

14. Dipsastraea maxima (Veron, Pichon & Wijsman- Best, 1977: 43, figs 67–73, 427, 445); holotype: NHMUK 1977.1 .1.1 (dry specimen) ; paratype: MTQ G59706 (dry specimen) ; paratype: RMNH 10732 (dry specimen); type locality: Nara Inlet , Hook Island, Whitsunday Islands, Australia, 5 m depth; phylogenetic data: molecular and morphology .

15. Dipsastraea mirabilis ( Yabe & Sugiyama, 1941: 72, pl. 61: figs 1, 1a–e); holotype: TIU 64330 (dry specimen); type locality: Yap Islands ; phylogenetic data: molecular and morphology .

16. Dipsastraea pallida ( Dana, 1846: 224, pl. 10: figs 13, 13a–e); syntype: USNM 30 (dry specimen; Fig. 10J ); syntype: YPM IZ 4282 (dry specimen); type locality: Fiji; phylogenetic data: molecular and morphology .

17. Dipsastraea rosaria ( Veron, 2000, vol. 3: 119, figs 3, 4) (see also Veron, 2002: 143, figs 264–268; ICZN, 2011: 164); lectotype (designated herein): MTQ G55822 (dry specimen); type locality: Milne Bay, Papua New Guinea, 10 m depth; phylogenetic data: molecular and morphology.

18. Dipsastraea rotumana ( Gardiner, 1899: 750, pl. 47: fig. 3); holotype: lost ( Wijsman-Best, 1972: 21); neotype: ZMA Coel. 5686, designated by Veron et al. (1977: 41) (dry specimen); type locality: New Caledonia; phylogenetic data: molecular and morphology.

19. Dipsastraea speciosa ( Dana, 1846: 220, pl. 11: figs 1, 1a–d); syntype: USNM 37 (dry specimen); type locality: ‘East Indies’ ( Dana, 1846: 220); phylogenetic data: molecular and morphology.

20. Dipsastraea truncata ( Veron, 2000, vol. 3: 113, figs 3–6) (see also Veron, 2002: 142, figs 260–263; ICZN, 2011: 164); lectotype (designated herein): MTQ G55823 (dry specimen); type locality: Milne Bay, Papua New Guinea, 5 m depth; phylogenetic data: molecular and morphology.

21. Dipsastraea veroni ( Moll & Best, 1984: 48, figs 1–3) (see also Veron et al., 1977: 49, fig. 81); holotype: RMNH 15209 (dry specimen); paratypes: RMNH 15210–15215 (six dry specimens); type locality: 100 m offshore of east Kudingareng Keke, Spermonde Archipelago, Indonesia, 2 m depth; phylogenetic data: none.

22. Dipsastraea vietnamensis ( Veron, 2000, vol. 3: 127, figs 3–5) (see also Veron, 2002: 146, figs 271–273; ICZN, 2011: 164); lectotype (designated herein): MTQ G55859 (dry specimen); type locality: Nha Trang, Vietnam, 10 m depth; phylogenetic data: none.

Taxonomic remarks

This is a large genus that, prior to Budd et al. (2012), had all its species distributed amongst Favia Milne Edwards & Haime, 1857, vol. 2: 426, and Barabattoia Yabe & Sugiyama, 1941: 72 . It was discovered through molecular phylogenetic analyses that Favia was actually comprised of at least two main lineages separated according to the geographical divisions of the Indo-Pacific and the Atlantic ( Fukami et al., 2004 a, 2008). As the type species of Favia is Madrepora fragum Esper, 1795: 79 , an Atlantic species (see Hoeksema, Roos & Cadée, 2012), it followed that a taxonomic split of the genus will involve reassigning the Indo-Pacific species into the resurrected genus Dipsastraea de Blainville, 1830: 338 .

Until the recent revision, Dipsastraea had never been applied since it was established. Wells (1936: 109) showed that all the species initially assigned to Dipsastraea had been placed in other genera, thus fixing Madrepora favus Forskål, 1775: 132 , as the lectotype by elimination, following the transfer of Madrepora favosa Esper, 1795: 34 into Favia ( Milne Edwards & Haime, 1857, vol. 2: 443). Matthai (1914: 79) subsequently moved Madrepora favus Forskål into Favia as well, effectively synonymizing Dipsastraea as Favia .

Here, we show that morphologically Madrepora favus Forskål falls well within the large clade of Indo- Pacific Favia , corroborating molecular results that show that these species are closely related ( Fukami et al., 2004 a, 2008; Huang et al., 2011; Kongjandtre et al., 2012). However, three major issues need to be addressed.

First, the synonymy of Barabattoia mirabilis Yabe & Sugiyama, 1941: 72 , as Barabattoia amicorum by Veron et al., 1977: 32, is untenable, as these are clearly two distinct species. The specimen shown in Veron et al. (1977: fig. 37), is incorrectly referred to as the ‘ holotype of Favia amicorum ’. We have verified that MNHN specimen IK-2010-470 is the holotype of Barabattoia amicorum ( Fig. 10D ), following the original description in Milne Edwards & Haime (1849b, vol. 12: 171) and illustration in Milne Edwards & Haime (1848b, vol. 10, pl. 9: fig. 9). All the molecular trees used here have essentially followed the taxonomy of Veron et al. (1977) when analysing Barabattoia mirabilis . We thus regard them both as valid species, and all molecular terminals identified as Barabattoia amicorum to be Barabattoia mirabilis , which has consistently been placed within the Indo-Pacific Favia clade ( Fukami et al., 2008; Huang et al., 2011; Arrigoni et al., 2012; Huang, 2012). Supported by recovery of two Barabattoia spp. (i.e. Barabattoia laddi and Barabattoia mirabilis ) in the same clade on the morphological phylogeny, we consequently consider Barabattoia as a synonym of Dipsastraea .

Second, Astrea (Orbicella) stelligera Dana, 1846: 216 , and Favites rotundata Veron, Pichon & Wijsman-Best, 1977: 64 , are more closely related to Goniastrea and Favites , respectively, than Favia (or Dipsastraea ), and we give separate accounts below based on their phylogenetic affinities.

Third, our results show that Trachyphyllia geoffroyi ( Audouin, 1826: 233) , Goniastrea aspera Verrill, 1866: 32 , and Favia palauensis Yabe & Sugiyama, 1936: 30 , are morphologically distinct from Dipsastraea , but molecular data have often placed these species within the latter ( Fukami et al., 2004 a, 2008; Huang et al., 2011; Arrigoni et al., 2012; Huang, 2012). On the basis of the morphological evidence and long molecular branch lengths leading to these species, we placed them in two other genera described here (i.e. Trachyphyllia Milne Edwards & Haime, 1848a , vol. 27: 492, and Coelastrea Verrill, 1866: 32 ).

Dipsastraea 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 eastwards from Hawai’i in the north.

Morphological remarks

We find no apomorphies for Dipsastraea that are consistent across data types, mainly because the nesting of Coelastrea and Trachyphyllia on the molecular tree results in distinguishing features being optimized as plesiomorphic traits. Unequal costosepta is the only synapomorphy on the morphological phylogeny. In spite of this, Dipsastraea can be differentiated easily from the aforementioned close relatives by its moderate amount of coenosteum, three cycles of septa, and weak to moderate paliform lobes, rather than fused, limited walls or phaceloid colonies, four septal cycles, welldeveloped septal lobes. Thin sections also reveal that Dipsastraea has more distinct costa centre clusters but weaker costa and septum medial lines than Coelastrea and Trachyphyllia .

Being conventionally grouped with the Atlantic Favia spp. previously, the distinction between Dipsastraea and Favia is much clearer with the characters analysed here. Even with macromorphology, the differences are substantial, with Dipsastraea possessing larger and deeper corallites (after losing Goniastrea stelligera ), fewer and less crowded septa, columellae that are smaller but denser, and paliform (single axis) instead of septal (fan-shaped) lobes. Of the 23 subcorallite characters used, 14 are distinct between them. Aside from the family-level synapomorphies associated with tooth shape, the walls of Dipsastraea are formed primarily by paratheca instead of septotheca as in Favia .

This genus is fairly well sampled, but most of the more recently described species of Veron (2000) are lacking data.

GENUS ECHINOPORA LAMARCK, 1816: 252

( FIG. 11 )

Synonyms

Acanthelia Wells, 1937: 73 (type species: Echinopora horrida Dana, 1846: 282 , pl. 17: figs 4, 4a–c; original designation, Wells, 1937: 73); Acanthopora Verrill, 1864: 54 (type species: Echinopora horrida Dana, 1846: 282 , pl. 17: figs 4, 4a–c; original designation, Verrill, 1864: 54); Echinastraea de Blainville, 1830: 343 (type species: Echinopora rosularia Lamarck, 1816: 253 = Madrepora lamellosa Esper, 1795: 65 , pl. 58: figs 1, 2; original designation, de Blainville, 1830: 344); Heliastraea Milne Edwards & Haime, 1857, vol. 2: 456 (type species: Madrepora astroites Forskål, 1775: 133 = Astrea forskaliana Milne Edwards & Haime, 1849b, vol. 12: 100; original designation, Milne Edwards & Haime, 1857, vol. 2: 457); Stephanocora Ehrenberg, 1834: 300 (type species: Stephanocora hemprichii Ehrenberg, 1834: 300 = Explanaria gemmacea Lamarck, 1816: 256 ; original designation, Ehrenberg, 1834: 300).

Type species

Echinopora rosularia Lamarck, 1816: 253 = Madrepora lamellosa Esper, 1795: 65 , pl. 58: figs 1, 2 (see Matthai, 1914: 50); original designation, Lamarck, 1816: 253; holotype: MNHN IK-2010-635 (dry specimen; Fig. 11A ); type locality: ‘les mers de la Nouvelle-Hollande’ ( Lamarck, 1816: 254).

Original description

‘Polypier pierreux, fixé, aplati et étendu en membrane libre, arrondie, foliiforme, finement striée des deux côtés. La surface supérieure chargée de petites papilles, et, en outre, d’orbicules rosacés, convexes, trèshérissés de papilles, percés d’un ou deux trous, recouvrant chacun une étoile lamelleuse.

Étoiles éparses, orbiculaires, couvertes; à lames inégales, presque confuses, saillantes des parois et du fond, et obstruant en partie la cavité.’ ( Lamarck, 1816: 252).

Subsequent descriptions

Lamouroux, 1821: 57; Lamarck, 1836: 395, 396; Dana, 1846: 277, 278; Milne Edwards & Haime, 1849b, vol. 12: 185; Milne Edwards & Haime, 1857, vol. 2: 621; Dana, 1859: 42; Klunzinger, 1879: 54, 55; Quenstedt, 1881: 1030–1031; Duncan, 1884: 117; Ortmann, 1890: 298, 299; Saville Kent, 1893: 170; Delage & Hérouard, 1901: 631; Gardiner, 1904: 782; Matthai, 1914: 48, 49; Hickson, 1924: 61; Faustino, 1927: 122; Yabe et al., 1936: 48; Vaughan & Wells, 1943: 175; Alloiteau, 1952: 625; Crossland, 1952: 118; Wells, 1956: F406; Nemenzo, 1959: 117; Chevalier, 1975: 68, 69; Veron et al., 1977: 182; Wijsman-Best, 1980: 239, 240; Scheer & Pillai, 1983: 135; Wood, 1983: 170; Veron, 1986: 526; Chevalier & Beauvais, 1987: 716; Sheppard, 1990: 16; Sheppard & Sheppard, 1991: 141; Veron, 2000, vol. 3: 252.

Diagnosis (apomorphies in italics)

Colonial, with extracalicular budding only. Corallites monomorphic and discrete (one to three centres); monticules absent. Coenosteum generally spinose (costate in Echinopora mammiformis ), extensive amount (≥ corallite diameter). Generally, calice width medium (4–15 mm), with low relief (<3 mm). Costosepta not confluent. Septa in three cycles (24–36 septa). Free septa regular. Septa spaced> 11 septa per 5 mm. Costosepta unequal in relative thickness. Columellae trabecular and spongy (> three threads), ≥ 1/4 of calice width, and discontinuous amongst adjacent corallites. Paliform (uniaxial) lobes weak or moderate. Epitheca well developed and endotheca low−moderate (tabular) ( Fig. 11A, D, G ).

Tooth base at midcalice circular. Tooth tip at midcalice irregular; tip orientation multiaxial. Tooth height low (<0.3 mm) and tooth spacing medium (0.3– 1 mm), with> six teeth per septum. Granules scattered on septal face; irregular in shape. Interarea smooth ( Fig. 11B, E, H ).

Walls formed by partial septotheca; abortive septa weak. 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. 11C, F, I ).

Species included

1. Echinopora lamellosa ( Esper, 1795: 65, pl. 58: figs 1, 2); holotype: lost ( Chevalier, 1975: 70; Scheer, 1990: 398); type locality: unknown; phylogenetic data: molecular and morphology.

2. Echinopora ashmorensis Veron, 1990: 152 , figs 58– 62, 87, 88; holotype: MTQ G32491 (dry specimen); type locality: Ashmore Reef , Western Australia, 2 m depth; phylogenetic data: none .

3. Echinopora forskaliana ( Milne Edwards & Haime, 1849b, vol. 12: 100); holotype: MNHN IK-2010- 406 (dry specimen); type locality: Red Sea; phylogenetic data: none .

4. Echinopora fruticulosa Klunzinger, 1879: 55 = Stephanocora hemprichii forma fruticulosa Ehrenberg, 1834: 301 ; holotype: ZMB Cni 749, see Matthai (1914: 56) (dry specimen); type locality: Red Sea; phylogenetic data: none.

5. Echinopora gemmacea ( Lamarck, 1816: 256) ; holotype: MNHN IK-2010-529 (dry specimen); type locality: ‘l’Océan indien?’ ( Lamarck, 1816: 256); phylogenetic data: molecular and morphology.

6. Echinopora hirsutissima Milne Edwards & Haime, 1849b, vol. 12: 187; holotype: MNHN IK-2010- 491 (dry specimen); type locality: ‘l’océan Indien’ ( Milne Edwards & Haime, 1849b, vol. 12: 187); phylogenetic data: morphology only.

7. Echinopora horrida Dana, 1846: 282 , pl. 17: figs 4, 4a–c; syntype: USNM 162 (dry specimen; Fig. 11D ); syntypes: YPM IZ 1980 A, B, 4307 (three dry specimens); type locality: Fiji; phylogenetic data: molecular and morphology .

8. Echinopora irregularis Veron, Turak & DeVantier, 2000 , vol. 3: 262, fig. 1 (see also Veron, 2002: 175, figs 318–321; ICZN, 2011: 163); lectotype (designated herein): MTQ G55835 (dry specimen); type locality: northern Red Sea coast of Saudi Arabia, 2 m depth; phylogenetic data: none.

9. Echinopora mammiformis ( Nemenzo, 1959: 112, pl. 14: fig. 2); holotype: UP C-99 (dry specimen; Fig. 11G ); type locality: Muelle , Puerto Galera, the Philippines; phylogenetic data: molecular and partial morphology .

10. Echinopora pacificus Veron, 1990: 150 , figs 55– 57, 86; holotype: MTQ G32490 (dry specimen); type locality: entrance to Kabira Bay , Ishigaki Island, Ryukyu Islands, Japan, 15 m depth; phylogenetic data: molecular and morphology .

11. Echinopora robusta Veron, 2000 , vol. 3: 263, figs 2–4 (see also Veron, 2002: 176, figs 322–324; ICZN, 2011: 163); lectotype (designated herein): MTQ G55849 (dry specimen); type locality: southern Sri Lanka, 2 m depth; phylogenetic data: none.

12. Echinopora taylorae ( Veron, 2000, vol. 2: 327, fig. 6) (see also Veron, 2002: 173, figs 315–317; ICZN, 2011: 163); lectotype (designated herein): UP MSI- 3005-CO (dry specimen); type locality: Calamian Islands, Palawan, the Philippines, 12 m depth; phylogenetic data: none.

13. Echinopora tiranensis Veron, Turak & DeVantier, 2000 , vol. 3: 265, figs 4, 5 (see also Veron, 2002: 178, figs 322–324; ICZN, 2011: 163); lectotype (designated herein): MTQ G55843 (dry specimen); type locality: Tiran Island, northern Red Sea coast of Saudi Arabia, 15 m depth; phylogenetic data: none.

Taxonomic remarks

Echinopora Lamarck, 1816: 252 , is a relatively large genus, with four new species only recently described ( Veron, 2000). It was first described as having an upper surface filled with small papillae – ‘la surface supérieure chargée de petites papilles’ ( Lamarck, 1816: 252) – a plesiomorphic trait shared with Cyphastrea . Togeth- er with Paramontastraea and Orbicella , these taxa have been consistently recovered at the base of the tree, either as paraphyletic ( Huang et al., 2011; Huang, 2012), or as a sister clade to the rest of Merulinidae ( Arrigoni et al., 2012) . The latter hypothesis appears to be more well supported with molecular data, and it also corresponds to the morphological tree topology obtained here.

It should be noted that the type species of Echinopora is Echinopora rosularia Lamarck, 1816: 253 , which has been synonymized as Echinopora lamellosa ( Esper, 1795: 65; Ranson, 1943: 118). The latter’s holotype is lost ( Chevalier, 1975: 70; Scheer, 1990: 398), but Lamarck’s holotype of Echinopora rosularia (MNHN IK-2010- 635; Fig. 11A ) should still be considered the type for this genus.

Echinopora is widely distributed on reefs of the Indo- Pacific, present as far east as the Tuamotu Archipelago in the Southern Hemisphere ( Glynn et al., 2007), but absent eastwards from Hawai’i in the north.

Morphological remarks

This genus is one of the most distinct and welldefined genera in Merulinidae , being supported by a high bootstrap value (93) and decay index (2) on the morphology tree. Synapomorphies inferred are large columellae (≥ 1/4 of calice width; likelihood of 0.86 based on the Mk1 model), extensive coenosteum (≥ corallite diameter; likelihood 0.77), and weak abortive septa (0.98). The latter two features distinguish Echinopora from the closely related genera of Cyphastrea , Paramontastraea , and Orbicella . Large columella is only shared with Orbicella amongst all Merulinidae taxa. Data are available only for six of the 13 species; the genus requires substantial additional sampling, particularly for the recently described species. None of the species described in Veron (2000) have been placed on the phylogeny.

Figures

Figure 10. Dipsastraea de Blainville, 1830, has discrete corallites that bud intracalicularly, equally thick costosepta, and spongy columellae. Septal teeth with medium height (0.3–0.6 mm) and spacing (0.3–1 mm). Walls formed by dominant paratheca and partial septotheca, with transverse septal crosses. A–C, Dipsastraea favus (Forskål, 1775), type species of Dipsastraea; macromorphology, lectotype ZMUC ANT-000466, Red Sea (A); micromorphology (scanning electron microscopy; B) and microstructure (transverse thin section; C), hypotype USNM 93662, Madang, Papua New Guinea. D, Dipsastraea amicorum (Milne Edwards & Haime, 1849b); macromorphology, holotype MNHN IK-2010-470, Tongatapu, Tonga (photo by P. Lozouet). E, F, Dipsastraea mirabilis (Yabe & Sugiyama, 1941); micromorphology (E) and microstructure (F), hypotype USNM 93642, Madang, Papua New Guinea. G–I, Dipsastraea laddi (Wells, 1954); macromorphology (G), micromorphology (H), and microstructure (I), holotype USNM 44942, Bikini Atoll, Marshall Islands. J–L, Dipsastraea pallida (Dana, 1846); macromorphology, syntype USNM 30, Fiji (J); micromorphology (K) and microstructure (L), hypotype USNM 44952, Bikini Atoll, Marshall Islands.

Figure 11. Echinopora Lamarck, 1816, has discrete corallites that bud extracalicularly, extensive coenosteum (≥ corallite diameter), medium-size (4–15 mm) and low-relief (<3 mm) calices, large (≥ 1/4 of calice width) spongy columellae, and weak/moderate paliform (uniaxial) lobes. Septal teeth are low (<0.3 mm) with medium spacing (0.3–1 mm) and multiaxial tips. Walls formed by partial septotheca and weak abortive septa. A–C, Echinopora lamellosa (Esper, 1795), type species of Echinopora; macromorphology, Echinopora rosularia Lamarck, 1816, holotype of Echinopora MNHN IK-2010-635, unknown locality (A); micromorphology (scanning electron microscopy; B) and microstructure (transverse thin section; C), hypotype USNM 89851, Alofi, Niue. D–F, Echinopora horrida Dana, 1846; macromorphology, syntype USNM 162, Fiji (D); micromorphology (E) and microstructure (F), hypotype RMNH 33956, Kepulauan Seribu, Indonesia. G, Echinopora mammiformis (Nemenzo, 1959); macromorphology, holotype UP C-99, Puerto Galera, the Philippines. H, I, Echinopora gemmacea (Lamarck, 1816); micromorphology, hypotype RMNH 17270, Watamu, Kenya (H); microstructure, hypotype USNM 1113168, Red Sea (I).