Favites, LINK, 1807: 162

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 : 314-317

GENUS FAVITES LINK, 1807: 162 ( FIG. 13 )

Synonyms

Aphrastrea Milne Edwards & Haime, 1848a, vol. 27: 495 (type species: Astrea deformis Lamarck, 1816: 264 = Madrepora pentagona Esper, 1795: 23 , pl. 39: figs 1, 2; original designation, Milne Edwards & Haime, 1848a, vol. 27: 495); Astrophyllia Ehrenberg, 1834: 322 (type species: not designated); Phymastrea Milne Edwards & Haime, 1848a, vol. 27: 494 (type species: Phymastrea valenciennesii Milne Edwards & Haime, 1849b, vol. 12: 124, vol. 10, pl. 9: figs 3, 3a; subsequent designation, Milne Edwards & Haime, 1849b, vol. 12: 124); Prionastrea Milne Edwards & Haime, 1848a, vol. 27: 495 (type species: Astrea abdita Lamarck, 1816: 265 ; original designation, Milne Edwards & Haime, 1848a, vol. 27: 495).

Type species

Favites astrinus Link, 1807: 162 = Madrepora abdita Ellis & Solander, 1786 (see Vaughan, 1901a: 21; Vaughan, 1918: 109); original designation, Link, 1807: 162.

Original description

‘Wabenkoralle. Unförmige, kalkartige Massen, mit oberflaechlichen zerstreuten sternförmig blaettrigen Öffnungen.’ ( Link, 1807: 162).

Subsequent descriptions

Verrill, 1901: 92; Vaughan, 1918: 109; Vaughan, 1919: 414; Hoffmeister, 1925: 24; Faustino, 1927: 134; Coryell & Ohlsen, 1929: 200; Yabe et al., 1936: 31; Vaughan & Wells, 1943: 167; Alloiteau, 1952: 616, 617; Wells, 1956: F402; Nemenzo, 1959: 93; Chevalier, 1971: 178; Wijsman-Best, 1972: 26; Veron et al., 1977: 50–53; Scheer & Pillai, 1983: 113; Wood, 1983: 147, 150; Veron, 1986: 468; Chevalier & Beauvais, 1987: 714; Veron & Hodgson, 1989: 271; Sheppard, 1990: 10; Sheppard & Sheppard, 1991: 126; Veron, 2000, vol. 3: 134, 135.

Diagnosis

Colonial, with intra- and extracalicular budding. Corallites monomorphic and discrete (one to three centres); monticules absent. Coenosteum costate, limited amount (includes double wall) or fused walls. Calice width medium (4–15 mm), but may be larger (> 15 mm), with medium relief (3–6 mm). Costosepta may be confluent. Septa generally in ≥ four cycles (≥ 48 septa). Free septa present but irregular. Septa spaced six to 11 septa per 5 mm. Costosepta unequal in relative thickness. Columellae trabecular and spongy (> three threads), <1/4 of calice width, and continuous amongst adjacent corallites. Paliform (uniaxial) lobes weak to well developed. Epitheca well developed and endotheca generally abundant (vesicular) ( Fig. 13A, 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. 13B, E, H, K ).

Walls formed by dominant paratheca and partial septotheca; abortive septa absent. Thickening deposits fibrous. Costa centre clusters generally strong; 0.3– 0.6 mm between clusters; medial lines weak. Septum centre clusters weak; 0.3–0.5 mm between clusters; medial lines weak or strong. Transverse crosses generally present. Columella centres clustered ( Fig. 13C, F, I, L ).

Species included

1. Favites abdita ( Ellis & Solander, 1786: 162, pl. 50: fig. 2); holotype: GLAHM 104005 (dry specimen; Fig. 13A ); type locality: ‘probablement les mers des Grandes-Indes’ ( Lamarck, 1816: 265); phylogenetic data: molecular and morphology.

2. Favites acuticollis ( Ortmann, 1889: 528, pl. 16: fig. 11); holotype: ZMB Cni 4793 (dry specimen); type locality: Sri Lanka; phylogenetic data: none .

3. Favites chinensis ( Verrill, 1866: 35) ; holotype: YPM IZ 1002 (dry specimen); type locality: Hong Kong; phylogenetic data: molecular and morphology .

4. Favites colemani ( Veron, 2000, vol. 3: 219, figs 6–11) (see also Veron, 2002: 164, figs 301, 302; ICZN, 2011: 164); lectotype (designated herein): UP MSI- 3008-CO (dry specimen); type locality: Calamian Islands, Palawan, the Philippines, 15 m depth; phylogenetic data: molecular and morphology.

5. Favites complanata ( Ehrenberg, 1834: 317) ; holotype: ZMB Cni 695 (dry specimen); type locality: Red Sea; phylogenetic data: molecular and morphology .

6. Favites flexuosa ( Dana, 1846: 227, pl. 11: figs 6, 6a–e); syntype: USNM 27 (dry specimen; Fig. 13D ); type locality: Fiji; phylogenetic data: molecular and morphology .

7. Favites halicora ( Ehrenberg, 1834: 321) ; holotype: ZMB Cni 733, lost ( Chevalier, 1971: 197; not found in ZMB), figured in Klunzinger (1879, pl. 4: fig. 1); type locality: Red Sea; phylogenetic data: molecular and morphology.

8. Favites magnistellata ( Chevalier, 1971: 293, pl. 9: fig. 3, pl. 34: fig. 2); holotype: H 78 m ( Chevalier, 1971: 293), MNHN status unknown; type locality: fringing reef near southwest Hugon Island, New Caledonia; phylogenetic data: molecular and morphology.

9. Favites melicerum ( Ehrenberg, 1834: 320) = Favites bestae Veron, 2000 , vol. 3: 140, figs 1, 2 (see also Veron, 2002: 150, figs 277–279; ICZN, 2011: 164); holotype: ZMB Cni 734, lost ( Wijsman-Best, 1972: 29; Veron, 2002: 150), figured in Matthai, 1914, pl. 36: fig. 4; neotype (designated herein): ZMA Coel. 5820 (dry specimen); type locality: southern New Caledonia, 5 m depth; phylogenetic data: none.

10. Favites micropentagonus Veron, 2000 , vol. 3: 137, figs 6–9 (see also Veron, 2002: 148, figs 274–276; ICZN, 2011: 164); lectotype (designated herein): UP MSI-3006-CO (dry specimen); type locality: Calamian Islands, Palawan, the Philippines, 12 m depth; phylogenetic data: none.

11. Favites monticularis Mondal, Raghunathan & Venkataraman, 2013: 4510 , figs 1, 2; holotype: ZSI/ ANRC-7410 (dry specimen); type locality: off Shibpur, Diglipur, North Andaman, 14 m depth; phylogenetic data: none.

12. Favites paraflexuosus Veron, 2000 , vol. 3: 155, figs 4–6 (see also Veron, 2002: 151, figs 280–282; ICZN, 2011: 164); lectotype (designated herein): WAM Z12911 (dry specimen); type locality: Houtman Abrolhos Islands, Western Australia, 15 m depth; phylogenetic data: molecular and morphology.

13. Favites pentagona ( Esper, 1795: 23, pl. 39: figs 1, 2); holotype: lost ( Chevalier, 1971: 216; Scheer, 1990: 390); type locality: ‘probablement l’Océan indien’ ( Lamarck, 1816: 264); phylogenetic data: molecular and morphology.

14. Favites rotundata Veron, Pichon & Wijsman-Best, 1977: 64 , figs 110–117, 436–438; holotype: NHMUK 1977.1 .1.6 (dry specimen; Fig. 13G, H ); paratype: RMNH 10734 (dry specimen); type locality: southwest Swain Reefs , Australia, 5 m depth; phylogenetic data: molecular and morphology .

15. Favites russelli ( Wells, 1954: 460, pl. 174: figs 7, 8); holotype: USNM 45004 (dry specimen); type locality: seaward slope of Bikini Atoll , Marshall Islands, 53–77 m depth; phylogenetic data: molecular and morphology .

16. Favites solidocolumellae Latypov, 2006: 149 , figs 38: 7, 8 (= Favites sp. 1 Latypov, 1995: 48 , pl. 8: fig. 1); holotype: FEBRAS 1/95118 (dry specimen); type locality: Nha Trang Bay, Chuong Island, Vietnam, 3 m depth; phylogenetic data: none.

17. Favites spinosa ( Klunzinger, 1879: 39, pl. 4: fig. 7, pl. 10: fig. 5); holotype: ZMB 2154 (dry specimen); type locality: Red Sea; phylogenetic data: none .

18. Favites stylifera Yabe & Sugiyama, 1937: 426 , fig. 1; holotype: NSMT (dry specimen); type locality: Yoronjima , Kagoshima, Japan; phylogenetic data: none .

19. Favites valenciennesi ( Milne Edwards & Haime, 1849b, vol. 12: 124, vol. 10, pl. 9: figs 3, 3a; see Article 58.14 of the Code); holotype: MNHN IK- 2010-696 (dry specimen; Fig. 13J ); type locality: unknown; phylogenetic data: molecular and morphology .

20. Favites vasta ( Klunzinger, 1879: 38, pl. 4: fig. 12, pl. 10: fig. 4a, b); holotype: ZMB Cni 2176 (dry specimen); type locality: ‘ Kossier’ (specimen label), Egypt, Red Sea; phylogenetic data: none .

Taxonomic remarks

Favites Link, 1807: 162 , has been a difficult genus to define. By convention, species tend to have ‘cerioid, occasionally subplocoid’ ( Veron, 2000, vol. 3: 134) corallites. There is now little doubt that the clade with the majority of Favites spp. , including the type species Favites abdita , also contains species with fully plocoid corallites such as Phymastrea valenciennesi Milne Edwards & Haime, 1849b, vol. 12: 124, and Montastrea colemani Veron, 2000 , vol. 3: 219 ( Arrigoni et al., 2012), whereas Montastraea magnistellata Chevalier, 1971: 293 , is sister to this clade ( Huang et al., 2011; Huang, 2012). In this sense, Favites has been a paraphyletic group. The solution proposed here is thus to move the three species above into Favites .

On the one hand, recovery of Favites pentagona , Favites russelli , and Favites peresi (a Goniastrea sp. according to Veron, 2000, vol. 3: 166) separately in distant lineages renders the genus polyphyletic ( Huang et al., 2011; Arrigoni et al., 2012). We resolve this partially by moving Favites peresi into the new genus Paramontastraea Huang & Budd. On the other hand, we find limited morphological basis for transferring Favites pentagona out of the genus because it is the sister group to the rest of Favites on the morphology tree. Favites micropentagonus ‘looks like a diminutive form of the well know [sic] Favites pentagona ’ ( Veron, 2002: 148) and is thus a likely sister species to Favites pentagona . For both species, further molecular sampling will clarify their affinities.

Favites bestae is a junior synonym of Astraea melicerum Ehrenberg, 1834: 320 described by Veron (2000, vol. 3: 140; see also Veron, 2002: 150), although the latter name is sometimes considered a synonym of Favites pentagona ( Matthai, 1914: 95; Chevalier, 1971: 215; see also Wijsman-Best, 1972: 30). The reason given for establishing this species is that the holotype of the senior synonym had been lost, and thus the name Favites melicerum is ‘unverifiable’. As Veron (2000, vol. 3: 140) deemed Favites bestae to be a separate species from Favites pentagona , by extension Favites melicerum is also regarded as distinct from Favites pentagona , a view held by Vaughan (1918: 112). The use of Favites bestae as a ‘new name’ or ‘nomen novum’ is considered unnecessary because it is neither a replacement for a preoccupied name (Article 60.3 of the Code; see Hoeksema, 1993) nor a substitute for an unavailable or invalid name (Article 23.3.5 of the Code). Nevertheless, a neotype needs to be designat- ed for its senior synonym Favites melicerum , a task we have undertaken above.

Favites 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

We find no apomorphies for Favites that are consistent across data types, primarily because of the recovery of Favites russelli and Favites pentagona in distant parts of the molecular phylogeny. Few characters sepa- rate them from other Favites spp. , such as the number of septa and distinctiveness of costa centre clusters, and further studies are warranted to determine if they should be distinguished as separate genera.

For reasons unknown, Favites rotundata Veron, Pichon & Wijsman-Best, 1977: 64 , was placed in Favia by Veron (2000, vol. 3: 124) although its ‘coralla are subplocoid’ ( Veron et al., 1977: 64). Morphological and molecular analyses consistently recover this species within the Favites clade (Fig. 2; Huang et al., 2011; Arrigoni et al., 2012; Huang, 2012), supporting its original placement within Favites . Favia marshae Veron, 2000 , vol. 3: 122, was described as a morphologically similar species (see also Huang, 2012), but without molecular data to justify this affinity, we preserve its membership within Dipsastraea .

The name Montastraea valenciennesi has been applied on two disparate plocoid species differing in the degree of separation between adjacent corallite walls ( Fukami & Nomura, 2009). Presumably, the ‘corallite-wall separate type’ is a Dipsastraea species , whereas the ‘corallitewall fusion type’ is the one recovered within the Favites clade ( Huang et al., 2011). Based on thin section observations, we find no difference in wall separation between the two types. However, two synapomorphies of the most inclusive Favites clade omitting Favites pentagona – septa in four or more cycles and strong costa centre clusters – are clearly present in the specimen recovered within the Favites clade (UP P1 L02131 ; Fig. 13L ). The specimen also possesses unequal costosepta and well-developed paliform lobes that are found in Milne Edwards & Haime’s holotype of Phymastrea valenciennesi . These traits are missing in the other type, which should therefore be considered as a cryptic Dipsastraea species.

Figures

Figure 13. Favites Link, 1807, has discrete corallites with double or fused walls, septa generally in ≥ four cycles (≥ 48 septa), weak to well-developed paliform (uniaxial) lobes, 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 strong costa centre clusters and transverse septal crosses. A–C, Favites abdita (Ellis & Solander, 1786), type species of Favites; macromorphology, holotype GLAHM 104005, unknown locality (A; photo by K. G. Johnson); micromorphology (scanning electron microscopy; B) and microstructure (transverse thin section; C), hypotype RMNH 10760, Pulau Air, Indonesia. D–F, Favites flexuosa (Dana, 1846); macromorphology, syntype USNM 27, Fiji (D); micromorphology (E) and microstructure (F), hypotype RMNH 14165, New Caledonia. G–I, Favites rotundata Veron, Pichon & Wijsman-Best, 1977; macromorphology (G) and micromorphology (H; photo by N. Santodomingo), holotype NHMUK 1977.1.1.6, southwest Swain Reefs, Australia; microstructure, hypotype MTQ G61874, Pelorus Island, Australia (I). J–L, Favites valenciennesi (Milne Edwards & Haime, 1849b); macromorphology, holotype MNHN IK-2010-696, unknown locality (J); micromorphology (K) and microstructure (L), hypotype UP P1L02131, Batangas, the Philippines.