Blastomussa omanensis ( Sheppard & Sheppard, 1991 )

Benzoni, Francesca, Arrigoni, Roberto, Waheed, Zarinah, Stefani, Fabrizio & Hoeksema, Bert W., 2014, Phylogenetic relationships and revision of the genus Blastomussa (Cnidaria: Anthozoa: Scleractinia) with description of a new species, Raffles Bulletin of Zoology 62, pp. 358-378 : 363-366

Blastomussa omanensis ( Sheppard & Sheppard, 1991)

( Figs. 1J–L , 6A–B , 7C , 8C , 9C )

Parasimplastrea omanensis — Sheppard, 1985 (nomen nudum)

Parasimplastrea omanensis Sheppard & Sheppard : 1991, fig. 147 (in synonymy of Parasimplastrea simplicitexta ); Pichon et al.: 2010, figs. 1–4

Parasimplastrea simplicitexta Sheppard & Sheppard, 1991 , fig. 147; not: Veron & Kelley, 1988 (partim): 49, fig. 16D. Not: Goniastrea simplicitexta Umbgrove, 1942 ; 35, pl. 12 fig. 5, pl. 13, Fig. 5

Parasimplastrea sheppardi Veron, 2000 : Volume 3, p. 239, figs. 7–10; Moothien Pillai et al.: 2002, figs. 1–3; Veron, 2002 partim: figs. 309, 311; Claereboudt, 2006: figs. 1–6

Type material. Holotype of B. omanensis ( BMNH 1991.6.4.150), Oman, Dhofar region, coll. C. Sheppard, 7 m (specimen illustrated in Sheppard & Sheppard, 1991, Fig. 147) ( Figs. 6A, B ), designation by monotypy.

Other material. Oman ( USNM 81272 ), Muscat, coll. C. Sheppard ; Yemen ( UNIMIB BAL037 ), Gulf of Aden, Balhaf , (13°58.163'N; 48°10.928'E), coll. F. Benzoni, 6 March 2007 , 14 m; ( UNIMIB BAL212 ), Gulf of Aden, Balhaf , (13°58.402'N; 48°12.410'E), coll. F. Benzoni, 23 September 2007; ( UNIMIB BAL230 ) , Gulf of Aden, Balhaf , (13°50.4167'N; 48°10.5167'E), 23 September 2007; ( UNIMIB Y571 ) , Gulf of Aden, Balhaf , (13°58.163'N; 48°10.928'E), coll. F. Benzoni, 6 March 2007; ( UNIMIB Y748 ) , Gulf of Aden, Balhaf , (13°50.4167'N; 48°10.5167'E), coll. F. Benzoni, 13 March 2008; ( UNIMIB MU094 ) , Gulf of Aden, Al Mukallah , (14°31.067'N; 49°10.335'E), coll. F. Benzoni, M. Pichon & C. Riva, 18 March 2007; ( UNIMIB MU160 ) , Gulf of Aden, Al Mukallah , (14°30.793'N; 49°10.339'E), coll. F. Benzoni, M. Pichon & C. Riva, 20 March 2007; ( UNIMIB MU205 ) , Gulf of Aden, Al Mukallah , (14°31.477'N; 49°07.855'E), coll. F. Benzoni, M. Pichon & C. Riva, 21 March 2007; ( UNIMIB BU016 ) , Gulf of Aden, Burum , (14°19.710'N; 48°59.903'E), coll. F. Benzoni, M. Pichon & C. Riva, 22 March 2007; ( UNIMIB SO010 ) , Arabian Sea, Socotra Island, Deubhil , (12°36.279'N; 54°21.053'E), coll. F. Benzoni & A. Caragnano, 11 March 2010; ( UNIMIB SO037 ) , Arabian Sea, Socotra Island, Ras Adho , (12°38.638'N; 54°16.147'E) coll. F. Benzoni & A. Caragnano, 13 March 2010; ( UNIMIB SO052 ) , Arabian Sea, Socotra Island, Ras Adho , (12°38.672'N; 54°16.043'E), coll. F. Benzoni & A. Caragnano, 13 March 2010 .

Description. Blastomussa omanensis forms encrusting subcerioid to cerioid coralla ( Figs. 1J, L , 6A, B , 7C ). Budding extra-tentacular, corallites are joined by secondary fusions, as described by Head (1978) for B. loyae , and the spaces between the partial fusions give the inter-corallite area a typical “groove and tubercule” appearance ( Sheppard & Sheppard, 1991). In specimens with tightly packed corallites these are not visible and corallite walls appear fused. Corallites are irregularly polygonal and 4–7 mm in largest diameter ( Figs. 1J, L , 6A, B , 7C ). Three cycles of septa are present, the first is generally complete and reaches the columella, the second can be incomplete and of variable length, the third reduced or incomplete ( Figs. 1J, L , 6A, B , 7C ). Septa are composed of one fan system thus margins are smooth ( Fig. 1L ). Septa are only slightly and equally exsert from the colony surface ( Fig. 1K ). Septal margins and sides finely granulated ( Figs. 1L ). Columella formed by loose trabecular processes and few papillae, seldom fused at the base ( Figs. 1L , 6B , 7C ).

Polyp tentacles and mantle vesicles expanded during the day until the polyps are mechanically disturbed and become retracted ( Fig. 8C ). Mantle vesicles smooth but forming lobes ( Fig. 9C ). Tentacles and vesicles uniformly brown, tentacle tips round and white, oral disc green. In contrast to its congenerics, this species shows a remarkably consistent colouration among colonies

This species is found in the same protected habitats as B. merleti with which it can co-occur.

Taxonomic remark. Blastomussa omanensis ( Sheppard & Sheppard, 1991) was originally presented as a nomen nudum in an unpublished report by Sheppard (1985). The species was formally redescribed by Sheppard & Sheppard (1991), who renamed it “ Parasimplastrea simplicitexta ( Umbgrove, 1939) ” because they erroneously assumed P. omanensis to be a synonym of Goniastrea simplicitexta Umbgrove, 1942 , based on remarks from J.E.N. Veron and J.W. Wells. Sheppard & Sheppard (1991) referred to Parasimplastrea omanensis in their synonymy of P. simplicitexta and they presented a photograph of a single coral (BMNH 1991.6.4.150) from Oman, which therefore became the holotype of P. omanensis by monotypy. Goniastrea simplicitexta was not described by Umbgrove in 1939, as mentioned by various authors ( Sheppard & Sheppard, 1991; Veron 2000). However, in that year Umbgrove described Simplastrea vesicularis Umbgrove, 1939 , which might have caused the confusion in the years. P. simplicitexta ( Umbgrove, 1942) is a valid species, which differs from P. omanensis (see Veron, 2000, 2002) and is only known from fossil corals found in Indonesia and Papua New Guinea ( Veron & Kelley, 1988). According to Budd et al. (2012: Table 1), P. omanensis is a synonym of G. simplicitexta but they do not give an explanation and do not mention the different view given by others ( Veron & Kelley, 1988; Sheppard & Sheppard, 1991: Veron 2000, 2002). We maintain the name Blastomussa omanensis instead of B. simplicitexta because no arguments are given to support their synonymy.

Veron (2000: Volume 3, p. 239) gave a new name, Parasimplastrea sheppardi , to Sheppard & Sheppard’s (1991) species and presented an unnumbered figure containing a black and white photograph of a coral skeleton without locality data and four colour photographs that were taken by others at Oman and Socotra Island. Hence, P. sheppardi Veron, 2000 , became an objective junior synonym of P. omanensis . Veron (2000) did not designate a holotype, but the black and white photograph distinctly shows a specimen of P. omanensis , which could serve as lectotype but its whereabouts are unknown. In a subsequent publication, Veron (2002) explained why a new name was given: “The name Parasimplastrea omanensis cannot be used because there is no holotype associated with it”. Instead of designating a neotype for P. omanensis, Veron (2002) designated a coral from Egypt (Red Sea) as “ holotype ” (= MTQ G 55860) for Parasimplastrea sheppardi . This designation is invalid because a holotype should have been introduced with the original description ( ICZN, 2011). Therefore, this coral should be considered neotype of P. sheppardi . Because this specimen actually belongs to Blastomussa loyae Head, 1978 , P. sheppardi is a subjective junior synonym of another species than intended by Veron. New names should only be given to species that have the same names as other species and when this homonymy would cause confusion ( Hoeksema, 1993).

Geographic distribution. Blastomussa omanensis has been recorded in the northern Gulf of Aden, the Arabian Sea, and Mauritius (see references mentioned above).

Blastomussa vivida , new species, Benzoni, Arrigoni & Hoeksema 2013 ( Figs. 1M–O , 2 , 3 , 7E , 8E , 9E )

Blastomussa wellsi Veron & Pichon, 1980 : fig. 768; Veron, 1986: Fig. 2 ; Veron, 2000: Vol. 3, pp. 6–7, Figs. 4–5 ; Hoeksema & van Ofwegen, 2004 partim; Wallace et al., 2009: Fig. 60A–B; Dai & Horng, 2009: 152; Turak & DeVantier, 2011: 163

Genus et Species nov.? Yabe et al., 1936, Pl. LII Fig. 2

Holotype. ( MNHN IK 2012 14226), New Caledonia, Canal Woodin, ST 332, coll. F. Benzoni & B.W. Hoeksema, 25 April 2012. IRD collection code HS3289 ( Figs. 2A, C ; 3A ).

The holotype consists of 2 corallites unequal in size growing on a fragment of biogenic rock encrusted with crustose coralline algae and bored by bivalves. The larger corallite (c 1 in Figs. 2A and 3A ) measures 1.5 cm in diameter. Septa are arranged in 5 cycles ( Fig. 2B ). The first three are complete and reach the columella, those of the fourth are less developed and those of the fifth cycle are less than ½ of the others in length. First two cycle septa are thicker than the remainder ( Figs. 2A–B ). Septa composed of multiple fan systems, and margins are dentated ( Figs. 2A–C ) ( Chevalier, 1975, Fig. 213). Septal margins and sides finely granulated ( Figs. 2C ). Part of the septa was broken when tissue was sampled for genetic analysis ( Fig. 2B at the bottom of the corallite). Columella well-developed and formed by trabecular processes from the inner margins of septa and papillae ( Fig. 2B ). The smaller corallite (c 2 in Figs. 2A , 3A ), still in the process of budding, is oriented 45 ° in relation to the calice surface plane of c1. At the time of collection the polyps were reddish- brown ( Fig. 3A ).

Paratypes. ( IRD HS3100 ), Banc de Touho , ST 1466, CC4, coll. F. Benzoni & B.W. Hoeksema, 15 April 2012 ( Fig. 2E ); ( RMNH Coel 40091), Brunei, Porter Patch, (04°53.55'N; 114°24.14'E), coll. B.W. Hoeksema, 28 April 2011 ( Figs. 1M , 2D ); ( UBDM.6.00002), E Littledale Shoal, (05°06.11'N; 114°46.00'E), coll. B.W. Hoeksema, 27 April 2011 ( Fig. 2F ); ( UBDM.6.00003), Hornet rock, (05°01.23'N; 114°43.90'E), coll. B.W. Hoeksema, 25 April 2011 ( Fig. 2H ) .

Paratypes RMNH Coel 40091 and UBDM.6.00003 are made of more than one corallite ( Figs. 2D and H , respectively), while IRD HS3100 and UBDM.6.00002 are made of one corallite ( Figs. 2E and F , respectively). Both colonial coralla clearly show the extratentacular budding process. In all paratypes at least 6 cycles of dentated septa with finely granulated margins, and well-developed columella. Paratype RMNH Coel 40091 is made of 7 corallites, rather irregular in outline and 2–2.5 cm in diameter ( Fig. 2D ). The columella is well-developed and with the typical fusion of trabecular processes and papillae forming a lamellar structure giving the columella a bilateral fashion. The columella is finely granulated like the septal sides. Paratype UBDM.6.00003 is the fragment of a larger colony, and includes two complete corallites irregular in outline. In this specimen, septa of the first cycle are thicker than the remainder ( Fig. 2H ). Paratype IRD HS3100 is the largest corallite known in the type specimens for this species which is 2.8 cm in diameter ( Fig. 2E ). There are 7 cycles of septa with those of the first two cycles slightly thicker than the others. Part of the specimen broken when tissue was sampled for genetic analysis. The single corallite of paratype RMNH Coel 40092 was part of a corallum comprising at least two more corallites small parts of which are visible at the bottom of the specimen ( Fig. 2F ). Septa in this specimen are thinner septa than in any other examined specimen. These are arranged in 6 cycles with those of the first two cycles slightly thicker than the others.

Other material. New Caledonia ( IRD HS3000 ), Canal Woodin, ST332, CC4, coll. F. Benzoni & B.W. Hoeksema, 25 April 2012 ( Fig. 8E ); ( IRD HS3263 ) ST1481, CC4, coll. B. Hoeksema, 23 April 2012 ( Fig. 3B ) ; Papua New Guinea ( UNIMIB PFB193 ), Sinub Island, (05°7.776'S; 145°48.804'E), coll. F. Benzoni, 15 November 2012; ( UNIMIB PFB241 ), Wonad Island , (05°8.16'S; 145°49.194'E), coll. F. Benzoni, 17 November 2012 ; Indonesia ( RMNH Coel. 33351), Bone Tambung, Spermonde Archipelago , South Sulawesi, (05°02'S; 119°16'E), coll. B.W. Hoeksema, 18 June 1997; ( MTQ G 60836), Selat Namatote , West Papua, (03°48.9'S; 133°55.6'E), coll. E. Turak, 21 April 2006, 22– 29 m ; Australia ( MTQ G 42939), Bullumbooroo Bay, Great Palm Islands , (18°46'S; 146°34'E), 2–15 m ; Japan ( MTQ AIMS Collection unregistered), Kushimoto, coll. J.E.N. Veron; ( MTQ AIMS Collection unregistered), Tosashimizu , coll. J.E.N. Veron ; Philippines ( MTQ AIMS Collection unregistered), coll. J.E.N. Veron ; Vietnam ( MTQ AIMS Collection unregistered), coll. J.E.N. Veron ; Brunei ( UNIMIB BLA01 ), Abana Rock (05°06.48'N; 115°04.22'E), coll. B.W. Hoeksema, 23 April 2011; ( RMNH Coel 40092), Hornet Rock (05°01.23'N; 114°43.90'E), coll. B.W. Hoeksema, 25 April 2011 ; Malaysia: North Sabah, Banggi Islands, TMPE, coll. B.W. Hoeksema ( RMNH Coel. 40108), Sta. TMP36, Patanunan Island , (07°05.995'N; 117°05.3517'E), 19 September 2012, 8– 29 m; ( RMNH Coel. 40109), Sta. TMP41, Kalang, (06°59.8017'N; 117°03.2233'E), 18 September 2012, 10 m; ( RMNH Coel. 40110), Sta. TMP16, SE Banggi Dangers, N Sibaliu, (07°11.5567'N; 117°23.6333'E), 11 September 2012, 15– 16 m; ( RMNH Coel. 40112), Sta. TMP37, Molleangan Besar Is., (07°05.12'N; 117°03.5633'E), 19 September 2012, 8 m; ( RMNH Coel. 40113), Sta. TMP15, SE Banggi Dangers, N Sibaliu, (07°12.6917'N; 117°28.2283'E), 12 September 2012, 15 m; ( RMNH Coel. 40114), Sta. TMP13, NW Tanjung Island , (07°05.6183'N; 117°16.13'E), 11 September 2012, 16 m; ( RMNH Coel. 40115), Sta. TMP27, SW Mangsee Great Reef, (07°27.4133'N; 117°13.36'E), 22 September 2012, 15 m; ( RMNH Coel. 40116), Sta. TMP38, W Carrington Reef, (07°07.8233'N; 117°13.6983'E), 20 September 2012, 23 m; ( RMNH Coel. 40117), Sta. TMP37, Molleangan Besar Island , (07°05.12'N; 117°03.5633'E), 19 September 2012, 8 m; ( RMNH Coel. 40118), Sta. TMP18, SW Bankanwan Reef, (07°11.3633'N; 117°17.6567'E), 12 September 2012, 14 m ; Malaysia: Peninsular Malaysia, east coast, Tioman Island, coll. B.W. Hoeksema ( RMNH Coel. 41517, 2 specimens) Sta. TIO-7, North Point (02°53'36"N; 104°09'26"E), 18 June 2013, 27– 30 m: ( RMNH Coel. 41518, colony 10 cm wide, largest calice 30 mm wide) Sta. TIO-12, east side, Tanjung Semanjin (02°48'41"N; 104°12'36"E), 19 June 2013, 18 m; ( RMNH Coel. 41519, 2 specimens), Sta. TIO-16, southeast point, Tanjung Asah (02°43'13"N; 104°12'53ʺE), 22 June 2013, 20– 25 m.

Skeletal variation. Overall, all examined coralla of B. vivida , new species, are formed by a variable though small number of corallites ranging between 1.5 and 2.8 mm (largest diameter). In some specimens, corallites, especially if newly formed, have septa of different cycles with the same thickness ( Fig. 2E, F ). However, in other specimens the first cycle can be visibly thicker and more exsert ( Figs. 2A–C, G, H ). Budding is mostly observed to happen sequentially ( Figs. 2D, G ). In some cases it can occur simultaneously around the first, and largest, corallite ( Fig. 3D ). Although the described bilateral fashion of the columella is very frequently observed, in some corallites only the trabecular processes can be observed ( Figs. 2B, E, H ). A number of colonial specimens have rounder and more regular corallites than the ones showed in Figure 2 (e.g., RMNH Coel. 33351).

Field characteristics. Blastomussa vivida , new species, can be solitary or form small colonies up to 10 cm long. The most striking characteristics in the field are the fleshy polyps and their colouration, which is usually bright orange to red ( Fig. 3C–E ), but can also be green to brown with contrasting oral discs ( Fig. 3F ), or nearly black ( Fig. 3B ). Mantle vesicles are well developed and rugged. This species lives in semi-protected to protected environments and it was often observed to grow on hard substrate covered in sediment.

Etymology. This species is named vivida (Latin, vividus = lively) after the typically flashy bright-coloured polyps.

Affinities. Blastomussa vivida , new species, is morphologically distinct from any other described species in the genus based on the combination of different characters including the primarily ceriod corallite organisation, large corallite size and high number of septa (the largest in the genus), and the development of a columella with a lamellar structure ( Table 2). This species has so far been confused with B. wellsi ( Veron & Pichon, 1980; Hoeksema & van Ofwegen, 2004; Wallace et al., 2009; Turak & DeVantier, 2011). The two species share the strongly dentated septa and a well-developed columella often attaining the above described bilateral structure. However, B. vivida can easily be told apart from B. wellsi , which has smaller corallites, a lower number of septa cycles, and a phaceloid corallite arrangement ( Table 2). Geographic distribution. The material listed in this study represents the known distribution of Blastomussa vivida , new species, which spans from New Caledonia and Australia in the south, Peninsular Malaysia in the west, and Japan in the north, including various localities in the Coral Triangle (for definition see Hoeksema, 2007; Veron et al. 2009).

Figures

Fig. 1. Morphology of examined specimens, Blastomussa merleti: A, top and lateral view of typically phaceloid corallites (IRD HS3264); B, side view of a corallite of the same specimen; C, detail of a corallite (IRD HS1686). B. wellsi: D, corallite arrangement (IRD HS3011); E, side view of a corallite; F, columella of the same specimen. Morphology of examined specimens, B. loyae: G, cerioid corallites (UN- IMIB DJ050); H, lateral view of the colony surface showing exsert septa devoid of dentation in the same specimen as in G; I, detail of a corallite B. omanensis: J, polygonal corallites showing the typical “groove and tubercule” appearance (UNIMIB MU094); K, side view of the same specimen as in J; L, detail of a corallite. Morphology of examined specimens, B. vivida, new species: M) ceriod corallites of a paratype (RMNH Coel. 40091; same specimen as in Fig. 2D); N, side view of septa of specimen IRD HS3000; O, columella of the same specimen. Scale bars A, D, G, J, M = 1 cm; B, C, E, F, H, I, K, L, N, O = 5 mm. Numbers 1–6 in front of the septa in C, E, I, L, and M indicate their cycle number.

Fig. 6. A, Corallum; and B, detail of corallites of the holotype of Blastomussa omanensis (BMNH 1991.6.4.150) collected by C. Sheppard in Oman (Sheppard & Sheppard, 1991: Fig. 147); C, neotype of Parasimplastrea sheppardi (MTQ G 55860); and D, close up of corallites; E, paratype of Blastomussa loyae (ZMA 8322); and F, close up of its corallites. Scale bars A, C, E = 1 cm; B, D, F = 5 mm.

Fig. 7. Skeleton macro-morphology of the examined taxa:A, Blastomussa merleti, holotype (USNM 45390); B, B. loyae, holotype (BMNH 1977.5.5.1); C, B. omanensis, specimen collected by C. Sheppard in Oman (USNM 81272); D, B. wellsi, holotype (ZMA 6905); E, B. vivida, new species, (RMNH Coel. 40092); F, Nemenzophyllia turbida, Philippines (RMNH Coel. 24246); G, Physogyra lichtensteini (UNIMIB MY007), Mayotte Island; H, Plerogyra sinuosa (UNIMIB MY006), Mayotte Island. Scale bars A, B, C = 5 mm; D–H = 1 cm.

Fig. 8. In situ images of the examined taxa: A, Blastomussa merleti, New Caledonia, ST117; B, B. loyae, Djibouti; C, B. omanensis, Yemen; D, B. wellsi, New Caledonia, ST1084; E, B. vivida, new species, New Caledonia (IRD HS3000), ST332; F, Nemenzophyllia turbida, Semporna, Malaysia; G, Physogyra lichtensteini, New Caledonia, ST1477; H, Plerogyra sinuosa, New Caledonia, ST1461. Scale bars = 1 cm

Fig. 9. Close up of the polyps showing mantle vesicles in the examined taxa: A, Blastomussa merleti, New Caledonia, ST117; B, B. loyae, Djibouti; C, B. omanensis, Yemen; D, B. wellsi, New Caledonia, ST1084; E, B. vivida new species, New Caledonia, ST332; F, Nemenzophyllia turbida, Semporna, Malaysia; G, Physogyra lichtensteini, New Caledonia, ST1477; H, Plerogyra sinuosa, New Caledonia, ST1461. White arrows indicate mantle vesicle.

Fig. 5. Phylogenetic tree of rDNA (spanning the entire ITS1, 5.8S, ITS2 and a portion of 28S and 18S) reconstructed with Bayesian Inference. Numbers at each node show percentages of Bayesian posterior probability (>70%) and MP bootstrap (>50%); – = no support. Filled circles indicate well-supported clades (bootstrap values ≥99 and posterior probability of 100).

Fig. 2. Blastomussa vivida, new species: A, the holotype (MNHN IK 2012 14226) (c1 and c2 are the larger and the smaller corallite, respectively); B, top view of corallite c1 of the same specimen, numbers 1–5 in front of the septa their cycle number; C, detail of the same corallite as in A and B showing the dentation and granulation of the septa; D, paratype (RMNH Coel. 40091); E, paratype (IRD HS3100); F, UBDM 6.0003; G, RMNH Coel. 40092; H, UBDM 6.0002. Scale bar = 1 cm

Fig. 3. Blastomussa vivida, new species in situ: A, holotype (MNHN IK 2012 14226) from New Caledonia (same specimen as in Figs. 2A–C); B, specimen IRD HS3100 from New Caledonia; C, colony from Kota Kinabalu, Malaysia showing the typically fleshy and bright coloured polyps; D, the same colony as in C with partially retracted polyps showing cerioid arrangement; E, specimen RMNH Coel. 40092 from Brunei (same specimen as in Fig. 2G); F, from Cebu, the Philippines. Scale bars = 1 cm

Fig. 4. Phylogenetic tree of mitochondrial gene COI reconstructed with Bayesian Inference. Numbers at each node show percentages of Bayesian posterior probability (>70%) and MP bootstrap (>50%); – = no support. Filled circles indicate well-supported clades (bootstrap values ≥99 and posterior probability of 100).