The genus Litophyton Forskal, 1775 (Octocorallia, Alcyonacea, Nephtheidae) in the Red Sea and the western Indian Ocean Author van Ofwegen, Leen P. text ZooKeys 2016 567 1 128 http://dx.doi.org/10.3897/zookeys.567.7212 journal article http://dx.doi.org/10.3897/zookeys.567.7212 1313-2970-567-1 6C7EADF3055D4219909EE37D218171FD Taxon classification Animalia Alcyonacea Nephtheidae Litophyton? savignyi (Ehrenberg, 1834) Figures 1A, 50, 51, 52, 53, 54, 55, 56, 57, 58 Nephthya Savignyi Ehrenberg, 1834: 284 (Red Sea). Spongodes Savignyi ; Klunzinger 1877 : 35 (Koseir). Dendronephthya savignyi ; Kuekenthal 1905 : 528. (Red Sea, Koseir, Tor); McFadden et al. 2011 : 25. Spongodes albida Holm, 1894: 30 (Red Sea, Gulf of Suez). Nephthya albida ; Kuekenthal 1903 : 160; Thomson and Mcqueen 1908 : 59. Litophyton? savignyi Not Nephthya albida ; Thomson and Dean 1931 : 82 (Indonesia). Litophyton? savignyi Not Nephthea albida ; Roxas 1933 : 413; Verseveldt 1966 : 9 (Indonesia); Tixier-Durivault 1970b : 298 (New Caledonia); Verseveldt 1974a : 96 (New Caledonia, listed); 1976: 499; 1977c : 175 (John Brewer Reef, Townsville, Qld, Australia; listed only); 1978: 50 (Pacific); Imahara 1991: 74 (Kerama Islands, Ryukyu Isl., Japan); 1996 : 25 (listed); Ofwegen 1996 : 209 (Papua New Guinea). Nephthya jaegerskioeldi Holm, 1904: 6 (Red Sea, Tor). Nephthea aberrans Verseveldt, 1968: 54 (Tany Kely, near Nosy Be , Madagascar); 1973 : 96 (Tany Kely, near Nosy Be , Madagascar; re-description). Nephthea tixierae Verseveldt, 1968: 55 (Nosy Ovy, Radama Is., Madagascar); 1973 : 94 (Nosy Ovy, Radama Is., Madagascar; re-description); Tixier-Durivault 1972 : 26 (Madagascar). Material examined. NHMW 2407, 2 specimens, Red Sea, Tor, Frauenfeld; UUZM 417, type Nephthya jaegerskioeldi , Red Sea, Tor, depth 0.5-0.65 m; RMNH Coel. 3906, Nephthea aberrans , holotype, Tany Kely, Madagascar, depth 10 m; RMNH Coel. 3907, Nephthea tixierae , holotype, Nosy Ovy, depth 8 m; RMNH Coel. 8044, paratype, Nosy Ovy, depth 8 m, 9; RMNH Coel. 42087, Egypt, Safaga, sample 44, 5 April 1997, coll. G. Reinicke; ZMTAU Co 25685 1965, Red Sea, South tip Sinai, Shab Mahmud, depth 0-21 m, 12 July 1987, coll. Y. Benayahu; ZMTAU Co 25688 1971, Red Sea, South tip Sinai, Shab Mahmud, 12 July 1987, coll. Y. Benayahu; ZMTAU Co 25690 1979, Red Sea, South tip Sinai, Shab Mahmud, 12 July 1987, coll. Y. Benayahu; ZMTAU Co 25825 1489, Red Sea, South tip Sinai, Shab Mahmud, depth 20-30 m, 9 July 1986, coll. Y. Benayahu; ZMTAU Co 25829 1523, Red Sea, South tip Sinai, Shab Mahmud, depth 20-30 m, 9 July 1986, coll. Y. Benayahu; ZMTAU Co 25833 1606, Red Sea, Gulf of Suez, Shag Rock, depth 0-20 m, 10 July 1986, coll. Y. Benayahu; ZMTAU Co 25840 1782, Red Sea, South tip Sinai, Tiran St., 13 July 1986, coll. Y. Benayahu; ZMTAU Co 25861 2048, Red Sea, Gulf of Suez, Shag Rock, depth 3-24 m, 14 July 1987, coll. Y. Benayahu; ZMTAU Co 25878 1625, Red Sea, Gulf of Suez, Shag Rock, depth 0-20 m, 10 July 1986, coll. Y. Benayahu; ZMTAU Co 25983 2618, Red Sea, Tiran Straits, Gordon andThomas reef, depth 12-16 m, 27 March 1988, coll. Y. Benayahu; ZMTAU Co 26061 (2748), Red Sea, Gulf of Suez, Shag Rock, 6 October 1988, coll. Y. Benayahu; ZMTAU Co 26062 (2750), Red Sea, Gulf of Suez, Shag Rock, depth 10 m, 6 October 1988, coll. Y. Benayahu; ZMTAU Co 26065 (2788), two specimens, Red Sea, Gulf of Suez, Shag Rock, 7 October 1988, coll. Y. Benayahu; ZMTAU Co 26068 (2830), Red Sea, Gulf of Suez, Shag Rock, depth 30 m, 7 October 1988, coll. Y. Benayahu; ZMTAU Co 26115 (3149), Red Sea, Gulf of Suez, Tawilla Is., 24 September 1989, coll. Y. Benayahu; ZMTAU Co 26131 (3236), Red Sea, Gulf of Suez, near Shaduan Is., 26 September 1989, coll. Y. Benayahu; ZMTAU Co 26189, Red Sea, Gulf of Aqaba, Ras Mamlakh, coll. Y. Benayahu, 12 March 1981; ZMTAU Co 26198 (1046), Red Sea, Gulf of Aqaba, Dahab southern oasis, depth 8 m, 4 November 1981, coll. Y. Benayahu; ZMTAU Co 26205 (1077), Red Sea, S tip of Sinai, S of Naama, "Amphores" , depth 16-20 m, 7. November 1981, coll. Y. Benayahu; ZMTAU 26208 (1088), Red Sea, Gulf of Aqaba, El Goz, depth 2-5 m, 8 November 1981, coll. Y. Benayahu; ZMTAU Co 26210 (1089), Red Sea, Gulf of Aqaba, Shurat el Manqata, depth 3-6 m, 9 November 1981, coll. Y. Benayahu; ZMTAU Co 26212 (1091), Red Sea, Gulf of Aqaba, Shurat el Manqata reef flat, 9 November 1981, coll. Y. Benayahu; ZMTAU Co 26214 (1097), two specimens, Red Sea, Gulf of Aqaba, Naqeb Shahin, depth 25 m, 29 November 1981, coll. Y. Benayahu; ZMTAU Co 26217 (1114), South tip Sinai, Sharm a Sheikh. "Amphores" , depth 20 m, 30 November 1981, coll. Y. Benayahu; ZMTAU Co 26219 (1118), Red Sea, S tip of Sinai, Naama garden, depth 30 m, 30 November 1981, coll. Y. Benayahu; ZMTAU 26233, Red Sea, Tiran Is., Favel Bay lagoon, depth 1-2 m, 22 September 1981, coll. Kerman; ZMTAU Co 26235, Red Sea, Gulf of Suez, Ras Tanaka, 25 September 1974, coll. Y. Benayahu; ZMTAU Co 26245 (327), Red Sea, Gulf of Aqaba, Taba km 179, depth 25 m, 9 October 1977, coll. Y. Benayahu; ZMTAU Co 26247 (350), Red Sea, Gulf of Suez, Sheikh Riach, depth 5 m, 18 November 1977, coll. Y. Benayahu; ZMTAU Co 26248 (389), Red Sea, Gulf of Suez, Sheikh Riach, depth 5 m, 18 November 1977, coll. Y. Benayahu; ZMTAU 26250 (446), Red Sea, Gulf of Suez, Ras Gahra, depth 1 m, 29 November 1977, coll. Y. Benayahu; ZMTAU Co 26253 (470), Red Sea, South tip Sinai, Marsa Khadamia, depth 30 m, 22 November 1977, coll. Y. Benayahu; ZMTAU Co 26254 (17923), Red Sea, coll. Y. Benayahu; ZMTAU NS 8295, Red Sea, Gulf of Suez, Ras Kanisa, 20 October 1971, coll. Fishelson; ZMTAU Co 30064, Eritrea, Heteroxenia bed near Eucus island, 15°53.884'N , 39°53.141'E , depth 3.5 m, 29 April 1997, coll. Y. Benayahu; ZMTAU Co 32965, Eritrea, Dahlak Archipelago, Shumma Is., 15°32.00'N , 40°00.00'E , depth 8-12 m, 16 February 2005, coll. Y. Benayahu; ZMTAU Co 34205-34207, Red Sea, Gulf of Aqaba, North Oil Jetty Elat, 29°31.41'N , 34°56.14'E , depth 15.2 m, 26 July 2007, coll. Y Benayahu; ZMTAU Co 34066-34067, Red Sea, Gulf of Aqaba, Elat, 29°30.14'N , 34°55.075'E , depth 18.3-22.9 m, 23 July 2007, coll. Y. Benayahu. Removed from the species. RMNH Coel. 8956, Red Sea, Gulf of Aqaba, Fara 'un Isl., 7 January 1968, coll. Hebrew Univ.- Smiths. Red Sea project 65/SLR 1204 (identified as Nephthea albida = Litophyton chabrolii ). Diagnosis. Litophyton where the polyps have a projecting supporting bundle and make an acute angle with the polyp stalk. Description of NHMW 2407. The colony is 3 cm high and wide, the colony stalk 2 cm high (Figure 50B). Polyps up to about 0.6 mm wide and high (Figure 52A). Supporting bundle projecting up to 0.7 mm, composed of 2-4 spindles (Figure 52A). These spindles are up to 3 mm long, with spines and projecting smooth tip (Figure 52D). Sclerites in polyp are irregularly distributed. Abaxial side of the polyp with curved spindles with spines or simple tubercles, up to 0.6 mm long, several with one smooth end (Figure 52B). Laterally less tuberculated spindles are present, up to 0.2 mm long (Figure 52B). Adaxially and in the tentacles flattened rodlets and ovals are present, up to 0.1 mm long (Figure 52C). The adaxial side of the polyp stalk has small rodlets, up to 0.05 mm long (Figure 52E). The amount of these rodlets varies per polyp, sometimes only a few are present (Figure 52A), others have the whole polyp stalk closely packed with them. Surface layer top of stalk. Spindles with simple tubercles, up to 2.5 mm long, some slightly unilaterally spinose. Surface layer base of stalk. Spindles and unilaterally spinose spindles with simple tubercles, shorter than in the top of the stalk, up to 1.5 mm long. Furthermore small rodlets, several unilateral spinose; smaller branched spindles, radiates and derivatives of these (Figure 53A). Interior base of stalk. The larger interior spindles are not very different from the surface ones, only slightly less tuberculate (Figure 52F). They are up to 1.5 mm long. Smaller, branched bodies also occur (Figure 53B). Colour. Colony is white. Distribution. Red Sea, Indian Ocean. Variability. Most colonies examined have slender branches and resemble species of Stereonephthya (Figure 51 A-B ); a few are more " Litophyton -like" (Figure 51C). Remarks. After the very short original description of Nephthya savignyi by Ehrenberg (1834 : 60), Klunzinger (1877 : 35) identified a specimen from Koseir (Red Sea), as Spongodes (= Dendronephthya ) savignyi . Kuekenthal (1905 : 529) examined many specimens, 20 all together, including two specimens from the Berlin museum, one of them Ehrenberg's, "originalexemplar" , and one from the Stuttgart museum, Klunzinger's, specimen. Kuekenthal mentioned little variability in all specimens examined and he also synonymized Holm's, Nephthya jaegerskioeldi and Nephthea jaegerskioeldi var. microspina with Ehrenberg's, Dendronephthya savignyi , based on the presence of polyps in bundles. One of the type specimens of Nephthea jaegerskioeldi has been re-examined; the colony and sclerites are presented in Figures 50D, 54-55, and I agree with Kuekenthal that the species should be synonymized with Litophyton? savignyi . During my visit to the Berlin museum I was unable to find Ehrenberg's, specimen, later on Dr. Goetz Reinicke was so kind to provide me with photographs of a specimen that could be that particular one, though with a question mark (Figure 50A). Indeed doubts remain about the status of the Berlin specimen since it is almost 12 cm wide, while Kuekenthal mentioned its width to be 8.5 cm. The specimen described above is from the Vienna Museum (NHMW 2407) (Figures 50 B-C , 52-53), and is probably one of the specimens examined by Kuekenthal , as it was found at Tor (Red Sea) and Kuekenthal (1905 : 531) also examined material from that locality. Sclerites of ZMTAU 26245 (Figure 51C) are presented to show their variation (Figures 56-58). Although not re-examined I consider Spongodes albida Holm, 1894 synonymous with Litophyton? savignyi . The specimen is only a few cm long but features all the characters of Litophyton? savignyi , i.e. projecting supporting bundle, many small rodlets in the polyp stalk and large interior spindles. In the Red Sea Litophyton? savignyi differs from all other Litophyton species in having polyps with a protruding supporting bundle giving the colony a prickly appearance. The polyps also make an acute angle with the stalk as is seen in the genus Stereonephthya . It can only be confused with two species of Stereonephthya , Stereonephthya acaulis Verseveldt, 1973, and Stereonephthya cundabiluensis Verseveldt, 1965. The latter always contains coloured sclerites but Stereonephthya acaulis can have white colonies with colourless sclerites ( Verseveldt 1973 : 153). But it differs from Litophyton savignyi in lacking oval tentacle sclerites, having differently shaped polyp stalk rodlets ( Stereonephthya -type; Figs 13a, 14a, 15a in Ofwegen and Groenenberg (2007) ), and having much smaller (up to 0.75 mm long), branched, less tuberculate, internal stalk spindles; see Verseveldt (1973 : figs 33n, o). ZMTAU Co 34066, identified by me as Dendronephthya savignyi , was used in a molecular study by McFadden et al (2011) . In that study it grouped with other Litophyton species, i.e. Litophyton striatum ( Kuekenthal , 1903) (identified by me as Litophyton elatensis (Verseveldt & Cohen, 1971), Litophyton acuticonicum Verseveldt, 1974, and Litophyton acutifolium Verseveldt, 1974 (= Litophyton viridis ) rather than with other Dendronephthya species included in the study. Unfortunately, Utinomi (1954a) designated Litophyton savignyi as the type species for Dendronephthya , a genus with more than 250 nominal species. Following strict nomenclatural priority would cause widespread confusion within nephtheid taxonomy. To avoid changing of generic combinations and the confusion that it would cause, a case will be submitted to the International Commission on Zoological Nomenclature (ICZN) to preserve the name Dendronephthya , in the meanwhile the species will be cited as Litophyton? savignyi . The two Nephthea species with projecting supporting bundle described by Verseveldt (1968) from Madagascar, Nephthea aberrans and Nephthea tixierae , I regard synonymous with Litophyton? savignyi . I consider the reported differences to represent intra-specific variation.