Research Article |
Corresponding author: Leen P. van Ofwegen ( ofwegen@naturalis.nnm.nl ) Academic editor: Bert W. Hoeksema
© 2016 Leen P. van Ofwegen.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
van Ofwegen LP (2016) The genus Litophyton Forskål, 1775 (Octocorallia, Alcyonacea, Nephtheidae) in the Red Sea and the western Indian Ocean. ZooKeys 567: 1-128. https://doi.org/10.3897/zookeys.567.7212
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The Litophyton species of the Red Sea and the western Indian Ocean are revised, which includes species previously belonging to the genus Nephthea, which is synonymized with Litophyton. A neotype for both Litophyton arboreum, the type species of Litophyton, and Nephthea chabrolii, the type species of Nephthea, are designated. The new species L. curvum sp. n. is described and depicted, and a key to all Litophyton species is provided. Of the 26 species previously described from the western Indian Ocean and Red Sea, 13 species are considered valid and 13 have been synonymized or placed in other genera.
Cnidaria , Anthozoa , Alcyonacea , Nephthea , revision, new species
This is the second in a series of publications in which nephtheid genera are revised. The first (
Nephthya savignyi Ehrenberg, 1834 is at present the type species of the genus Dendronephthya, as designated by
Examination of the types of most species of Litophyton and Nephthea and examination of many recently collected specimens proved that species of this genus have restricted distributions and that therefore the revision can be split based on different geographic regions. The 17 nominal species of Litophyton (van
NBC
ZMTAU
For the technical terms used in the descriptions, see the glossary compiled by
Four permanent microscope slides have been made for each specimen examined, which are kept at
Sclerite drawings have been made using the permanent microscope slides. As most old museum specimens examined contained a large amount of broken sclerites, SEM images were produced of recently collected material.
All the Nephthea and Litophyton type specimens available have been re-examined. In addition, more recent material from the
Litophyton Forskål, 1775: 139.
Ammothea Lamarck, 1816: 410.
Nephthee Savigny, 1817: pl. 2 fig. 5 (plates of the text of Andouin)
Nephthea Audouin, 1828: 49.
Nephthya Ehrenberg, 1834: 284.
Neptaea Blainville, 1834: 523.
Nephtya Van Beneden, 1867: 197.
Amicella Gray, 1869: 123.
Verrilliana Gray, 1869: 130.
Litophytum Kükenthal, 1903: 106.
Nephtheids with bushy and arborescent colonies. Polyps clustered at the end of the terminal branches, forming catkins. Polyps non-retractile, without or with supporting bundle, sometimes completely unarmed. Sclerites of surface layer of branches, stem and stalk are spindles and unilateral spinose spindles, the colony stalk also contains capstans and derivatives of capstans. Interior of the stalk has sparsely tuberculated spindles. Colonies zooxantellate.
Litophyton arboreum Forskål, 1775, by monotypy.
Because of the synonymy of Nephthea with Litophyton, for many species a spelling emendation needed to be made to comply with ICZN Art. 31.2 in relation to gender agreement between generic and species names.
Litophyton species are known to have extreme intraspecific variation in colony shape and sclerites (
Colony shape did not prove to provide a reliably constant character. A good example is L. ? savignyi, which may resemble some other Litophyton species but can also have a colony shape like that seen in some species of Stereonephthya (Figures
The polyp armature showed some useful characters but some sclerite arrangements were observed in various species: L. chabrolii (Figure
The sclerites of Litophyton species show a staggering morphological variation, with those in the polyp and stem and stalk surfaces varying most in shape. Notably, the shape of the stalk surface sclerites is different depending on the height on the stalk. The least variable sclerites are the spindles of the internal canals. As with the polyp armature some species have the same types of spindles, which limits the usefulness of these sclerites.
L. acuticonicum (Verseveldt, 1974), Red Sea
L. arboreum Forskål, 1775 (type lost), Red Sea and Socotra
L. bumastum (Verseveldt, 1973), Madagascar
L. chabrolii (Andouin, 1828) (type lost), Red Sea
L. curvum sp. n., Red Sea
L. filamentosum (Verseveldt, 1973), Madagascar
L. laevis (Kükenthal, 1913), Red Sea
L. lanternarium (Verseveldt, 1973), Madagascar
L. maldivensis (Hickson, 1905), Red Sea and Maldives
L. ? savignyi (Ehrenberg, 1834), Red Sea and Madagascar
L. simulatum (Verseveldt, 1970), Red Sea, Socotra and Chagos Archipelago
L. striatum (Kükenthal, 1903), Red Sea, Socotra, Chagos Archipelago and Madagascar
L. viridis (May, 1898), Red Sea and East Africa
Nephthea aberrans Verseveldt, 1968 = L. savignyi
Spongodes albida Holm, 1894 = L. savignyi
Nephthya armata Thomson & Henderson, 1906 = Stereonephthya
BM 1912.2.25.12 (holotype) and BM 1933.3.13.148 (holotype fragment) were re-examined. Despite disintegrated sclerites, it was obvious that the species belongs to Stereonephthya
Nephthea galbuloides Verseveldt, 1973 = L. striatum
Nephthea tixierae Verseveldt, 1968 = L. savignyi
Litophyton acutifolium Kükenthal, 1913 = L. viridis
Litophyton crosslandi Thomson & McQueen, 1908 = L. viridis
Nephthea elatensis Verseveldt & Cohen, 1971= L. striatum
Nephthea hirsuta Tixier-Durivault, 1966 = Stereonephthya
Nephthya jaegerskioeldi Holm, 1904 = L. savignyi
Ammothea sanderi May, 1899 = L. viridis
Ammothea stuhlmanni May, 1898 = L. viridis
Nephthya zanzibarensis Thomson & Henderson, 1906 = Stereonephthya
BM 1933.3.13.150 (part of syntype) was re-examined. Despite disintegrated sclerites it was obvious that the species belongs to Stereonephthya
1 | Supporting bundle projecting | L. savignyi |
– | Supporting bundle not projecting | 2 |
2 | Polyp stalk with rodlets | 3 |
– | Polyp stalk without rodlets | 5 |
3 | Internal spindles of the base of the stalk mostly with blunt ends | L. arboreum |
– | Internal spindles of the base of the stalk mostly with pointed ends | 4 |
4 | Base of stalk surface sclerites straight | L. filamentosa |
– | Base of stalk surface spindles curved | L. curvum |
5 | Polyps without sclerites or mostly small rodlets | 6 |
– | Polyps with spindles | 7 |
6 | Internal spindles of the base of the stalk heavily branched. | L. viridis |
– | Internal spindles of the base of the stalk not branched, several with blunt ends | L. maldivensis |
7 | Adaxial side of polyps without sclerites | L. bumastum |
– | Adaxial side of polyps with sclerites | 8 |
8 | Internal spindles of the base of the stalk very small, most about 0.5 mm long or shorter | 9 |
– | Internal spindles of the base of the stalk at least 1.0 mm long | 10 |
9 | Internal spindles of the base of the stalk slender, up to 0.15 mm wide, few in number | L. laevis |
– | Internal spindles of the base of the stalk wide, up to 0.25 mm wide, many present; spindles with blunt ends can be present | L. simulatum |
10 | Larger internal spindles of the base of the stalk heavily branched | L. striatum |
– | Larger internal spindles of the base of the stalk not branched | 11 |
11 | Internal spindles of the base of the stalk up to 2.0 mm long | L. acuticonicum |
– | Internal spindles of the base of the stalk up to about 1.0 mm long | 12 |
12 | Internal spindles of the base of the stalk mostly unbranched, with regular tuberculation | L. chabrolii |
– | Internal spindles of the base of the stalk often branched, with irregular tuberculation | L. lanternarium |
Nephthea acuticonica Verseveldt, 1974b: 28, figs 20–21, pl. 9 (El Kura’, Gulf of Aqaba, Red Sea);
Litophyton acutifolium;
Not Litophyton acutifolium Kükenthal, 1913: 12 (= L. viridis).
Nephthea striata;
Litophyton with branched spindles in the surface layer of the base of the stalk and large internal spindles in the base of the stalk, up to 2.0 mm long, the largest not branched.
Red Sea: Gulf of Suez, Gulf of Aqaba.
The species is sufficiently described by
The species is characterized by the presence of branched spindles of the surface layer of the base of the stalk and the presence of large internal spindles in the base of the stalk, sometimes with blunt ends or branched. L. arboreum also has internal spindles with blunt ends, but here the surface spindles of the base of the stalk are never branched. L. simulatum can also have spindles with blunt ends but these are always less than 0.5 mm long, while in L. acuticonicum they are up to 1.5 mm long. Sometimes also spindles with side branches are present in the interior of the base of the stalk (ZMTAU 26239, Figure
Litophyton arboreum Forskål, 1775: 139 (Red Sea);
Ammothea virescens Lamarck, 1816: 411; Savigny 1817: pl. 2 fig. 6;
Nephthea cordierii Audouin, 1828: 48 (Savigny's, pl. 2 fig. 6).
Neptaea inominata Blainville, 1830: 487 (Savigny's, pl. 2 fig. 6);
Ammothea arborea Klunzinger, 1877: 31, pl. 2 fig. 4 (Red Sea);
Litophytum arboreum;
Litophyton viride;
Not Litophyton arboreum;
ZMTAU Co 26246, neotype, Red Sea, Gulf of Aqaba Eilat Marine lab, 12 m depth, 20 March 1978, coll. Y. Benayahu;
Litophyton with many internal spindles of the base of the stalk with blunt ends. The polyp stalk with scales.
The neotype is 5 cm high and 7.5 cm wide; the colony stalk is 2–3 cm high (Figure
The polyps (Figure
Surface layer top of stalk. Spindles, radiates, and derivatives of these, merging into unilaterally spinose spindles; all sclerites with simple tubercles (Figures
Surface layer base of stalk. Sclerites similar to those of the top of the stalk but with longer and sharper spines (Figure
Interior base of stalk. Spindles, up to 1.2 mm long, with simple sparse tubercles (Figure
Colour. The colony is white.
Red Sea, Socotra.
The microscope slide of the stalk of ZMTAU Co 26234 only has internal sclerites of the stalk because the specimen has the surface layer missing.
Two of the 14 microscope slides of
L. arboreum is characterized by having large spindles with blunt ends in the interior of the stalk. L. acuticonicum and L. simulatum also have this type of sclerites. L. acuticonicum differs in having branched, unilaterally spinose spindles in the surface layer of the stalk, which are also twice as long as the unbranched spinose spindles of L. arboreum. L. simulatum also differs in having twice as long unilaterally spinose spindles in the surface layer of the stalk. Moreover, L. arboreum has small oval scales in the polyp stalk, a type of sclerite not present in L. acuticonicum and L. simulatum.
Nephthea bumasta Verseveldt, 1973: 98, figs 22-23 (Madagascar).
Litophyton with adaxial side of polyps without sclerites; interior stalk with pointed spindles up to 1.5 mm long; a few have blunt ends.
Madagascar.
The species is sufficiently described by
L. bumastum is the only Litophyton species in the western Indian Ocean described with the adaxial side of the polyps lacking sclerites.
Nephthea chabrolii Andouin, 1828:49 (explanation for Savigny's, “Description de l’Egypte ...”, 1817, pl. 2 fig. 5; ?
Neptaea Savignyi;
Not Nephthya Savignii;
Nephthya chabrolii;
Not Spongodes (Nephthya) chabrolii;
Not Spongodes (Nephthya) Chabrolii var. ternatana Kükenthal, 1895: 428 (Ternate, Indonesia).
Not Spongodes (Nephthya) Chabrolii var. molukkana Kükenthal, 1895: 428 (Ternate, Indonesia).
Not Nephthya chabrolii var. ternatana;
Not Nephthya chabrolii var. moluccana;
Not Nephthya chabrolii;
Not Nephthea chabrolii;
ZMTAU Co 26244, neotype, Red Sea, Gulf of Aqaba Wadi Magrash km 207, 20 July 1974, coll. Y. Benayahu (second specimen in the bottle is L. simulatum);
Litophyton with polyps with spindles. Internal spindles of the base of the stalk up to about 1.0 mm long, mostly unbranched and with very regular tuberculation.
The neotype is 4 cm high and 6.5 cm wide; the colony stalk is 1 cm high (Figure
The polyps are up to about 0.5 mm wide and high (Figure
Surface layer top of stalk. Spindles, radiates, and derivatives of these, spindles, and unilaterally spinose spindles; sclerites with simple or complex tubercles (Figure
Surface layer base of stalk. Sclerites similar to those of the top of the stalk but the unilaterally spinose sclerites with slightly longer spines (Figure
Interior base of stalk. Spindles, up to 1.2 mm long, with simple, regular, sparse tubercles (Figure
Colour. The colony is white.
Gulf of Aqaba.
The species resembles L. lanternarium and L. simulatum but differs in having mostly unbranched internal stalk spindles with very regular tuberculation.
It is noteworthy that
ZMTAU Co 28555 (E167), holotype and seven paratypes, Eritrea, Dahlak Archipelago, Dur Ridgrig, depth 8 m, 15 October 1993, coll. Y. Benayahu; paratypes: ZMTAU Co 25670 1873, Red Sea, South tip Sinai, Shab el Utaf, depth 0–20 m, 11 July 1987, coll. Y. Benayahu; ZMTAU Co 26223, Red Sea, Gulf of Aqaba, 10 km south of Dahab, 24 July 1972, coll. L. Fishelson; ZMTAU Co 26225, Red Sea, South tip Sinai Ras um Sud, 11 April 1972, coll. Y. Benayahu; ZMTAU Co 28549 (E241); Eritrea, Dahlak Archipelago, Sarad, depth 3 m, 17 October 1993, coll. Y. Benayahu; ZMTAU Co 28552 (E261), Red Sea, Dahlak Archipelago, Daliacus, depth 3 m, 18 October 1993, coll. Y. Benayahu; ZMTAU 32929, Eritrea, Dahlak Archipelago, between Nocra Is. and Dahlak Is., southern entrance to the channel, 15°41.36'N, 39°56.08'E, depth 0–5 m, 14 February 2005, coll. Y. Benayahu; ZMTAU Co 32964, Eritrea, Dahlak Archipelago, Shumma Is., 15°32.00'N, 40°00.00'E, depth 8–12 m, 16 February 2005, coll. Y. Benayahu.
Litophyton with the internal spindles of the base of the stalk mostly with pointed ends. Polyp stalk with scales. Surface layer of the stalk with straight and curved sclerites.
The flabby holotype ZMTAU Co 28555 is 5.5 cm long and wide (Figure
The polyps are up to about 0.5 mm wide and 0.6 mm high (Figure
Surface layer top of stalk. Radiates, derivatives of these, spindles and unilaterally spinose spindles (Figure
Surface layer base of stalk. Radiates, derivatives of these, spindles and unilateral spinose spindles (Figures
Interior base of stalk. Spindles, up to 1.0 mm long, with widely spaced simple tubercles (Figure
The Latin “curvum”, curve, curved object or line, refers to the curved spindles from the surface of the stalk.
Red Sea: Gulf of Aqaba, Dahlak Archipelago.
ZMTAU Co 26223, ZMTAU Co 26225 and ZMTAU Co 28552 are slightly different from the holotype. They show less compressed colony shapes (Figure
To show variation, the sclerites of ZMTAU Co 28552 are also presented (Figures
The species can be confused with L. chabrolii (Andouin, 1828), but that species has stiffer colonies, stronger polyp armature, and wider, more regular shaped internal stalk spindles. L. laevis (Kükenthal, 1913) is also similar to this species, but lacks the curved spindles and unilateral spinose spindles in the surface layer of the base of the stalk. Moreover, both these species do not have the polyp stalk scales present in L. curvum.
Nephthea filamentosa Verseveldt, 1973: 141, figs 24–25, pl. 6 (Tany Kely, near Nosy Bé, Madagascar).
Not Nephthea filamentosa;
Litophyton with the internal spindles of the base of the stalk mostly with pointed ends. Polyp stalk with scales, surface of the stalk with straight spindles and unilaterally spinose spindles.
Only known from the type locality Madagascar.
The species is sufficiently described by
The species mostly resembles Litophyton curvum but differs in having straight sclerites in the surface layer of the base of the stalk and very spiny, almost spheroidal, sclerites in the surface layer of the base of the stalk.
Nephthya laevis Kükenthal, 1913: 20, figs 9-13, pl. 2 fig. 5 (Red Sea, Jeddah).
Nephthea laevis;
Not Nephthea laevis;
Litophyton with the internal spindles of the base of the stalk short and slender, up to 0.15 mm wide and 0.5 mm long.
The holotype is 8 cm high and 5 cm wide; the short colony stalk divides in several main stems shortly above its base (Figure
The polyps are up to about 0.7 mm high and 0.6 mm wide (Figure
Surface layer top of stalk. Rods and spindles, up to 0.45 mm long, with simple tubercles (Figure
Surface layer base of stalk. Radiates and derivatives of these, up to 0.15 mm long, with simple tubercles; a few are unilaterally spinose (Figure
Interior base of stalk. Spindles, up to 0.5 mm long, with simple sparse tubercles (Figure
Colour. The colony is whitish.
Red Sea: Gulf of Suez, Dahlak Archipelago.
The species can be confused with L. simulatum, but the latter has wider, more branched internal spindles.
ZMTAU Co 26126 3211 has been used for SEM images of sclerites (Figures
Nephthea lanternaria Verseveldt, 1973: 147 (Madagascar).
Nephthea amentacea;
Not Nephthya amentacea
Litophyton with the internal spindles of the base about 1.0 mm long, often branched, with irregular distribution of tubercles.
Only known from the type locality Madagascar.
The species is sufficiently described by
The species mostly resembles L. chabrolii but differs in having internal spindles in the base of the stalk with irregular tuberculation; several spindles branched.
The specimens from Madagascar identified by Verseveldt as Nephthea amentacea (
Eunephthya maldivensis Hickson, 1905: 824, fig. 12 (Maldives, Kolumadula Atoll);
Litophyton maldivensis; Hickson 1908: 173–176.
BMNH 1962.7.20.123, syntype; BMNH 1962.7.20.124, syntype; ZMTAU Co 26249, Red Sea, Gulf of Suez, Ras Gahra, depth 2 m, 19 November 1977, coll. Y. Benayahu; ZMTAU Co 26221, Red Sea, Gulf of Suez, A-Tur, 20 September 1967, coll. L. Fishelson; ZMTAU Co 26252, Red Sea, Gulf of Suez, Ras Gahra, depth 2 m, 20 November 1977, coll. Y. Benayahu; ZMTAU Co 28548 (E262), Red Sea, Dahlak Archipelago, Daliacus; depth 3 m, 18 October 1993, coll. Y. Benayahu;
Litophyton with polyps with small rodlets. Internal spindles of the base of the stalk short, mostly unbranched, several with blunt ends.
The holotype is 3.5 cm high and 5 cm wide (Figure
The polyps have small rodlets and spindles, situated in the tentacles and both the lateral and abaxial parts of the polyp (Figure
Lobes. Surface and interior with narrow spindles up to 0.5 mm long (Figure
Surface layer top of stalk. Radiates, derivatives of these, and spindles (Figure
Surface layer base of stalk. Radiates, derivatives of these, spindles and unilateral spinose spindles (Figure
Interior base of stalk. Spindles with widely spaced simple tubercles (Figure
Maldives, Red Sea, Socotra.
The characteristics of specimen BMNH 1962.7.20.124 agree with the description of
The species can be confused with L. arboreum Forskål, 1775, as that species has also many blunt spindles in the interior of the base of the stalk. But they are longer, have more regularly spaced tubercles and do not include smaller, smoother forms. Also the polyps are more strongly armed. Litophyton maldivensis can also be confused with specimens of L. simulatum in terms of having short sclerites in the interior of the stalk. But L. simulatum, like L. arboreum, differs in lacking the smooth smaller internal spindles and having more strongly armed polyps. Moreover, it has many branched internal spindles.
SEM images of the sclerites of ZMTAU Co26249 (Figure
Nephthya Savignyi Ehrenberg, 1834: 284 (Red Sea).
Spongodes Savignyi;
Dendronephthya savignyi;
Spongodes albida Holm, 1894: 30 (Red Sea, Gulf of Suez).
Nephthya albida;
Not Nephthya albida;
Not Nephthea albida;
Nephthya jaegerskioeldi Holm, 1904: 6 (Red Sea, Tor).
Nephthea aberrans Verseveldt, 1968: 54 (Tany Kely, near Nosy Bé, Madagascar);
Nephthea tixierae Verseveldt, 1968: 55 (Nosy Ovy, Radama Is., Madagascar);
Litophyton where the polyps have a projecting supporting bundle and make an acute angle with the polyp stalk.
The colony is 3 cm high and wide, the colony stalk 2 cm high (Figure
Polyps up to about 0.6 mm wide and high (Figure
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
Interior base of stalk. The larger interior spindles are not very different from the surface ones, only slightly less tuberculate (Figure
Colour. Colony is white.
Red Sea, Indian Ocean.
Most colonies examined have slender branches and resemble species of Stereonephthya (Figure
After the very short original description of Nephthya savignyi by
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
The specimen described above is from the Vienna Museum (
Sclerites of ZMTAU 26245 (Figure
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 L. ? savignyi, i.e. projecting supporting bundle, many small rodlets in the polyp stalk and large interior spindles.
In the Red Sea L. ? 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, S. acaulis Verseveldt, 1973, and S. cundabiluensis Verseveldt, 1965. The latter always contains coloured sclerites but S. acaulis can have white colonies with colourless sclerites (
ZMTAU Co 34066, identified by me as Dendronephthya savignyi, was used in a molecular study by
Unfortunately,
The two Nephthea species with projecting supporting bundle described by
Nephthya striata (in part) Kükenthal, 1903: 166, pl. 7 fig. 12, pl. 9 fig. 60 (Red Sea).
Nephthea simulata Verseveldt, 1970: 221, figs 7–8, pl. 2 fig. 1 (Et Tur, Gulf of Suez).
Litophyton with internal spindles of the base of the stalk short, mostly up to 0.5 mm long, up to 0.25 mm wide. Spindles with blunt ends and branched spindles present.
(after
The polyps are up to about 0.65 mm wide and 0.8 mm high. Supporting bundle not projecting, composed of spindles with simple tubercles, outer side and distal end with more tubercles. Length of these spindles is up to 1.1 mm. Polyp body sclerites irregularly arranged, the smallest are present adaxially; abaxially they merge into the smaller spindles of the supporting bundle and have larger tubercles on the outer side. The tentacle sclerites resemble the smallest adaxial polyp sclerites.
Surface layer top of stalk. Spindles, radiates, and derivatives of these, merging into unilaterally spinose spindles; all sclerites with simple tubercles. The spindles are up to 0.25 mm long.
Surface layer base of stalk. Sclerites similar to those of the top of the stalk but longer, up to 0.4 mm long; the unilaterally spinose sclerites having longer spines.
Interior base of stalk. Spindles, up to 1.0 mm long, with simple sparse tubercles. Several spindles have one or more side branches, the smaller spindles are more often branched than the larger ones. A few spindles have blunt ends.
Colour. The colony is grey.
Red Sea, Socotra, Chagos Archipelago.
The microscope slides of transparent polyps made by
The species also resembles L. striatum as that species has also branched interior spindles in the base of the stalk. However, the sclerites of the interior of the base of the stalk of L. striatum always include spindles with many small side branches or extra tall tubercles, while those of L. simulatum have just one or a few.
For showing variation sclerites of ZMTAU Co 25874 (Figures
Nephthya striata (in part) Kükenthal, 1903: 166, pl. 7 fig. 12, pl. 9 fig. 60 (Red Sea);
Not Nephthea striata;
Nephthea galbuloides Verseveldt, 1973: 144, figs 26–28 (Andraikarekabé, Nosy Komba, near Nosy Bé, Madagascar; Pointe Ambarionaomby, Nosy Komba, near Nosy Bé, Madagascar; Tany Kely, near Nosy Bé, Madagascar).
Not Nephthea galbuloides;
Nephthea elatensis Verseveldt & Cohen, 1971: 53, fig. 1 (Red Sea);
SMF 1279, syntype Nephthya striata, Rotes Meer; Rüppell leg. 1832;
Litophyton with the large internal spindles of the base of the stalk at least 1.0 mm long and heavily branched.
The stiff lectotype is 7 cm long and 5 cm wide (Figure
The polyps are up to 0.80 in height and up to 0.90 in width (Figure
Surface layer top of stalk. Radiates and derivatives of these, spindles and unilaterally spinose spindles, which are up to 0.6 mm long. Several sclerites with some side-branches (Figure
Surface layer base of stalk. Sclerites similar to those of the top of the stalk but with longer spines (Figure
Interior base of stalk. Spindles with widely placed simple spines and tubercles; the spindles often have side-branches (Figure
Colour. Colony cream.
Red Sea, Socotra, Chagos Archipelago, Madagascar.
In the catalogue of the
The lectotype SMF 1279 has some supporting bundle spindles with a somewhat leafy projecting end (not depicted); ZMTAU 26194 and ZMTAU 26195 have some with a smooth spine (not depicted).
ZMTAU Co 25851 (Figures
The type material of Nephthea galbuloides has been re-examined (Figures
ZMTAU CO 34112–34113, identified by myself as Nephthea elatensis, have been used by
Ammothea viridis May, 1898: 33 (Muemba Island; East Africa);
Litophytum viridis;
Litophyton viridis;
Not Litophytum viride;
Not Litophytum viridis;
Not Litophyton viridis;
Ammothea stuhlmanni May, 1898: 34 (East-Africa);
Litophytum stuhlmanni;
Not Litophytum stuhlmanni;
Not Litophyton stuhlmanni;
Ammothea sanderi May, 1899: 141, pl. 3 fig. 26, pl. 5 fig. 12 (Zanzibar).
Litophytum sanderi;
Litophyton sanderi; Verseveldt and Benayahu 1983: 4 (Eilat, Gulf of Aqaba, 40-45 m, leg. Ch. Lewinsohn; listed only).
Litophytum crosslandi Thomson & McQueen, 1908: 56 (Red Sea, Coral reef of Khor Delaweb, 3-4 feet).
Litophytum acutifolium Kükenthal, 1913: 12, fig. 1, pl. 1 fig. 1 (Egyptian Red Sea coast, Berenice);
Not Litophyton acutifolium;
Colonies flabby, end lobes finger-like. Polyps with irregularly arranged, smooth rodlets adaxially and spiny rodlets abaxially; these rodlets are up to 0.1 mm long. Sometimes a few spindles are also present in the polyp stalk; sometimes the polyps are unarmed. Surface base of stalk with radiates, derivatives of these, and unilateral spinose spindles, the latter up to 0.5 mm long; many with side branches. Interior base of stalk with spindles up to 1 mm long; they can have side branches.
Colony flabby, 10.5 cm long and 7 cm wide (Figure
Polyps and branches. Without sclerites (Figure
Surface layer top of stalk. Capstans, spindles and unilateral spinose spindles; all with closely set tubercles; length up to 0.15 mm (Figure
Surface layer base of stalk. Capstans, spindles and branched spindles; the spindles up to 0.85 mm long (Figure
Interior base of stalk. Spindles with widely placed simple tubercles (Figure
Colour of colony. Cream.
Red Sea, East Africa.
Syntype
The difference between Litophyton viridis, L. acutifolium, L. crosslandi, L. sanderi, and L. stuhlmanni is only based on the polyps, those of L. viridis having no sclerites at all, while the other four species have few sclerites in the polyps. I regard the polyps without sclerites of L. viridis an extreme case of a species with a few sclerites in the polyps and synonymize L. acutifolium, L. crosslandi, L. sanderi, and L. stuhlmanni with L. viridis.
Apart from being much smaller (Figure
The colony fragment of Litophyton crosslandi present in the Natural History Museum (BM 1933.3.13.193) is only part of the colony originally described. The total length of the fragment is 5.3 cm (Figure
The holotype of Litophyton acutifolium (Kükenthal, 1913: pl. 1 fig. 1) was cut into pieces, and these are now stored as
ZMTAU Co 32941 and 33091 are only fragments of the top of colonies, the flabby nature of the fragments together with the sclerites matching those of L. viridis made me identify them as this species.
ZMTAU Co 34114 (previously identified as L. acutifolium by me) has been used by
ZMTAU 26193 (Figure
L. maldivensis and L. acuticonicum both also have polyps with limited amount of sclerites. L. acuticonicum differs from L. viridis in having much larger interior stalk sclerites (up to 2 mm long). L. maldivensis has overall much smaller interior stalk sclerites which mostly have blunt ends. L. striatum has similar looking sclerites in the interior of the base of the stalk as ZMTAU 26193 but in that species the polyps are much stronger armed.
Litophyton simulatum (Verseveldt, 1970),
Litophyton striatum (Kükenthal, 1903),
ZMTAU Co 25672 1894, Red Sea, South tip Sinai, Shab Mahmud, beacon rock, depth 0–20 m, 12 July 1987, coll. Y. Benayahu; disintegrated sclerites.
ZMTAU Co 26081, Red Sea, South tip Sinai Ras um Sud, 9 October 1988, coll. Y. Benayahu; stalk missing.
ZMTAU Co 26255, Red Sea, South tip Sinai, Ras Zaatir, 10 October 1989, coll. Y. Benayahu; consists of seven fragments all lacking a stalk.
ZMTAU Co 28609, Red Sea, Gulf of Aqaba, Eilat (Marin Lab.), depth 3 m, 1 August 1984, coll. Y. Benayahu; disintegrated sclerites.
ZMTAU Co 33090, Israel, Gulf of Aqaba, nature reserve, May 2000, coll. Y. Benayahu; fragments of branches, no internal sclerites found.
ZMTAU Co 25827 1460, Red Sea, South tip Sinai, Shab el Utaf, depth 10 m, 8 July 1986, coll. Y. Benayahu; base missing.
ZMTAU Co 26238, 4 small colonies, Red Sea, Gulf of Aqaba Wadi Magrash km 207, 17 April 1979, coll. Y. Benayahu; large supporting bundle spindles unlike other species, base looks a bit like L. maldivensis.
I thank Dr. Yehuda Benayahu (ZMTAU) for loan of the extensive material examined here, and Dr. Goetz Reinicke, Deutsches Meeresmuseum, Stralsund, Germany for material from Socotra and the Chagos Archipelago. Mr. Koos van Egmond (NCB) and Mr. A. Shlagman (ZMTAU) are thanked for curatorial assistance. I thank Dr. Phil Alderslade (Commonwealth Marine and Atmospheric Research, Tasmania) and Dr. Yehuda Benayahu (ZMTAU) for useful comments on the manuscript.