Echinodermata : Holothuroidea

Zootaxa 1245: 1–51 (2006) www.mapress.com/zootaxa/

ISSN 1175-5326 (print edition)

Copyright © 2006 Magnolia Press

ISSN 1175-5334 (online edition)

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New species and new records of dendrochirotid and dactylochirotid holothuroids (Echinodermata : Holothuroidea) from off the east coast of South Africa AHMED S. THANDAR School of Biological and Conservation Sciences, University of KwaZulu-Natal, Private Bag X54001, Durban 4000, South Africa. E-mail: [email protected]

Table of contents Abstract ............................................................................................................................................. 2 Introduction ........................................................................................................................................ 3 Systematic account ............................................................................................................................ 5 Order Dendrochirotida Grube .................................................................................................... 5 Family Phyllophoridae Oestergren ............................................................................................ 5 Subfamily Thyoninae Panning ................................................................................................... 5 Genus Thyone Oken ................................................................................................................... 5 Thyone avenusta Cherbonnier, 1970 .......................................................................................... 5 Thyone comata Cherbonnier, 1988 ............................................................................................ 7 Genus Stolus Selenka ............................................................................................................... 12 Stolus pseudoalbescens Thandar, 2005 .................................................................................... 12 Stolus cf pseudoalbescens Thandar, 2005 ................................................................................ 13 Subfamily Semperiellinae Heding & Panning ......................................................................... 14 Genus Selenkiella Heding & Panning ...................................................................................... 14 Selenkiella paradoxa Cherbonnier, 1970 ................................................................................. 14 Family Sclerodactylidae Panning ............................................................................................. 17 Subfamily Sclerothyoninae Thandar ........................................................................................ 17 Genus Sclerothyone Thandar ................................................................................................... 17 Sclerothyone velligera (Ludwig and Heding, 1935) ................................................................ 17 Subfamily Cladolabinae Heding & Panning ............................................................................ 19 Genus Ohshimella Heding & Panning ..................................................................................... 19 Ohshimella ehrenbergii (Selenka, 1868) .................................................................................. 20 Ohshimella ocula (Cherbonnier, 1988) comb. nov. ................................................................. 22 Family Cucumariidae Ludwig ................................................................................................. 25 Subfamily Cucumariinae Ludwig ............................................................................................ 25 Genus Pawsonellus Thandar .................................................................................................... 25

Accepted by A. Dartnall: 22 May 2006; published: 26 Jun. 2006

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Pawsonellus africanus Thandar, 1986 ..................................................................................... 25 Subfamily Colochirinae Panning ............................................................................................. 26 Genus Trachasina Thandar ...................................................................................................... 26 Trachasina crucifera (Semper, 1869) ...................................................................................... 26 Genus Plesiocolochirus Cherbonnier ....................................................................................... 28 Plesiocolochirus dispar (Lampert, 1889) ................................................................................ 28 Plesiocolochirus tessellara (Cherbonnier, 1970) comb. nov. ................................................. 30 Genus Staurothyone H.L. Clark ............................................................................................... 33 Staurothyone rosacea (Semper, 1869) ..................................................................................... 33 Family: Psolidae Ed.Perrier ..................................................................................................... 35 Genus Psolidium Ludwig ......................................................................................................... 35 Psolidium multipes sp. nov. ..................................................................................................... 35 Psolidium acorbulum sp. nov. ................................................................................................. 38 Genus Ekkentropelma Pawson ................................................................................................. 41 Ekkentropelma groovia sp. nov. ............................................................................................... 41 Order Dactylochirotida Pawson & Fell .................................................................................... 44 Family Vaneyellidae Pawson & Fell ........................................................................................ 44 Genus Psolidothuria Thandar .................................................................................................. 44 Psolidothuria yasmeena sp. nov. ............................................................................................. 44 Acknowledgements ......................................................................................................................... 46 Literature cited ................................................................................................................................ 47

Abstract Numerous unidentified specimens of Holothuroidea received from the Natal and South African Museums or present in the collections of the former University of Durban-Westville, contain several new species and many that are new to the fauna of South Africa. This paper includes descriptions of new species or new records of several dendrochirotid holothuroids and a dactylochirotid from off the east coast of South Africa, extending from Kosi Bay in the north to the Port St Johns-East London area, in the Eastern Cape Province, the latter region represents the transition zone between the southern African subtropical and warm-temperate zoogeographical provinces. A few other species have been added if they represent juvenile material, or considered extensions of ranges (bathymetric or otherwise) of previously known species, or confirmation of a species previously recorded from a single locality in the region under consideration. Of the 16 species here treated, four are new to science, seven new to the fauna of South Africa, whereas the rest are either extension of ranges or additional records. The four new species are Psolidium multipes, P. acorbula, Ekkentropelma groovia and Psolidothuria yasmeena. The following six species are additions to the fauna of South Africa: Thyone avenusta Cherbonnier, 1970; T. comata Cherbonnier, 1988; Selenkiella paradoxa Cherbonnier, 1970; Ohshimella ocula (Cherbonnier, 1988); Plesiocolochirus tessellarus (Cherbonnier, 1970) and Staurothyone rosacea (Semper, 1869). Pseudocolochirus dispar (Lampert, 1889) is new to the fauna of southern Africa, recorded for the first time from Mozambique. The remaining species include Stolus pseudoalbescens Thandar, 2005; Sclerothyone velligera (Ludwig & Heding, 1935); Ohshimella ehrenbergii (Selenka, 1868); Pawsonellus africanus Thandar, 1986 and Trachasina crucifera (Semper, 1869).

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© 2006 Magnolia Press

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Key words: Holothuroidea, Dendrochirotida, Dactylochirotida, South Africa, new species, new combinations

Introduction The first comprehensive report of the southern African holothuroid fauna is that of H.L. Clark (1923), which included 30 species of which three were new to science. He, however, despite the poor quality of his material from the South African Museum, provided a key to all the species then known from southern Africa. Prior to this work only a few (ca.7) species were recorded from southern Africa, excluding Mozambique (see Pallas 1766, Quoy & Gaimard 1833, Ludwig 1875 & 1882, Lampert 1885, Théel 1886a, Vaney 1908a & b), Britten 1910, Gilchrist 1920), mostly based on early European expeditions. E cological surveys carried out along the southern African coast by the University of Cape Town from the early thirties led to reports of the holothuroids by Heding (1938), John (1939), Deichmann (1944 & 1948) and Cherbonnier (1952, 1953, 1954 & 1970). Deichmann’ s (1948) revision was very comprehensive and also based on material present in the Museum of Comparative Zoology, Harvard, USA. Although she described no new species and excluded all reference to tropical species from Mozambique and truly deepsea forms, her work includes diagnoses, keys and distribution records of altogether 22 species then valid. Cherbonnier’ s (op. cit.) contributions led to descriptions of no fewer than 22 new species and re-descriptions of many others. Although he provides neither keys nor diagnoses, his descriptions are detailed and well illustrated but his earlier publications (1952–1954) suffered much from lack of any consideration given to Panning’ s (1949) revision of the family Cucumariidae, which preceded those works. Of the 22 new species described by Cherbonnier as many as 18 appear to be valid but as many as 12 were based on single specimens. Our knowledge of the Mozambique holothuroid fauna stems largely from the works of Semper (1868), Bell (1884), Pearson (1910), H.L. Clark (1923) and Cherbonnier (1952, 1970) and from reports of the University of Witwatersrand ecological surveys at Inhaca Island (Kalk 1958; Macnae & Kalk 1958, 1962,) and on the shores of northern Mozambique (Kalk 1959). Prior to Thandar’ s (1984) revision of the southern African holothuroid fauna our knowledge of the southern African deep-sea species was very scanty and based on collections made by the ‘Valdivia’ expedition (Ludwig & Heding 1935), the ‘Galathea’ expedition (Hansen 1975) and those described by H.L. Clark (1923), Cherbonnier (1952) and A.M. Clark (1977). Thandar’ s revision was never published in its entirety but numerous papers resulted from this revision as well as from his and his co-workers’ later works. The taxonomic ones of these mostly revise several families pertinent to southern Africa, south of the Tropic of Capricorn, describe new taxa or new records, or report on deep-sea collections (see Thandar 1977, 1985, 1986, 1987a & b, 1989a, b & c, Thandar &

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Rowe 1989, Thandar 1990, 1991, 1992, 1994, 1996, 1998, Rajpal & Thandar 1998, 1999, Thandar & Rajpal 1999, Thandar 1999, Thandar 2001, Thandar & Moodley 2003, Samyn & Thandar 2003, Massin et al. 2004, Thandar & Samyn 2004, Thandar 2005). These additions still await a monographic treatment, which can only be completed once several unidentified collections, currently in the hands of the author, have been studied and recorded. These collections come from waters ranging from the littoral to the deep sea and originate from the South African and Natal Museums and the former University of Durban-Westville. They contain several hitherto undescribed species and many new records for the South or southern African region. Most of the materials have been collected as a result of deep-water dredgings on the R.V. ‘Meiring Naude’ by researchers from the South African and Natal Museums, while others result from either the Natal Museum’ s NMDP project, which extended from Kosi Bay on the north-east coast of KwaZulu-Natal to False Bay, east of the Cape Peninsula, or are a result of miscellaneous collections present in the collections of the above Museums and the former University of DurbanWestville. The current paper contains descriptions of new species and new records of several dendrochirotid holothuroids and a dactylochirotid from off the east coast of South Africa, extending from Kosi Bay in northern KwaZulu-Natal to the Port St. Johns-East London area in the Eastern Cape Province. A few other species have been added if they represent juvenile material or considered extensions of ranges of previously known species or confirmation of a species previously recorded from a single locality in the region under consideration. For the latter detailed descriptions are avoided if the species was previously fully described. Five new species of dendrochirotids from this region and based on these collections have already been described elsewhere (see Rajpal & Thandar 1998, 1999; Thandar & Rajpal 1999, Thandar 2005) while descriptions of a new dendrochirotid species and one originally known from only Madagascar and the Comores but also now discovered from northern KwaZulu-Natal have also been published (see Samyn & Thandar 2003, Thandar & Samyn 2004). Altogether a total of 16 species are here treated, of which four are new to science, six new to the fauna of South Africa, one to that of Mozambique and five are additional records. Acronyms: The following acronyms are used throughout the text to denote source of material and/or location of types: MNHNP—Museum National d’ Histoire Naturelle, Paris; NM—Natal Museum, Pietermaritzburg, South Africa; NMDP—Natal Museum Dredging Programme; SAMSouth African Museum, Cape Town; UCT—University of Cape Town; UDW—University of Durban-Westville; ZMUA—Zoological Museum, University of Amsterdam.

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Systematic account

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1245 Order Dendrochirotida Grube Family Phyllophoridae Oestergren Subfamily Thyoninae Panning Genus Thyone Oken Thyone avenusta Cherbonnier, 1970 Figure 1 Thyone avenusta Cherbonnier, 1970: 286, fig 4 A–M; Thandar, 1990: 214, fig. 5 a–g; Cherbonnier, 1988: 197, fig 85 A–M.

Type MNHNP. Type locality Morrumbene, Mozambique, 35 cm. Previous South African record None Material examined SAM — A23721, R.V. ‘Meiring Naude’ St. XX 94, 30° 13.8’ S, 30° 54.2’ E, between Green Point and Widdenham, KwaZulu-Natal, 8.vii.1986, 70 m., 1 spec. Description Specimen small, slightly curved with anterior and posterior ends turned upwards; length 11 mm, breadth in mid-body about 3.5 mm; colour, in alcohol, uniformly white with a slight tinge of yellow at the extreme anterior end. Mouth and anus terminal. Tentacles retracted; anal teeth distinct, flanked by subterminal podia. Dissection revealed 10 whitish tentacles - eight elongated but of unequal size, ventral two much reduced. Podia well developed, especially ventrally, where they are arranged mostly in the ambulacra with a few also scattered in the interambulacra; suckers of ventral podia with diameter much larger than that of podia; dorsal podia shorter, scattered. Body wall thin, soft. Calcareous ring (Figure 1I) well developed, extremely long, reaching almost twothirds length of body from anterior end; total length of ring, including introvert, ca. 6.5 mm, tubular, with the radial plates deeply incised, i.e. bifurcating before the posterior border of the interradial plates; both radial and especially interradial plates broken into a HOLOTHUROIDEA


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mosaic; anterior projections of all plates of more or less equal length, with those of the radials being bifid for the attachment of retractor muscles, posterior processes of radial plates distally curved and fragmented into a single series of segments. Polian vesicle single, elongated. Stone canal single, short, straight and free, terminating in cotyledonous madreporic body (Figure 1J). Respiratory trees well branched, both uniting before opening into an elongate cloaca. Gonad immature, in two tufts of unbranched, white tubules. Longitudinal muscles unpaired, retractor muscles well developed, originating from longitudinal muscles in anterior third of body, more anteriorly in the ventral ambulacra.

FIGURE 1. Thyone avenusta Cherbonnier, 1970. SAM-A23721. A. tables from body wall; B. tables from dorsal podia; C. perforated rods from dorsal podia; D. end-plate from dorsal podium; E . rosettes from introvert; F. rosette-like rods from tentacle; G. rosettes from tentacles; H. rods from tentacles; I. calcareous ring; J. madreporite. A-H scale a.

Spicules of body wall comprise typical Thyone tables (Figure 1A) with small discs (38–48 µm, mean 44 µm) with smooth or slightly spinose margins and perforated by 4–5 (occasionally 6) large holes; spire short (33–43 µm, mean 38 µm), two-pillared, terminating in a crown of teeth usually developed as two clusters; spire sometimes reduced to knobs on surface of disc. Podial deposits comprise few tables, numerous rods and an endplate (Figure 1D). Tables (Figure 1B) usually with an irregular disc (ca. 50 µm long), pierced by more than four holes, and a short, occasionally arched spire provided with tuberosities; rods (Figure 1C) 62–76 µm (mean 69 µm), slightly curved with smooth

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margins and perforated by 1–3 holes at each end; endplates of the ventral podia better developed than dorsal (105–129 µm, mean 119 µm cf. 86–91 µm, mean 89 µm), with small holes in the centre, larger ones outside these in a single series, and small ones outside these in one or two concentric series. Tentacles supported by curved, elongated, slender rods (49–110 µm, mean 79 µm) (Figure 1H) with one or more perforations at expanded and/or digitated ends, other smaller rosettes-like rods (35–52 µm, mean 44 µm) (Figure 1F) as well as by rosette-shaped bodies (22–34 µm, mean 28 µm) (Figure 1G), quite unlike the typical rosettes of the introvert. Introvert spicules developed only as closed rosettes (22–34 µm, mean 29 µm) (Figure 1E). Distribution Madagascar, Mozambique and east coast of South Africa, up to 70 m. Habitat Mangrove, sand. Remarks The specimen at hand corresponds well with the description of the type given by Cherbonnier (1970), except that the podial deposits comprise, in addition to a few characteristic tables, also large curved rods. Thandar (1990) examined a single specimen of this species that was part of the type-material and found body wall tables with up to 16 holes on the discs, no rods in the podia and recorded the presence of anal teeth, reported as absent in the type. Thandar concluded that these differences were perhaps age variations as the type measured only 30 mm whereas his specimen was 47 mm, with spicules restricted to the posterior end. The present specimen is also small (11 mm) and hence its body wall tables are similar to those of the type, both in size and the number of holes in the disc. The presence of rods in the podia may be a juvenile feature. A characteristic of this species is the presence of three types of spicules in the tentacles (smooth rods, rosette-like rods, and rosettes). T. avenusta can be confused with T. propinqua Cherbonnier, 1970 because of its characteristic four-holed table discs but the latter species has plate-like spicules, instead of rosettes, in the introvert and large multilocular rods in the tentacles. The present specimen, the first record of this species from South Africa, taken at a depth of 70 m, is the deepest yet recorded for the species as the type came from mangrove off the Mozambican coast while the Madagascar material was taken from the reef at Tuléar.

Thyone comata Cherbonnier, 1988 Figure 2 Thyone comata Cherbonnier, 1988: 198–190, fig. 80.

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MNHNP, 3592. Type locality Tuléar (Madagascar), 8–50 m. Previous South African record None Material examined SAM-A27889, NE of Kosi Bay, KwaZulu-Natal, 26° 52.8’ S, 32° 54.8’ E, NMDP St. ZA 33, 3 vi 1990, 45 m., 1 spec.; SAM-A27890, 26° 52.4’ S, 32° 55.2’ E, NMDP St. ZA 38, 3 vi 1990, 45–47 m., 1 spec.; SAM-A27891, Gypsy Hill, Leadsman Shoal, 27° 49.7’ S, 32° 38.2’ E, ‘Meiring Naude’ St. ZK 1, 8.vi.1988, 47–50 m., 1 spec. Description Best preserved specimen (SAM-A27890) swollen anteriorly and in the middle, narrowing posteriorly, posterior end turned upwards. Length of ventral surface excluding turned up posterior end 22 mm (35 mm including posterior end), width of mid-body 8 mm. Largest specimen about 38 mm long along ventral surface, 10 mm broad in mid-body. All specimens whitish in colour, including podia. Podia elongated, evenly scattered and often not retracted, slightly more numerous ventrally, giving the skin a villose appearance, sucking discs small. Mouth anterior, anus dorsally directed and in at least two specimens encircled by minute calcareous teeth. Tentacles retracted, number and size variation could not be ascertained despite dissection of calcareous ring of two individuals. Calcareous ring complex, tubular, with radial plates carrying long bifurcate processes, both ring and processes fragmented, the latter in a single series; fragmentation of radial and interradial plates more or less similar. Both radial and interradial plates triangular, the anterior process of each radial plate with shallow indentation for insertion of retractor muscle, that of interradial plates pointed. Anterior process of each plate flanked at tip by thickened alae, those of the radials not meeting, hence each radial plate appears anteriorly notched; in addition, each radial plate deeply incised posteriorly, i.e. bifurcating before the posterior border of the corresponding interradial plate. Calcareous ring, including processes, about 10 mm in length in the smaller specimen and about 15 mm in the distorted specimen. Polian vesicle single, elongated. Stone canal not detected in both dissected specimens. Spicules of dorsal and ventral body wall (SAM-A27890) identical, comprising twopillared tables (Figure 2A,B & C) with an irregular disc, perforated by usually four central and a varying number of marginal holes of more or less equal size to the central holes. Spire of moderate height (Figure 2B), terminating in 4(–6) teeth. Length of dorsal table discs 80–130 µm (mean 98 µm) with 8–13 holes, spire height about 40 µm; length of

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ventral table discs 90–110 µm (mean 100 µm) with 7–14 holes, spire height 40–44 µm. Deposits of dorsal and ventral podia identical, comprising two-pillared tables with curved discs perforated by four large central holes and several small holes at each end. Spire usually terminating in a single tooth or rarely in two teeth. Tables of dorsal podia (Figure 2G) with discs 62–96 µm (mean 81 µm), spire height 27–46 µm (mean 34 µm); table discs of ventral podia (Figure 2F) 68–101 µm (mean 85 µm), spire height 22–36 µm (mean 29 µm). Endplates (Figure 2D & E) well developed with small central holes and 2–3 series of larger marginal holes with the most peripheral of marginal series always smaller than those within; diameter of endplates 102–135 µm (mean 120 µm). Anal region with broken periproctal plates and small tables similar to those of body wall. Introvert deposits include only two-pillared tables (Figure 2I, J) with multilocular discs (59–94 µm, mean 76 µm) of varying shapes and with short spires (30–40 µm), terminating in 4–6 teeth. Tentacle stalks with delicate, perforated rods (Figure 2K) (40–69 µm, mean 50 µm), tentacle tips with rosettes (Figure 2L) (23–32 µm, mean 27 µm). Longitudinal muscles with delicate, perforated and branching rods (45–81 µm, mean 67 µm) (Figure 2J). Dorsal table discs of largest specimen 89–132 µm long (mean 113 µm) with 9–15 holes, spire height 44–60 µm; ventral table discs 94–141 µm long (mean 114 µm) with 10–19 holes, spire height 41–50 µm. Largest specimen with table discs of dorsal podia 62–139 µm long (mean 100 µm), spire height 27–57 µm (mean 35 µm); table discs of ventral podia 65–128 µm long (mean 86 µm), spire height 24–41 µm (mean 32 µm); endplates 115–168 µm in diameter, larger in dorsal podia.

FIGURE 2. Thyone comata Cherbonnier, 1988. SAM-A27889. A. tables from antero-dorsal body wall; B. tables from side; C. tables from postero-ventral body wall; D. end-plate from dorsal podium; E. end-plate from ventral podium; F. tables from ventral podia; G. tables from dorsal podia; H. tables from proximal introvert; I. tables from distal introvert; J. rods from longitudinal muscles; K. small perforated rods from tentacle stalk; L. rosettes from tentacle tip. A–K scale a; L scale b.

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TABLE 1. Distribution, depth records and morphological characters of four related species of Thyone, including the two South African specimens of T. comata. T. pedata (ex Liao 1995)

T. villosa (holotype)

T. comata T. comata T. comata (ex Cherb. (A27889) (A27890) 1988)

T.crebrapodia (ex Cherb. 1988)

Distribution

China Sea

Philippines

Madagascar East coast East coast South Africa & Comores South Africa

Madagascar

Depth (m)

55

26

8–32

45

45–47

shallow

Length (mm) 18–22

20

5–27

27

18

12–24

Anal teeth

absent

present

present

present

undetected

?

Body wall tables disc length (µm)

119–128

93–122

100–130

89–141

79–128

55–60

spire heigth (µm)

?

37–56

35–45

41–60

39–44

35–40

Podial tables 81–106 disc length (µm)

93–194

80–100

62–139

62–101

65–85

spire heigth (µm)

31–41

37–59

20–35

24–57

22–46

20–30

Introvert deposits

rosettes (in heaps)

tables, discs tables, discs tables, tables, discs (90–103 multilocular discs (59–94 µm), µm), multilocul multilocular multilocular ar

tables, discs 70–100µm), multilocular

Tentacle deposits

stout rods

irregular branching perforated rods and plates

branched rods, perforated at ends, some rosette-like and rosettes

slender rods, perforated at ends and rosettes



Longitudinal ? muscles

thick, thin to thick thin, thin, ? conspicuous inconspicu inconspicuous ous

Longitudinal ? muscle deposits

absent

?

slender, slender, ? perforated, perforated, branching branching rods rods

Distribution Southwest Indian Ocean, 8–50 m.

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Habitat Sand, fine sand, coral sand, algae.

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Remarks The specimens here studied come very close to Thyone villosa Semper, 1868, from Cebu (Philippines), re-described from the type by Panning (1949), T. comata Cherbonnier, 1988 from Tuléar (Madagascar) and Mayotte Island (Comores) and, on the bases of body wall deposits alone, to T. pedata Semper, 1868, recently described by Liao (in Liao & Clark 1995) from new material collected from the Gulf of Tonkin, South China Sea. In fact, so much is the resemblance of the present material to the above three species that I was at first inclined to consider T. pedata and T. comata to be junior subjective synonyms of T. villosa, a species not encountered since its original description. However, judging from an examination of the holotype of T. villosa, received by courtesy of the Hamburg Museum, and from Cherbonnier’ s description of T. comata and that of Liao’ s of T. pedata, there is some justification in keeping the three species separate. The southern African material comes very close to T. comata, to be almost indistinguishable. It differs from T. villosa in the form of the tentacle deposits, the thinness of the longitudinal muscles and the presence therein of slender branching rods (absent in T. villosa) and from T. pedata in the presence of anal teeth and slender rods and rosettes in the tentacles and only tables (instead of rosettes) in the introvert.

FIGURE 3. Thyone villosa Semper, 1868. Holotype. A. body wall tables; B. podial tables; C. tables from distal introvert; D. rods from tentacle tip; E. plates from tentacle tip. A–C scale a; D & E scale b.

It is a pity that Cherbonnier (1988) compared his T. comata only with T. pedata while his T. crebrapodia, also from Tuléar, he compared with T. villosa Semper and T. dura Koehler & Vaney. The latter species, described from the deep Arabian Sea (at 264 m), according to Koehler and Vaney (1908), shares some similarities with T. pituitosa Sluiter, 1901 from the deep tropical West Pacific (at 310 m). An examination of the holotype of the latter species at the ZMUA, showed that its calcareous ring is different from typical

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members of Thyone in being of the sclerodactylid rather than the phyllophorid type (see Thandar 1989c). This is also true for the calcareous ring of T. dura, judging from the figure given by Koehler & Vaney. Thus, Cherbonnier’ s T. crebrapodia cannot be compared with either T. pituitosa or T. dura. In fact, his T. comata resembles T. villosa more than does his T. crebrapodia. The latter species differs from the former two in the smaller size of its body wall and podial tables and in its tentacle deposits. Table 1 lists the characters of the above four nominal species of Thyone as well as those of the southern African material. The body wall, podial, tentacle and introvert spicules of the holotype of T. villosa are illustrated in Figure 3. Semper (1868) separated his T. pedata from T. villosa on the basis of the absence of anal teeth and this system was adopted by both Lampert (1885) and Théel (1886a). Regrettably, Panning (1949) overlooked T. pedata in his revision of the Cucumariidae. However, as shown above and in the table below, there is enough justification in maintaining the distinctiveness of both these species.

Genus Stolus Selenka Stolus pseudoalbescens Thandar, 2005 Stolus pseudoalbescens Thandar, 2005: 161, fig. 1, 3A & C, 6A & B.

Type SAM A-24194. Type locality Durban, off Mngeni River Mouth (29° 49.8’ S, 31° 12.2’ E). Previous records Off KwaZulu-Natal, South Africa (see Thandar 2005). Additional records SAM — A27892, Natal, S.E. of Umzimbazi River, 30° 8.2’ S, 30° 56.5’ E, R.V. ‘Meiring Naude’, St. XX 102, 8.vii.1986, 65 m., fine sand, 1 spec.; SAM — A27893, Natal off Durban, 29° 50.2’ S, 31° 12.3’ E, R.V. ‘Meiring Naude’, St. XX 113, 9.vii.1986, 95 m., fine, slightly muddy sand, 1 spec.; SAM — A27894, Natal off Durban, 29° 49.2’ S, 31° 14.3’ E, R.V. ‘Meiring Naude’, St. XX 117, 9.vii.1986, 104-110 m., muddy sand and broken shells, 1 spec.; SAM — A27895, 32° 14.8’ S 29° 00.8’ E, R. V. ‘Meiring Naude’, St. SM 239, 25.vi.1979, 90 m., 1 spec.; SAM — A27896, Ballito, 29° 34.4’ S, 31° 17.9’ E , R.V. ‘Meiring Naude’, St. XX 130, 7.vi.1988, 60 m., 2 spec.; SAM — A27897, Gobey’ s Point, 27° 26.0’ S, 32° 44.5’ E, R.V. ‘Meiring Naude’, St. ZG 2, 3.vi.1987, 55-100 m., 1 12


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spec; SAM — A27898, off Cooper Lighthouse, Durban, Natal 29° 56.2’ S, 31° 10.2’ E, R.V. ‘Meiring Naude’, St. XX 118, 9.vii.1986, 88 m., firm grey muddy sand, 1 spec.; SAM - A 23720, off Durban, Natal, 29° 50.0’ S, 31° 14.2’ E, R.V. ‘Meiring Naude’, St. XX 115, 9.vii.1986, 130 m., 1 spec. Distribution Only known from the east coast of South Africa, 55–130 m. Habitat Mud, stones, coarse sand, muddy sand and shell debris.

Stolus cf pseudoalbescens Thandar, 2005 Stolus cf pseudoalbescens Thandar, 2005:119, fig. 2, 3B.

Previous record Off KwaZulu-Natal, South Africa (see Thandar 2005). Additional record SAM — A27899, South of Umlaas River, 30° 0.8’ S, 31° 2.6’ E, R.V. ‘Meiring Naude’, St. XX 73, 10.vii.1985, 100 m., 1 spec. Distribution Only known from the east coast of South Africa, 85–103 m. Habitat Fine muddy sand, shell debris. Remarks The above 10 specimens of Stolus pseudoalbescens, contained in the ‘Meiring Naude’ material received from the SAM, were collected with the type material of the species from the Natal Museum (see Thandar 2005:117, 119). While nine of the specimens are clearly identical with S. pseudoalbescens, one is without tubercles on the skin and referable to S. cf. pseudoalbescens also described in the same paper. Hence another description of the species is unnecessary.

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Subfamily Semperiellinae Heding & Panning

1245 Genus Selenkiella Heding & Panning Selenkiella paradoxa Cherbonnier, 1970 Figure 4 Selenkiella paradoxa Cherbonnier, 1970: 291–293, fig. 6 (A–P); Day, 1974: 94; Cherbonnier, 1988: 234–235, fig. 103; Thandar, 1990:219–221, fig. 9 a–f.

Type MNHNP. Type locality Morrumbene, Mozambique, 35 cm. Previous South African record None Material examined SAM-A27900, SE of Durban Bluff, 29° 59.9’ S, 31° 03.8’ E, ‘Meiring Naude’, St. XX 146, R. Kilburn, 13 vi 1989, 60 m, 1 spec. (female); SAM-A27901, off Mngeni River Mouth, Durban, 29° 49.8’ S, 31° 12.2’ E, ‘Meiring Naude’, St.XX 149, R. Kilburn, 13 vi 1989, 95 m, 1 spec. Description Larger of the two specimens barrel-shaped, slightly bloated ventrally; length approximately 41 mm, width of mid-body 10 mm, height about 14 mm. Colour, in alcohol, a variegated greyish-brown, suckers white; skin wrinkled. Mouth anterior, anus terminal, the latter flanked by minute calcareous teeth, not easily visible. Podia scattered, not confined to ambulacra, more numerous ventrally, suckers well developed. Tentacles 25, retracted, arranged in three rings of 10+10+5, however, first and second rings not clearly defined. All tentacles well branched, brownish in colour with lighter stalks. Body wall soft, smooth to the touch. Calcareous ring complex (Figure 4U), tubular, with unbroken radial plates, slightly higher than interradial plates and carrying short, thick, paired, inwardly curled processes fragmented into few segments, several anterior notches and a depression for insertion of retractor muscle; interradial plates roughly triangular, not notched, but anteriorly pointed, and broken into two pieces; calcareous ring, including processes, about 7 mm long. Polian vesicle single, large, sac-like. Stone canal short, white, embedded in dorsal mesentery; madreporic body (Figure 4T) somewhat oval, well calcified, also lodged in mesentery. Respiratory trees well branched, right one slightly longer than left, both reaching anterior

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third of body, uniting before opening into cloaca. Cloaca narrow, elongate, suspensors well developed. Gonad (ovary) in two tufts of short, unbranched tubules full of eggs, left tuft better developed than right. Longitudinal muscles thick, unbranched. Retractor muscles well developed, originating from anterior third of longitudinal bands, slightly more anteriorly in the two dorsal ambulacra.

FIGURE 4. Selenkiella paradoxa Cherbonnier, 1970. SAM-A27900. A. pseudobuttons from lateral body wall; B. side view of pseudobuttons from lateral body wall; C. reduced tables from body wall; D. three types from rosettes from body wall; E. rods from body wall; F. plates from ventral podia; G. plates from dorsal podia; H. rods from dorsal podia; I. plate-like rods from dorsal podia; J. rods from ventral podia; K. minute rods from ventral podia; L. rosettes from introvert; M. mulberry-like rosettes from tentacles; N. rods from tentacles; O. rods from gonad; P. tables from anal region; Q. side view of tables from anal tables; R. rosette-like plates and rods from cloacal muscles; S. part of end-plate; T. madreporite; U. calcareous ring. A-R scale a.

Spicules of the dorsal, lateral and ventral body wall identical, comprising minute rods (29–76 µm, mean 54 µm), pseudobuttons, tables with reduced spires (47–69 µm, mean 63 µm) and rosettes (36–61 µm, mean 45 µm). Rods (Figure 4E.) numerous, smooth, with or without perforations, either representing complete spicules or developmental stages of pseudobuttons and tables. Pseudobuttons (Figure 4A, B) numerous, with 1–6 holes. Table discs (Figure 4C) subcircular to oval, with usually a smooth rim, two or more holes and commonly a pair of nodules indicating reduced spires. Rosettes (Figure 4D) equally developed dorsally and ventrally, originating from branched, anastomosing rods, which may or may not leave perforations — larger rosettes usually with perforations, smaller

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ones without. Anal region with better-developed tables (Figure 4P, Q), with usually an oval to round disc (50–98 µm, mean 76 µm) with an undulating rim, four central holes and a short (14–25 µm, mean 21 µm) spire of four pillars, ending in usually a quadrangular crown bearing a few blunt teeth. Podial deposits include rods (Figure 4H, K) and pseudobuttons, identical to those of body wall but slightly larger (31–164 µm, mean 79 µm), other more specialized plate-like rods (82–200 µm, mean 130 µm) (Figure 4I, J), better developed ventrally, with a series of perforations along their length or with one or two perforations at ends, as well as typical, elongated (112–216 µm, mean 156 µm), multilocular plates (Figure 4F, G). Endplates up to 407 µm in diameter, with an irregular margin and numerous small to large holes scattered without any regular order (Figure 4S). Tentacle deposits only comprise straight or curved slender rods (Figure 4N) (40–168 µm, mean 85 µm) with slightly expanded, digitated ends carrying one or two perforations, and rosettes (Figure 4M) similar to those of body wall but slightly larger (26–93 µm, mean 51 µm). Introvert with only rosettes developed as minute (44–76 µm, mean 55 µm) branched rods (Figure 4L). Cloacal suspensors with branched rods (35–82 µm, mean 52 µm) developed as ‘pseudobuttons’ or rosette-like spicules (Figure 4R). Gonad with slender rods bearing one or more processes (Figure 4O), sometimes developed as apophyses, resembling those of the spicules of some elasipodids, notably Elpidia species. Retractor muscles without spicules. Distribution East coast of southern Mozambique and South Africa and west coast of south Madagascar, up to 95 m. Habitat Fine sand, mud and seaweeds. Remarks This species was based only on the holotype collected from Morrumbene (Mozambique) at 35 cm (Cherbonnier 1970) but Cherbonnier (1988) states that the holotype came from a depth of 35 m. The latter depth record hence appears to be a lapsus calami as is also Thandar’ s (1990) depth record of 3–5 m for the holotype. The species is also reported to occur at Inhaca Island, off Maputo (Mozambique), but this could not be confirmed. Hence the second definite record of the species is that of Cherbonnier (1988), based on seven specimens collected from Tuléar ( Madagascar). The present specimens from around Durban thus extend the known distribution of the species, and also double its recorded size. Both Cherbonnier (1970) and Thandar (1990) compare this wellcharacterised species with its two congenors, S. siamense Heding and Panning 1954, the type species, and S. malayense Heding and Panning 1954. Cherbonnier (1970, 1988) states that the tentacles are in two circles of 15 large tentacles on the outside and 10 small

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tentacles on the inside. However, in the large specimen, described above, there are clearly five very small tentacles on the inside and 20 large tentacles, apparently arranged in two rings, on the outside. This difference may not be very significant as the arrangement cannot be clearly discerned if the tentacles are retracted. Also insignificant is the fragmentation of the interradial plates. These differences and a few others concerning the spicules may be geographic variations in this species which is well characterised by its pseudobuttons and rosettes in the body wall, and elongated multilocular plates in the podia.

Family Sclerodactylidae Panning Subfamily Sclerothyoninae Thandar Genus Sclerothyone Thandar

Sclerothyone velligera (Ludwig and Heding, 1935) Figure 5 Cucumaria? velligera Ludwig and Heding, 1935: 70, text fig. 49. Pentamera velligera Deichmann, 1948: 351. Neothyone? velligera Panning, 1949: 458. Sclerothyone velligera Thandar, 1989c: 295, figs. 3 & 9b.

Type ?Bonn, Germany. Type locality Cape Point, South Africa. Previous southern African records West coast of Cape Peninsula. Material examined SAM-A27902, Off East London, Eastern Cape Province, 32° 28.6’ S, 28° 58.8’ E, R.V. ‘Meiring Naude’, St. SM 226, 24.vi.1979, 710–775 m., 1 spec. (male). Description Specimen tiny but an almost mature male; form cucumber-like with an extended tentacle crown. Length, including tentacle crown, 6 mm, tentacle crown 2 mm, breadth in mid-body 2 mm. Colour, in alcohol, a dull beige-grey. Anal papillae five, teeth-like,

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densely packed with spicules but not itself calcified. Tentacles 10, bushy, well-developed, mid-ventral two reduced. Podia non-retractile, restricted to ambulacra in 2–3 rows, well developed, giving impression of being crowded, especially on the dorsal surface; suckers well developed, diameter larger than diameter of podium. Calcareous ring (Figure 5H) well developed, non-tubular, compact, radial and interradial plates only connected to each other at their bases, radial plates anteriorly notched and bearing paired, faintly subdivided posterior prolongations, twice as long as radial plates; interradial plates triangular. Polian vesicle single, anteriorly directed; stone canal very short, free; madreporite (Figure 5H) minute, not well calcified. Gonadal tubules unbranched, full of spermatozoa. Respiratory trees poorly developed with sacciform endbranches at their base.

FIGURE 5. Sclerothyone velligera (Ludwig & Heding, 1935). SAM-A27902. A. tables from body wall; B. podial tables; C. podial plates; D. part of end-plate; E. introvert plates; F. plates from tentacle stalk; G. plates from tentacle tip; H. calcareous ring and madreporite. A–G scale a.

Spicules of body wall comprise tables (Figure 5A) of two types: commonly those with oval discs (65–106 µm, mean 87 µm), with four large central holes and a two-pillared spire (27–39 µm, mean 32 µm), terminating in a single point or with three or more teeth; others with larger, more robust, rectangular to sub-rectangular discs (79–88 µm, mean 85 µm), with 4–6 holes and bearing either an arched spire (ca 25 µm) or a two-pillared spire terminating in several teeth. No handle to body wall tables. Podia supported by two18


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pillared, toothed tables (Figure 5B) with a curved disc (67–110 µm, mean 88 µm) and a short spire (23–29 µm, mean 27 µm) as well as by curved, multilocular plates/rods (60–94 µm, mean 71 µm) with a spinose/serrated margins (Figure 5C). Endplates reduced (Figure 5D). Introvert supported by multilocular plates (58–94 µm, mean 76 µm) with jagged margins (Figure 5E). Tentacles also supported by multilocular plates (67–189 µm, mean 98 µm) with more jagged margins than those from introvert (Figure 5F, G). Distribution East and west coast of South Africa, 360–775 m. Habitat Rock. Remarks This species, described by Ludwig & Heding (1935), was rediscovered by Thandar (1989c), who pointed out several differences between his specimen and that of the type but despite this, identified his material with Ludwig & Heding’ s species. The differences, according to him, might have warranted the establishment of a new species but hesitated to do so. The current material, also taken from deep waters but off the South African southeast coast, differs from that described by Thandar from off the west coast, but less so than the differences between the west coast specimen and the type. In the present specimen the radial and interradial plates of the calcareous ring are fused at their bases, the processes of the radial plates not clearly fragmented, the body wall tables lack a handle/half ring on one side, and the introvert and tentacle spicules have more pronounced jagged margins. Thandar (1989c) did not mention the presence of endplates in his single specimen but these are recorded in the original manuscript as being well developed. In the present specimen endplates are rare or absent. These differences are perhaps geographic, age or individual variations and more material is required to confirm this. In the meantime the name S. velligera must also suffice for the east coast form. Ludwig & Heding did not comment on the maturity of their specimen but like the west coast specimen, the present specimen is also mature and hence it appears unlikely that this species reaches any appreciable size. Sclerothyone was especially erected to accommodate only Ludwig and Heding’ s species and hence it still remains monotypic. Perhaps Pawson and Miller’ s (1981) Thyone adinopoda and other similar species could be referred to this genus if the generic diagnosis is amended to also include species with scattered podia.

Subfamily Cladolabinae Heding & Panning Genus Ohshimella Heding & Panning

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Ohshimella ehrenbergii (Selenka, 1868) Figure 6 Urodemas ehrenbergii Selenka, 1868: 14, figs. 6–8. Phyllophorus n. sp. Semper, 1868: 245, pl. 30, fig. 21. Phyllophorus ehrenbergi (sic) Lampert, 1885: 181; Théel, 1886a: 151. Phyllophorus frauenfeldi Ludwig, 1875: 95, fig. 22; H.L. Clark, 1923: 417. Cucumaria turbinata Pearson (non Hutton, 1878: 307), 1903: 189, pl. 1, figs. 2–6; 1910: 169, text figs. 13, 14; Heding and Panning, 1954, 137, text fig. 59 (synonymy). Orcula torrense Helfer, 1913: 433, text figs. 1–7. Urodemella ehrenbergii Deichmann, 1944: 733; 1948: 358. Ohshimella ehrenbergii Heding and Panning, 1954: 133, text figs. 57–59; Thandar, 1989c: figs. 6b, 8 & 9f. Ohshimella ehrenbergi (sic) Clark and Rowe, 1971: 182 (dist.), pl. 30, fig. 5; Cherbonnier, 1988: 216, figs. 94 A–J. ?Urodemas gracile Selenka, 1868: 114; Heding and Panning, 1954: 137.

Type ?Gottingberg Museum. Type locality Red Sea. Previous South African records East coast of KwaZulu-Natal. Material examined SAM-A27903, SE of Mission Rocks, Zululand, 28° 16.4’ S, 32° 31.4’ E, ‘Meiring Naude’, St. ZN 4, R. Kilburn, 10 vi 1988, 30 m, 1 spec; SAM-A27904, off Boteler Point, KwaZulu-Natal, 27° 00.4’ S, 32° 55.2’ E, NMDP, St. ZB 18 6.vi.1990, 66 m, 1 spec; SAM A23175, Coconut Bay, Mozambique, 17.v. 1973, 1 spec. Distribution Indian Ocean, up to 60 m. Habitat Fine sand, rock, stones, coral. Remarks All three specimens in the collection are perhaps juvenile of this well-known Indian Ocean species. The depth of the one from Coconut Bay has not been recorded but the other two come from 30 m and 60 m. The smallest individual is in a poor state of preservation (previously dried up), measures 10 mm x 2.5 mm, and is of greyish-brown colouration; the

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slightly larger specimen is in an excellent state of preservation with the tentacles extended and measures 21 mm x 5 mm and of whitish colouration; the largest measures 34 mm x 12 mm and is of a dull creamish-white colouration. The two smaller specimens show an ambulacral restriction of the podia in 2–3 rows (a juvenile feature), whereas the largest specimen has the podia scattered. In the specimen with extended tentacles there are nine large tentacles in the outer ring and 10 much smaller tentacles arranged in pairs in the inner ring. The calcareous ring is typical of the species (Figure 6K) with the radial prolongations subdivided.

FIGURE 6. Ohshimella ehrenbergii (Selenka, 1868) (Juvenile). SAM-A27903. A. rods from ventral body wall; B. table discs from ventral podia; C. tables from ventral podia; D. tables from ventral body wall. SAM-A23175. E. rods from dorsal body wall; F. rosettes from dorsal body wall; G. plate from dorsal podium; H. rods from dorsal podia; I. rods from tentacles; J. rosettes from tentacles; K. calcareous ring. A-D scale a; E–J scale b; K scale c.

The body wall spicules comprise the typical spinous rods, measuring 67–81 µm (mean 73 µm) in length in SAM-A27903 (Figure 6A) and 54–68 µm in length (mean 59 µm) in SAM A23175 (Figure 6E). In the podia and anal region of the two smallest individuals two types of rudimentary tables are present, both resembling those found in members of the genus Thyone. The first type (Figure 6C) is similar to that found in the podia of the Thyone species — i.e. with curved discs, 157–181 µm in length (mean 168 µm) in SAM-A27903, with one or two perforations at each end and a two-pillared spire, 62–71 µm high (mean 67

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µm), with the pillars uniting at base and ending in paired clusters of teeth. The second type of tables (Figure 6D) resemble those found in the body wall also of some Thyone species — i.e. with quadrilobed and quadrilocular discs, 81–105 µm long (mean 95 µm), with one or two clusters of teeth. Often the tables are reduced to simple discs with a varying number of holes (Figure 6B). Endplates present. Podia of larger specimen (SAM A23175) are supported by rods perforated at one or both ends (Figure 6H), rosette-like rods (64–134 µm, mean 108 µm) (Figure 6F), multilocular plates (132–210 µm, mean 176 µm) (Figure 6G) and endplates (ca 350 µm). Rosettes (18–36 µm, mean 24 µm), which characterise the species, only occur in the body wall of the largest specimen. Tentacle spicules (in SAM A 23175) comprise rosettes (Figure 6J), curved rods of variable length (40–240 µm, mean 99 µm) and thickness (Figure 6I), the smallest ones with usually a single terminal perforation and/or tuberosities, the larger ones with several perforations at each end as well as rosettes (22–38 µm, mean 29 µm). This is the first report of rudimentary tables, of the form here illustrated, present in the body wall of this species. These tables differ from those illustrated by Thandar (1989c) from the anal region of his South African material and by Pearson (1910) from his Querimba specimen. Thandar speculated that the anal tables of this species are perhaps juvenile deposits lost with age. It appears that the rudimentary tables of the body wall suffer the same fate. From the material at hand it appears that the rosettes develop after the loss of tables. In fact, Pearson’ s (1903 & 1910) C. turbinata, respectively from Ceylon and Querimba (Mozambique), suspected by Heding & Panning (1954) and Thandar (1989c) to be a synonym of O. ehrenbergii, showed the presence of rosettes in the larger specimen (48 mm) from Querimba and not in the smaller specimen (25 mm) from Ceylon. Incidentally, Pearson’ s Ceylon specimen presumably possessed no tables, as they are not described. Apparently this is a very variable species whose posterior prolongations of the calcareous ring may or may not be fragmented, and its body wall and podial spicules may or may not include tables and/or rosettes, depending on age, depth or geographic location.

Ohshimella ocula (Cherbonnier, 1988) comb. nov. Figure 7 Phyllophorus (Urodemella) oculus Cherbonnier, 1988 : 222–224, fig. 97 (A–G).

Type MNHNP, 2978. Type locality Mozambique Channel, 20–25 m.

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Previous South African record None.

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Material examined SAM-A27905, off Sodwana Bay, KwaZulu-Natal, 27° 32.8’ S, 32° 42.8’ E, NMDP, St. ZH 19, 2 vi 1990, 77 m, 1 spec.; SAM-A27906, S.E. of Lala Nek, KwaZulu-Natal, 27° 14.1’ S, 32° 49.2’ E, NMDP, St. ZDD 8, 8 vi 1990, 84–97 m, 1 spec. Description Both specimens barrel-shaped; colour in alcohol dirty off-white, slightly darker dorsally. Larger of the two specimens (off Sodwana Bay) approximately 45 mm in length and 23 mm in breadth; smaller specimen (Lala Nek) approximately 20 mm in length and 10 mm in breadth. Podia well developed and of same colouration as body wall — those of the larger specimen scattered in both ambulacra and interambulacra, more numerous ventrally; those of smaller specimen mostly confined to ambulacra in 2–3 rows with a few also in the interambulacra. Mouth anterior, anus posterior. Anal teeth absent, no special anal papillae. Tentacles 20, in two rings of 15+5; those of outer ring large but of unequal size (ventral ones slightly reduced), largest tentacle 10 mm in length; those of the inner circle small. Smaller specimen also with 20 tentacles, those of outer ring comprising 10 large and five small tentacles, the latter alternating with and of same size as the five of inner ring, giving the impression that there are 20 tentacles in two rings of 10+10; large tentacles beige-brown with darker basal branches and tips, smaller tentacles uniformly beige-brown. Body wall soft, slightly rough to the touch. Calcareous ring complex, both radial and interradial plates well developed, somewhat fused basally and not fragmented, the former with a deep anterior notch for the attachment of retractor muscle; interradial plates triangular, slightly shorter than radial plates and slightly concave posteriorly. Posterior prolongations of radial plates short, fragmented but fragmentation often obscured by thick membrane. Calcareous ring of larger specimen, including processes, about 12 mm long. Polian vesicle single, well developed, saccular, 5 mm long. Stone canal single about 3 mm long, straight, anteriorly directed and partially attached to dorsal mesentery. Madreporite spherical and yellowish. Respiratory trees well branched, right one reaching anterior end of body, left one shorter, about half the length of the right one, both uniting before entering cloaca. Cloaca elongate, suspensors well developed. Most gonadal tubules lost in larger specimen, the remaining ones branched. Longitudinal muscles unpaired. Retractors short, arising anteriorly from longitudinal muscles and may or may not bifurcate before inserting on radial plates. Spicules of dorsal and ventral body wall of same form, slightly larger ventrally, comprising numerous rosettes (Figure 7C) and relatively few spinous rods (Figure 7A, B, D), the latter developed as crosses with or without holes, holes when present usually two, rarely one. Spines developed only on arms of cross or only on its main body, or both.

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Rosettes rounded, mulberry-like. Rods of the dorsal body wall of larger specimen 55–82 µm long (mean 69 µm), ventral rods 52–85 µm (mean 65 µm). Dorsal rosettes of larger specimen 23–45 µm (mean 32 µm), ventral rosettes 19–33 µm (mean 29 µm). Podial deposits include rosettes, scarce tables and multilocular plates, the latter surrounding large endplates. Rosettes (Figure 7F) 16–40 µm (mean 28 µm); multilocular plates (Figure 7E) 145–250 µm (mean 210 µm), of variable form, perforated by 8–35 holes; endplates 400–500 µm in diameter (Figure 7G), with no regular arrangement of large and small holes. Tables rare, delicate (Figure 7H), reduced, only four observed in about 10 preparations of podial deposits. Tentacle deposits include rosettes (Figure 7J) and slightly curved rods (Figure 7I). Rods 48–145 µm (mean 98 µm) long, with usually one, sometimes more, terminal perforations, larger ones with also terminal digitations/ tuberosities; larger rods in tentacle shaft, smaller ones in the terminal branches where rosettes predominate. Rosettes 22–58 µm (mean 42 µm). Introvert deposits include minute rods (Figure 7K), 38–90 µm (mean 63 µm) and rosettes (Figure 7L), 30–40 µm (mean 36 µm). Smaller specimen with identical spicules but no introvert deposits. Distribution South-West Indian Ocean. Habitat Dead coral and rubble; coarse sand, muddy sand.

FIGURE 7. Ohshimella ocula (Cherbonnier, 1988). SAM-A27906. A. rods from body wall; B. rods/crosses from same; C. rosettes from ventral body wall; D. side view of cross from body wall; E . plates from dorsal podia; F. rosettes from dorsal podia; G. part of dorsal end-plate; H. table from ventral podium; I. rods from tentacles; J. rosettes from tentacles; K. rods from introvert of SAMA27905; L. rosettes from introvert of SAM-A27905. A–F & H–L scale a; G scale b.

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Remarks The two specimens described above correspond well with Cherbonnier’ s (1988) description of the type of Phyllophorus (Urodemella) oculus from the Mozambique Channel. On the basis of its spicules the species clearly falls in the genus Ohshimella and not Phyllophorus as it lacks table deposits of the type found in the latter genus. This is supported by Yves Samyn (pers. com.). Perhaps Cherbonnier was misled by some similarities in the spicules of his new species with that of P. (U.) brocki (Ludwig), which precedes his description of P. oculus. However, in the former species the deposits are clearly tables or table derivatives, which is not the case in O. oculus. Incidentally, both species have a more or less similar calcareous ring, rosettes and tentacle rods. What is more surprising is that Cherbonnier states that his new species bears some affinities with Phyllophorus purpureopunctata (Sluiter, 1901). However, the spicules of both these species are remarkably different. Regrettably Cherbonnier did not compare his P. oculus with his Ohshimella castanea Cherbonnier, 1980 from New Caledonia. Although both these species may not be synonymous there is a striking resemblance in their body wall, podial and tentacle spicules. In fact, the podial and tentacle deposits of both are identical. I examined the holotype of O. castanea, received from MNHNP, and am of the opinion that there are sufficient differences between the two species to keep them separate. However P. (U.) oculus is here transferred to the genus Ohshimella.

Family Cucumariidae Ludwig Subfamily Cucumariinae Ludwig Genus Pawsonellus Thandar Pawsonellus africanus Thandar, 1986 Pawsonellus africanus Thandar, 1986: 485, fig 1 a–n; 1991:136, figs. 2c, 15d.

Type SAM A-23442. Type locality 16 km south of Durban, 36 m. Previous record Type locality only.

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Material examined SAM-A27907, Park Rynie, KwaZulu-Natal, K. S. Ganga, March 1984, LWS, 1 spec. Distribution Southern KwaZulu-Natal, South Africa, 0–36 m. Habitat Reef sand, seaweed, bryozoans. Remarks This species was based on numerous specimens taken from subtidal waters, south of Durban. This is the first record of the species from intertidal waters, also south of Durban. The species is well characterised by its ambulacral restriction of the podia in two rows, the two mid-ventral tentacles reduced to stumps with bifurcating tips, and multilocular, platelike body wall spicules.

Subfamily Colochirinae Panning Genus Trachasina Thandar & Moodley Trachasina crucifera (Semper, 1869) Figure 8 Cucumaria crucifera Semper, 1869: 121, pl. 1, fig. 1 a–c; Ludwig, 1887:1235, pl. 1, 5–11; Deichmann, 1948: 343, pl. 18, fig. 1–3. Trachythyone crucifera Panning, 1949: 427, fig. 15; Clark & Rowe, 1971: 182 (distrib.), fig. 92; Thandar, 1991: 131, fig. 10, 15a (synonymy & records). ?Trachythyone crucifera Panning, 1964: 170, fig. 9; 1971: 37, 40 (passim). Trachasina crucifera Thandar & Moodley, 2003: 284 (passim).

Type Hamburg Museum, Germany, 2760. Type locality Aden, Red Sea. Previous South African record Southern KwaZulu-Natal. Material examined SAM-A27908, Cracker Reef, Aliwal Shoal, KwaZulu-Natal, D. Herbert, SCUBAdive, 30. iv. 1989, ±23 m., 1 spec.(juv.). 26


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Distribution West Indian Ocean, from South East Arabia to the east coast of South Africa, including Madagascar and islands of the West Indian Ocean, up to 23 m. Thandar’ s (1991) record of the species occurring in the Bay of Bengal is erroneous and should be discounted. Habitat Coral and rock. Remarks From the available literature it appears that this is perhaps the deepest record yet for this well-known West Indian Ocean species, which has been well re-described by Panning (1949, 1964) and many other workers and hence does not require a detailed description. Thandar (1991) remarked on the species and Thandar & Moodley (2003) transferred the species to the monotypic genus Trachasina erected for this purpose. The specimen, here examined, is obviously a juvenile, judging from its size, distribution of podia and immaturity of the gonad. It is whitish in colour, including the podia and tentacles, and measures only 18 mm in length. Podia are confined to the ambulacra in single or double rows and are much longer ventrally. The 10 tentacles are of unequal length. The calcareous ring is as previously illustrated for the species (see Thandar 1991: fig. 10i and Cherbonnier 1988: fig. 10g).

FIGURE 8. Trachasina crucifera (Semper, 1869). SAM-A27908. A. crosses from body wall; B. rosettes from base of dorsal podia; C. plate of dorsal body wall; D. plates and rods from dorsal podia; E. end-plate from dorsal podium; F. rosettes from tentacle tips and branches; G. small rods from tentacle tips and branches; H. large rods from tentacle stalk. All drawn to same scale.

The body wall spicules are typically spinous crosses (Figure 8A), 65–120 µm long (mean 88 µm) and thick, multilocular plates (Figure 8C), 205–410 µm long (mean 300 HOLOTHUROIDEA


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µm) and 140–230 µm wide (mean 176 µm), with up to 16 holes. Rosettes described for the type by Semper (1869) and perhaps over-looked by other writers or not present in their material, were detected at the base of the podia, but are scarce (Figure 8B). Podial deposits include rods with several perforations at the ends and/or in the expanded middle (Figure 8D). The pedicel endplates (Figure 8E) are small, of same size dorsally and ventrally, with 3–4 series of holes, larger medially and smaller peripherally. The tentacle shaft comprises slightly curved rods (Figure 8H), 127–337 µm long (mean 210 µm), with terminal perforations, while smaller, more delicate rods (Figure 8G), 3–113 µm long (mean 48 µm), and rosettes (Figure 8F), 10–47 µm long, (mean 26 µm), are present in the branches. No deposits, with the exception of a few rosettes, were detected in the introvert. It appears that rosettes of the body wall are juvenile features and so are also the large-size and multilocular nature of the plates.

Genus Plesiocolochirus Cherbonnier Plesiocolochirus dispar (Lampert, 1889) Figure 9 Colochirus dispar Lampert, 1889:820, fig. 4; Ekman, 1918:32. pl. 3, fig. 25, pl. 4, fig. 26. Pentacta dispar Clark & Rowe, 1971 :180 (dist.); Cherbonnier, 1988:164, fig. 68. Colochirus gravieri Vaney 1905:187. Pentacta gravieri Cherbonnier, 1955:162, pl. 48, figs. k–s. Pentacta trimorpha Clark, H.L. 1921:171, pl. 37 (1–8). Plesiocolochirus dispar Rowe (in Rowe & Gates 1995):278 (syn.)

Type Syntypes perhaps in Stuttgart, Germany. Type locality Mermaid Strait, northwest Australia. Previous southern African record None. Material examined SAM - A23175, Coconut Bay, Mozambique, 17.v.1973, 1 spec. Description Form barrel-shaped, with dorsal surface arched and ventral flattened, sole-like, but sole not clearly defined; anterior end broad, posterior narrow and turned upwards. Length about 10 mm, diameter of anterior and posterior ends 5 mm and 2.5 mm respectively. 28


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Colour uniformly white, including podia and tentacles. Podia restricted to ambulacra in double rows but most of those of mid-ventral ambulacrum retracted, except at anterior and extreme posterior ends; those of ventro-lateral ambulacra mostly well extended; dorsal podia indistinct because of scales which envelope most of dorsum. Ventral podia with well-developed suckers. Mouth star-shaped, boarded by five, distally calcified valves, especially the mid-ventral one which appears as a tooth-like structure. Tentacles 10, midventral two reduced. Anus terminal, surrounded by five teeth, each flanked by subterminal podia. Skin thin and rough to the touch.

FIGURE 9. Plesiocolochirus dispar (Lampert, 1889).SAM-A23175. A. dorsal body wall scales; B. small-knobbed buttons; C. large-knobbed buttons; D. part of end-plate from ventral podia; E. closed baskets from dorsal body wall; F. smooth plates from dorsal podium; G. knobbed plates from dorsal podium; H. small rods from tentacles; I. rosettes from tentacles; J. large rods from tentacle stalk; K. madreporite; L. calcareous ring. B–H & J scale a; I scale b.

Calcareous ring (Figure 9L) simple, well calcified, radial and interradial plates fused, radial plates with a bifid anterior prolongation for attachment of retractor muscle and a notched posterior surface; interradial plates shorter than radial plates, with a triangular anterior end and slightly indented posterior surface. Polian vesicle single, sac-like; stone canal short, free; madreporite (Figure 9K) spherical, grooved, lodged in the posterior notch of mid-dorsal interrradial plate. Respiratory trees delicate, extremely transparent but well branched, right tree reaches about one third the body length from anterior end, left one shorter. Gonad not observed. Longitudinal muscles unpaired, thin, well developed anteriorly, fused to body wall for most of their length. Retractor muscles arise as single

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strands from the anterior third of the longitudinal muscles. Dorsal body wall invested in imbricating scales and, besides these, include deposits of four types: multi-layered buttons with large knobs; simple buttons with small knobs; usually complete baskets; and developing spicules of various shapes and sizes. Scales (Figure 9A) rather complex, oval, 105–875 µm long; small-knobbed buttons of various shapes and sizes (Figure 9B), 48–65 µm (mean 56 µm); large-knobbed buttons (Figure 9C), 58–71 µm (mean 63 µm), with four or more holes; baskets characteristic, flat (Figure 9E), 39–65 µm (mean 51 µm), with four large and four smaller alternating holes, surface spinose. Dorsal podia supported by perforated rods and plates (Figure 9F), 84–165 µm (mean 94 µm), some with marginal tuberosities (Figure 9G) and reduced endplates, 90–145 µm (mean 132 µm). Ventral podia with similar deposits but larger endplates, 170–240 µm (mean 202 µm) (Figure 9D). Tentacles supported by elongated, perforated rods, 42–258 µm (mean 107 µm), smaller in the branches (Figure 9H) and larger in the stalk (Figure 9J), as well as rosettes (Figure 9I), 16–32 µm (mean 24 µm). Distribution Tropical Indo-West Pacific, up to 20 m. Remarks The single specimen was at first thought to represent Psolidium ornatum (E. Perrier, 1893). However, Dr Rowe (pers. comm.) drew my attention to Plesiocolochirus dispar (Lampert), a species which he thinks the specimen represents. I checked several descriptions of P. dispar and its synonyms and I concur with Dr Rowe without reservation. P. dispar has been recorded from the Red Sea and Somalia (see Clark & Rowe 1971) but this is its first record from southern Africa. However, the species was reported from Tuléar and Nosy Bay (Madagascar) by Cherbonnier (1988) who also provides a detailed description of the species. The current specimen agrees well with that material.

Plesiocolochirus tessellara (Cherbonnier, 1970) comb. nov. Figure 10 Pentacta tessellara Cherbonnier, 1970:282, fig. 2 (A–O); Thandar, 1991:123, fig. 8 (A–J). Colochirus minutus Macnae & Kalk, 1958:130 (non Ludwig, 1875).

Type MNHNP. Type locality Mouth of Rio Coche, near Inhambane, Mozambique, 3–5 m, sand, gravel and shells. Previous South African record None 30


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Material examined SAM-A27909, Cracker Reef, Aliwal Shoal (KwaZulu-Natal), D. Herbert, SCUBAdive, 30.iv.1989, ±23 m., 2 spec.(juv.). Distribution Only known from Mozambique and east coast of South Africa. Habitat Coral, rock. Remarks This species was assigned to Pentacta by both Cherbonnier (1970) and Thandar (1991). However, Rowe (in Rowe & Gates 1995) restricted the genus Pentacta to include only the type species, Actinia doliolum Pallas, 1766 and perhaps Cucumaria capensis Theel, 1886a. He characterised the genus as having podia restricted to the ambulacra ventrally but scattered dorsally, ten tentacles of more or less equal size and spicules comprising multilocular, knobbed plates and shallow, rugose baskets with basic quadrilocular design. These characters still present a problem as in some specimens of P. doliolum the two ventral tentacles or some others may be reduced and the podia in C. capensis are confined to the ambulacra and not dorsally scattered. However, both these are endemic southern African temperate species (shallow-water and deep-water respectively), not known to extend into the warmer subtropical waters of KwaZulu-Natal. P. tessellarus, on the other hand, is from tropical-subtropical waters and has the ventral surface somewhat sole-like with prominent podia, the ventral-most two tentacles reduced and a brittle body wall packed with spicules comprising multi-layered scales, nodular perforated plates and simple shallow baskets. These characters accord well with those of Plesiocolochirus enunciated by Rowe (in Rowe & Gates 1995). However, the genus Plesiocolochirus was initially erected for only the type species, P. spinosus (Quoy & Gaimard) characterised by evenly distributed radial and interradial podia, latero-ventral tubercles and radial plates of the calcareous ring bearing distinctive paired posterior prolongations(see Cherbonnier 1946; O’ Loughlin & Alcock 2000). Hence the generic placement of P. tessellarus in Plesiocolochirus is problematic. It is here only tentatively referred to this genus until a more suitable genus is found to accommodate it. This species, originally known only from Mozambique, is well described by Cherbonnier (1970). Thandar (1991) examined a paratype of the species and provided measurements of its spicules. The current material is the first record of the species from South Africa. The two specimens here examined are obviously juvenile, judging from their size (28 mm & 35 mm), distribution of podia, state of development of the tentacles and calcareous ring, and the absence of the gonad. Both are white in alcohol with a yellowish tinge at the anterior end of the larger specimen, and hence differing in colour

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from that recorded for the species by Cherbonnier (!970) and Thandar (1991) - perhaps due to depth, age or preservation. The tentacles are retracted in both specimens, the anus encircled by minute teeth in the larger specimen, and the podia restricted to the ambulacra, better developed ventrally in 1–2 rows per ambulacrum but in single row in each dorsal ambulacrum. The skin is rough to the touch and the imbricating plates easily visible with a stereoscope.

FIGURE 10. Plesiocolochirus tessellarus (Cherbonnier, 1970). SAM-27909. A. large-knobbed buttons from dorsal body wall; B. side view of buttons; C. plate/scale from dorsal body wall; D. baskets from dorsal body wall; E. side view of baskets from dorsal body wall; F. baskets from dorsal podia; G. rods from dorsal podia; H. small-knobbed plates from dorsal podia; I. end-plate from dorsal podium; J. part of end-plate from ventral podium; K. plates from tentacles; L. tentacle rods. A-C & G-L scale a; D-F scale b.

The body wall spicules comprise baskets (15–25 µm, mean 19 µm), always provided with small nodules on the surface and a spinous or “jagged” rim (Figure 10D, E); buttons (75–95 µm, mean 86 µm) that are small, medium to large- knobbed (Figure 10A, B); and large (405–475 µm, mean 433 µm) imbricating, multi-layered, knobbed plates/scales (Figure 10C). The podia are supported by: elongated, slightly curved rods (112–287 µm, mean 199 µm) with several perforations and often with a distinct knobbed surface (Figure 10G); baskets (16–29 µm, mean 20 µm), similar to those of the body wall (Figure 10F); and perforated smooth to knobbed plates (39–116 µm, mean 65 µm) (Figure 10H). The dorsal endplates are circular with large central holes and a single series of peripheral holes 32


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(Figure 10I), whereas the ventral endplates are larger (136 µm cf. 120 µm), multi-lobed (Figure 10J), with similar arrangement of holes except that instead of a single series of holes on the periphery there are numerous small holes in the lobes. The tentacle stalk contains elongated, delicate rods (39–90 µm, mean 65 µm) with few holes (Figure 10L) whereas perforated plates (87–184 µm, mean 154 µm) occur in the branches (Figure 10K). No rosettes were detected in the tentacles, nor any deposits in the introvert. The present specimens differ from the type material in size, colouration, distribution of podia, poor calcification of the calcareous ring, absence of rosettes in the tentacles and lack of spicules in the introvert. These can be attributed to the immaturity of the material and hence here considered to be of no taxonomic significance. Thandar (1991) commented that this species comes quite close to Pentacta minuta (Ludwig) (junior subjective synonym of Plesiocolochirus australis according to Rowe (in Rowe & Gates 1995), originally described from North Australia but also recorded by Macnae and Kalk (1958) from Inhaca Island (Mozambique) as Colochirus minutus, on material identified but not described by Cherbonnier. Regrettably this material is no longer extant. Hence, P. minuta (Macnae & Kalk, 1958, non Ludwig, 1875) is here relegated to the synonymy of P. tessellarus.

Genus Staurothyone H.L. Clark Staurothyone rosacea (Semper, 1869) Figure 11 Thyone rosacea Semper, 1868: 242 (nom.nud.); 1869: 122, pl.1, fig. 2 a–c; Lampert, 1885: 159, pl. 1, fig. 3. Thyone sargassi Lampert, 1889: 840, fig. 10. Staurothyone rosacea Panning, 1949: 418, fig. 5; 1964: 159, fig. 1.

Type Hamburg Museum, Germany, 2901. Type locality Red Sea. Previous South African record None Material examined SAM-A27910, between Bhanga Nek and Kosi Bay, No. 13 reef, KwaZulu-Natal, D. Herbert, SCUBA-dive, 12–20 vii 1987, 6–18 m, 1 spec.

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Description Specimen previously dried up but prolonged soaking in alcohol softened the body wall to some extent. Form cylindrical, slightly U-shaped, measuring 22 mm along trivium and 3.5 mm in breadth in mid-body. Colour dark reddish brown ventrally, paler dorsally. Podia usually restricted to ambulacra, two rows dorsally and 3–4 rows ventrally, with few also scattered in interambulacra of ventral surface. Mouth anterior, tentacles extremely bushy but number and size variation could not be determined. Anus terminal, encircled by five calcareous teeth, easily demonstrated. Calcareous ring (Figure 10C) well developed, interradial plates longer than radial plates with a shallow depression on their outer surface. Polian vesicle single, sac-like. Stone canal not observed. Spicules of body wall include large crosses (Figure 11A) and minute rosettes (Figure 11B). Podial deposits include rods with digitated ends. No indication of any secondary branchings to the two primary dichotomous branches of the crosses. Distribution West Indian Ocean, including Red Sea.

FIGURE 11. Staurothyone rosacea (Semper, 1869). SAM-A27910. A. crosses from body wall; B. rosettes from body wall; C. calcareous ring.

Remarks Of the three other species currently classified in Staurothyone, S. vercoi (Joshua & Creed, 1915) is apparently a 20-tentacled form and hence referable to another genus (see Rowe & Gates 1995); S. distincta (Clark, 1938) from North Australia is distinct in possessing, in addition to the crosses and rosettes in the body wall, peculiar minute ‘oblong particles’; and S. inconspicua (Bell, 1887) is a South Australian-Tasmanian species, very close to the Indo-West Pacific S. rosacea and suspected by both Clark (1938) and Panning (1949) to be synonymous with it. However, according to O’ Loughlin (1991) and Materia et al. (1991), S. inconspicua is a seasonal coelomic brooder, while Rowe (pers. comm.) is of the opinion that there are also differences in proportions of crosses in the two species. The current specimen differs from the holotype, re-described by Panning (1949, 1964), in the ambulacral restriction of the podia, the shape of the plates of the calcareous ring, especially the anterior tip of the interradial plates, and the posterior margin of the ring. These are perhaps growth characteristics since the type (40 mm long) is almost twice as large as the specimen here described.

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Family: Psolidae Ed.Perrier

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1245 Genus Psolidium Ludwig Psolidium multipes sp. nov. Figure 12 Diagnosis Small species, holotype 11 mm long; colour in alcohol white. Sole not well defined; oral valves indistinct. Tentacles ?10, ventral two reduced. Anal teeth present. About 10 scales between mouth and anus, 8–9 laterally. Dorsal podia distinct, emitting through and between scales; ventral podia crowded, 4–5 rows in each ventro-lateral ambulacrum and 3–4 rows in midventral ambulacrum. Dorsal scales imbricating, complex, multi-layered, up to 1.5 mm long; other dorsal deposits include buttons, baskets and rosettes. Buttons knobbed, 68–84 µm, with 4–6 holes, knobs of various sizes, central ones often united by a half-ring on one side; baskets mostly incomplete; rosettes minute, mulberry-like. Ventral scales also complex but smaller than dorsal ones, up to 660 µm long; other ventral deposits include knobbed buttons, up to 94 µm, with three or more holes and mostly complete baskets and rosettes. Type SAM-A23175. Type locality Coconut Bay, Mozambique. Material examined Holotype only, collected 17.v.1973. Etymology The specific name refers to several rows of well-developed podia both dorsally and ventrally. Description Specimen small, length approximately 11 mm, width of mid-body about 5 mm. Colour beige-white mid-dorsally, white elsewhere, including oral and anal ends. Dorsal surface arched, ventral flattened, but sole not clearly defined. Mouth anterior, oral valves indistinct but mouth surrounded by scales appearing as pairs in the five radii; buccal membrane distinct. Tentacles nine, bushy, white, few protruding through mouth, seven well developed, mid-ventral two reduced. Anus situated on a dorsally directed cone, encircled by five calcareous teeth. Dorsal and lateral surfaces invested in large, imbricating scales

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covered with granules. About 10 scales mid-dorsally, between mouth and anal cone, 8–9 scales laterally. Dorsal podia small, distinct, numerous, scattered, with about 2–3 emitting from each scale but most between scales, spreading to the dorso-lateral radii where they are smaller and fewer. Ventral podia retracted, crowded, in 4–5 rows on each ventro-lateral radius and 3–4 rows in mid-ventral radius, increasing to five mid-ventrally. Ventral suckers well developed, dorsal less so.

FIGURE 12. Psolidium multipes sp. nov. Holotype. SAM-A23175. A. scale from dorsal body wall; B. large-knobbed buttons from ventral body wall; C. small-knobbed buttons from ventral body wall; D. plates from ventral podia; E. rosettes from ventral body wall; F. closed baskets from ventral body wall; G. open baskets from dorsal podia; H. rods and plates from tentacle branches and tips; I. rods from tentacle stalk; J. end-plate from ventral podium; K. stone canal (frontal and lateral view); L. calcareous ring. B–J scale a.

Calcareous ring (Figure 12L) well calcified with radial and interradial plates simple, of more or less the same length, the former anteriorly bifid with a concavity for insertion of radial muscle; all plates with a posterior concavity but no posterior processes. Polian vesicle single, bulbous, arising ventrally; stone canal short, only slightly coiled; madreporite (Figure 12K) poorly calcified, lodged in the posterior notch of mid-dorsal interradial plate. Respiratory trees delicate, well branched, both trees extending to anterior end. Gonad (testis) as two tufts of unbranched, white, sausage-like tubules, filling most of body cavity; gonadal stolon short, lodged in dorsal mesentery. Longitudinal muscles unpaired, retractor muscles arise from anterior third of longitudinal bands.

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Dorsal scales (Figure 12A) large, up to 1.5 mm long, complex, multi-layered, those surrounding mouth elongated, specialised, up to 1.3 mm long; other dorsal deposits include buttons, baskets and rosettes. Buttons of three types: robust ones, 84–113 µm (mean 95 µm), pierced by 4–6 holes, with large knobs and with central knobs developed as a half-ring; other robust buttons, 68–84 µm (mean 76 µm), perforated by usually four (sometimes 2–5) holes, also with large knobs but without half-ring; remaining buttons small, ca. 70 µm, fewer, small- knobbed and usually perforated by four holes and with varying number of knobs. Baskets in the form of spinose, often cup-like crosses (Figure 12G), with spines usually restricted to the branches; complete baskets not detected amongst dorsal deposits. Rosettes minute, mulberry-like. Ventral deposits also comprise complex, multi-layered scales, much smaller (194–660 µm, mean 395 µm) than the dorsal ones, and buttons, baskets and rosettes. Buttons usually small-knobbed (Figure 12C), 45–74 µm long (mean 57 µm), with four or more holes and a varying number of knobs; other knobbed buttons fewer but slightly larger (58–94 µm, mean 73 µm), with large knobs and three or more holes (Figure 12B). Ventral baskets (26–48 µm, mean 38 µm) mostly complete (Figure 12F), with spinose margins and surface, few may be incomplete and similar to those of dorsal body wall. Ventral rosettes (Figure 12E) 16–39 µm (mean 25 µm), similar to those of dorsal body wall but commoner. Podial deposits (Figure 12D) comprise marginally-knobbed, perforated plates and smooth rods (ventrally 87–184 µm, mean 130 µm; dorsally 74–110 µm, mean 91 µm); dorsal endplates reduced, ventral endplates (Figure 12J) larger (77 µm cf. 187 µm). Tentacle branches with minute plates and perforated rods of varied shapes and sizes (36–187 µm, mean 93 µm) (Figure 12H), as well as both open and closed rosettes (10–29 µm, mean 17 µm). Tentacle stalks supported by larger rods (265–426 µm, mean 349 µm)(Figure 12I). Deposits of the buccal membrane comprise large-knobbed rods (ca 185 µm), perforated with several holes, baskets similar to those of the dorsal body wall but smaller (26–29 µm, mean 27 µm), and closed rosettes (13–26 µm, mean 20 µm). No spicules detected in gonad. Distribution Type locality only. Remarks In the presence of dorsal podia emitting through or between scales, the distribution of the ventral podia and the presence of four distinct types of body wall deposits besides the scales, this species appears unique. Its body wall deposits come quite close to Psolidium ornatum (Ed. Perrier, 1893), re-described by Cherbonnier (1988) from designated neotype, but the new species differs in its distribution of podia. Cherbonnier (1988) describes ventro-lateral ambulacra with two rows of podia and the mid-ventral ambulacrum with three rows. Further, in P. ornatum the plates of the calcareous ring appear separate while in the new species, they are partially fused. Hence, the new species appears to be very close

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to P. ornatum but not identical with it. The only other species where the podia are arranged in crowded bands is P. nigrescens Clark, 1938 from south-east Australia and P. dorsipes Ludwig (1887) from Chile and Patagonia. However, the dorsal deposits of the former species are stated to be scarce and also include numerous triradiate bodies. Such bodies are also found in the sole together with knobbed, perforated plates unlike those found in the new species. In P. dorsipes, on the other hand, there are tower-like deposits in the dorsal body wall and the baskets are of a different kind.

Psolidium acorbulum sp. nov. Figure 13 Diagnosis Minute, sub-globose species, holotype (almost mature) 4 mm long, with distinct ventral sole; oral valves absent, scales completely concealing mouth and anus. About 10–12 scales between mouth and anus, about 12 scales laterally. Tentacles eight, fingershaped, without lateral branches. Dorsal podia distinct, emitting from between scales. Sole bordered by a single row of podia laterally, mid-ventral ambulacrum without podia except a couple both anteriorly and posteriorly. Dorsal scales imbricating, up to 342 µm long, simple, consisting of a single layer of calcareous material, multilocular, smooth or slightly knobbed; knobs, when present, restricted to one side of plate. Sole deposits exclusively knobbed plates or buttons, up to 133 µm long, with 4–5 (sometimes 2–10) holes. Baskets and rosettes absent. Material examined Holotype SAM A-A27911; Paratype SAM-A27912. Type locality Between Port St. Johns and East London, 32° 28.6’ S, 28° 58.8’ E, R.V. ‘Meiring Naude’, St. SM 226, 24.vi.1979, 710–775 m. Etymology The specific name refers to the absence of basket-like deposits from the body wall, including the sole. Description Both specimens minute, sub-globose, but distinctly psolid-like (Figure 13M) with well-arched dorsal surface and a flattened ventral surface forming a distinct sole. Oral cone not distinct, no oral valves. Scales completely conceal both oral and anal apertures. Largest specimen (holotype) 4 mm long; the other (paratype) 3.5 mm, width of mid-body of both approximately half body length. Colour, in alcohol, yellowish-white. Dorsal and 38


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lateral surfaces covered by simple, imbricating scales, about 10–12 scales between oral and anal apertures, decreasing in size posteriorly. Only eight tentacles detected, of unequal length, finger-shaped, without lateral branches. Dorsal podia distinct, non-retractile, about 50, scattered, emitting from between scales and appearing as spire-like projections from body wall. About 12 scales laterally. Sole covered by thin, translucent membrane extending from just behind the anterior end to extreme posterior end, boarded by a single row of about 10 podia on each side and two anteriorly. Mid-ventral ambulacrum without podia, except for the two anterior ones and those of the lateral series forming an arc posteriorly. Ventral podia better developed than dorsal and with well-formed suckers.

FIGURE 13. Psolidium acorbulum sp. nov. Holotype. SAM-A27911. A. scale from dorso-lateral body wall; B. scale from dorsal body wall; C. plates from ventral body wall; D. plates from dorsal podia; E. rods/plates from ventro-lateral podia; F. end-plate from the ventro-lateral podium; G. rods from tentacle stalk; H. rods from tentacle branches and tips; I. calcareous ring showing mid-ventral radial plate; J. dorsal radial plate; K. single tentacle; L. polian vesicle and stone canal; M. Paratype. SAM-A27912, entire. A & B scale a; C–H scale b; I–K scale c.

Calcareous ring (Figure 13I, J) simple but well developed, delicate. Radial plates with a triangular anterior projection, a depression for attachment of retractor muscle and a slightly concave posterior margin; interradial plates of much the same form. Polian vesicle single slightly elongated; stone canal short, madreporite funnel-like (Figure 13L), attached to dorsal mesentery. Gonad developed as three unbranched, slightly coiled tubules, attached to dorsal mesentery at anterior end, in process of maturity. Respiratory trees simple, short with few terminal branches. Spicules of dorsal body wall only comprise simple, single-layered, imbricating scales (Figure 13A, B), up to 342 µm, perforated by numerous holes and with or without a HOLOTHUROIDEA


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slightly-knobbed surface; knobs, when present, restricted to one end of plate. Spicules of sole exclusively knobbed plates or buttons (Figure 13C) 71–133 µm long (mean 98 µm), with usually 4–5 (sometimes 2–10) holes, with an uneven, spinous margin; knobs when present minute and rather faint. Ventral podia with well-developed endplates (ca. 97 µm) (Figure 13F) and other densely packed plates (Figure 13E), 73–123 µm long (mean 89 µm) with up to eight holes and a spinose or slightly knobbed margin; endplates may also bear one or two tiny knobs on surface. Dorsal podia with similar deposits (Figure 13D). Tentacle stalk supported by rods and plates (Figure 13G), 90–198 µm long (mean 134 µm), the former more numerous, frequently curved and with several holes, usually restricted to the ends but sometimes also in the middle, which may bear a short arm, with or without perforations; plates few, restricted to the terminal end, with a spinose or slightly-knobbed margin and pierced by up to 10 holes. Tentacle branches supported by rods of varying shapes (Figure 13H). Distribution Type locality only. Remarks In its size, simplicity of the tentacles and dorsal plates, presence of a single row of marginal podia on each side of the sole, and the absence of mid-ventral tube feet in combination, this species differs from all nominal species of the genus. In the nature of its spicules it comes quite close to Psolidium vitreum Ohshima from the north-east Pacific and to the Psolidium sp. (Theel, 1886b) from the Mexican Gulf. It, however, differs from both these species by the simplicity of its tentacles and the presence of a single, instead of a double row of marginal podia. Thandar (1999) hesitantly identified a single specimen from the south-west coast of South Africa as P. vitreum despite the fact that it had only a single row of marginal podia and no mid-ventral podia, and also referred Thėel’ s (1886b) Psolidium sp. to the synonymy of P. vitreum, thus supporting the suspicions of Ohshima (1915). This view can no longer be upheld as the writer is now of the opinion that the distribution of podia is an important taxonomic feature in Psolidium species. It is therefore possible that Thandar’ s P. vitreum is perhaps another species similar but not identical to the new species here described as it possesses bushy tentacles, complex multi-layered dorsal scales and more pronounced knobs on the ventral buttons. Rowe (pers. comm.) thinks that the new species, because of its simple, finger-like tentacles, should be referred to the order Dactylochirotida, perhaps to Psolidothuria, or it may represent a young Ypsilothuria. However, this is inadmissible as the form of the body and the presence of a distinct sole with marginal podia are psolid characters. Further, the gonad appears too well developed for the specimen to be a young Ypsilothuria. Incidentally, Pawson (1971) also described digitate rather than dendritic tentacles in the psolid Ekkentropelma brychia and Pawson & Valentine (1981) described weakly dendritic

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to unbranched tentacles in juvenile of Psolidium prostratum. Simple, weakly developed dendritic tentacles are also present in the new Ekkentropelma species described below. E vidently then the psolids may bridge the gap between dendro- and dactylochirotid holothuroids.

Genus Ekkentropelma Pawson Ekkentropelma groovia sp. nov. Figure 14 Diagnosis Small U-shaped species, mature holotype 8.75 mm along ventral surface, oral or anal cones absent. Colour in alcohol yellowish-white. Sole distinct, enclosed within groove that is displaced slightly to right, extending from just behind the mouth to terminate just in front of anus. Dorsal podia few, apparently restricted to ambulacra and emitting from between scales. Podia of sole larger, fewer, 3–5 on each side. Anal teeth minute. Tentacles about 10, finger-shaped, with minute lateral branches. Dorsal body wall encased in simple, smooth or slightly knobbed, one-layered, imbricating scales, up to 380 µm long, and much smaller, up to 220 µm long, smooth or slightly knobbed plates with up to 10 holes and often with one end slightly prolonged; baskets absent. Type SAM A-27913. Type locality Between Port St. Johns and East London, 32° 28.6’ S, 28° 58.8’ E, R.V. ‘Meiring Naude’, St. SM 226, 24.vi.1979, 710–775 m. Material examined Holotype (female). Etymology The specific name is with reference to the groove in which the sole is embedded. Description Specimen (Figure 14I) small, somewhat U-shaped, cylindrical, without oral and anal cones, mouth and anus terminal. Mouth not bordered by valves; tentacles retracted; anus terminal but downwardly directed. Length along ventral surface 8.75 mm, along dorsal surface 8 mm, width in mid-body 2 mm, anterior and posterior ends narrower, more so the latter. Colour uniformly yellowish-white. Dorsal surface slightly arched, invested in HOLOTHUROIDEA


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imbricating scales, which extend laterally and also ventrally. Sole distinct, enclosed within a groove that is displaced slightly to the right, extending from about 2 mm behind the mouth to terminate about 2 mm in front of the anus; covering membrane of sole damaged. Dorsal podia few, poorly developed, emitting between scales, restricted to the radii, about 10 per radius; sole bordered by larger podia, five on the right and three on the left; with the one on each end of the groove extremely well developed. Anal teeth minute.

FIGURE 14. Ekkentropelma groovia sp. nov. Holotype. SAM-A27913. A. plates from dorsolateral body wall; B. plates from sole; C. plates and rod from dorso-lateral podia; D. plates and rods from ventral podia; E. end-plate from ventral podium; F. rods from tentacles; G. single tentacle; H. calcareous ring and associated structures; I. whole animal. A–F scale a; G & H scale b.

Calcareous ring (Figure 14H) simple, well calcified but delicate; radial plates triangular with an anterior, slightly bifid projection and a shallow depression for the retractor muscle, posterior surface slightly concave; interradial plates only slightly shorter but with deeper posterior concavity. Tentacles nine, of unequal length - five large, a single dorsal one of medium size, three ventral remarkably reduced; all tentacles finger-shaped (Figure 14G) with minute lateral branches. Polian vesicle (Figure 14H) single, tubular; stone canal remarkably short; madreporite poorly calcified, form difficult to determine. Gonad (ovary) (Figure 14H) developed as two sacciform tubules attached to dorsal mesentery in anterior part of gut, each tubule full of ripe eggs. Retractor muscles originate from anterior third of longitudinal muscle bands. Dorsal and lateral body wall covered with imbricating scales and much smaller perforated plates (Figure 14A). Scales large, up 380 µm, single-layered, smooth,

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occasionally with 1–5 minute knobs; perforated plates, 150–220 µm (mean 180 µm), of irregular shapes, either smooth or faintly-knobbed and with up to 10 holes, commonly less, sometimes with one end slightly prolonged into a branched projection. Sole deposits small plates (Figure 14B), 104–170 µm (mean 140 µm), perforated by 5–10 holes, similar in size and shape to those of dorsal body wall, but with nodules restricted to usually one side of plate which is devoid of handle-like projection. Dorsal podia densely packed with minute, 73–130 µm (mean 97 µm), plates and rods (Figure 14C), usually with two or more perforations, occasionally none, endplates absent; ventral podia with endplates (ca 130 µm) (Figure 14E) and other plates and rods (Figure 14D) of similar size (69–125 µm, mean 97 µm) and shape as those of the dorsal podia. Tentacle deposits include mostly perforated rods (Figure 14F) of varied form and size (54–170 µm, mean 94 µm). Distribution Type locality only. Remarks The single specimen represents a new species here referred to the monotypic genus E kkentropelma with E. brychia Pawson, 1971, as type species by original designation. The genus is characterised by a rigid U-shaped body form with a well- defined sole that does not occupy the greater curvature of the body, hence mouth and anus lying in a plane perpendicular to the usual plane of bilateral symmetry, and sole deposits only buttons and no cups (Pawson 1971). Pawson states that the sole, although not occupying the greater curvature of the body, is still ventral in position with its centre in the mid-ventral radius. He, therefore, states that the mouth and anus are displaced, lying in a plane perpendicular to the plane, which contains the mid-ventral radius and the mid-dorsal interradius. In the type species some specimens had the sole displaced to the left while others to the right of the mouth-anus plane, making the sole functionally lateral. In the present specimen the sole does not appear as distinct as in other psolids since it is deeply enclosed within a groove which is displaced to the right of the mouth-anus plane. The new species also differs from the type species in having dorsal podia arranged in a single series in each ambulacrum and very few marginal podia bordering the sole, and no cup-like deposits in the dorsal body wall. Further, the smaller plates of the body wall have smooth or slightly-knobbed margins but without projections described for the type species, and the gonad is developed as two sacciform tubules and not as tufts of tubules as in the type species. These differences clearly warrant a new species of Ekkentropelma, especially if we consider that the type species came from the Antarctic Peninsula, with depths in excess of 2000 m. With its single row of marginal podia the new species comes quite close to the preceding species but in its body form and some features of the sole deposits it is very different. According to Pawson, Ekkentropelma is closely related to Psolidium. However, besides this new species, the preceding one, and the southern African form

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erroneously referred to P. vitreum by Thandar (1999), it appears that no other psolid species has yet been described with a single row of marginal podia on each side of the sole.

Order Dactylochirotida Pawson & Fell Family Vaneyellidae Pawson & Fell Genus Psolidothuria Thandar Psolidothuria yasmeena sp. nov. Figure 15 Diagnosis Small, up to 8 mm long, U-shaped to fusiform species, with tapering posterior end. Colour in alcohol white to yellowish-white. Tentacles eight, a few ventral ones reduced. Anal teeth/papillae absent. Podia sparse, apparently non-retractile, emitting from between scales, arranged in rows in ambulacra, better developed ventrally with 5–7 podia per ventral ambulacrum. Calcareous ring simple, delicate, ribbon-like. Spicules of body wall comprise simple, multilocular, imbricating, often faintly knobbed thecal plates, up to about 450 µm long. Plates of oral and anal siphons usually without knobs; more holes in dorsal (14–27) than in ventral plates (10–24). Type SAM A-279134, paratypes SAM-A27915. Type locality Between Port St. Johns and East London, 32° 28.6’ S, 28° 58.8’ E, R.V. ‘Meiring Naude’, St. SM 226, 24.vi.1979, 710–775 m. Material examined Holotype and 24 paratypes. Etymology This species is named after my younger daughter Yasmeen, whose continuous interest in my work, even after my retirement, has been enough to keep me active for so long. Description Holotype (male) U-shaped to fusiform (Figure 15G), anterior end short, posterior end elongated, tapering. Length along ventral surface 8 mm, along dorsal surface 5.5 mm. Tentacles 8 - five large, ventral three small, retracted - all tentacles finger-shaped but finely branched (Figure 15H). Anal papillae/teeth absent. Colour white, yellowish-white in 44


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regions where viscera are visible through translucent body wall. Body wall covered by thecal plates from between which emerge relatively few podia, arranged in rows in ambulacra, better developed ventrally. Left and right ventral ambulacra with about five podia each, odd ambulacrum with 6–7 podia (mid-ventral ones paired), left and right dorsal ambulacra with 6–7 reduced podia each. Podia apparently non-retractile, whitish, with reduced suckers, decreasing in size anteriorly and moreso posteriorly. Calcareous ring ribbon-like (Figure 15I), delicate, poorly calcified; radial plates without posterior processes but with a posterior concavity and a triangular anterior projection; interradial plates slightly narrower than radial plates, more or less of same shape, but anterior projections slightly shorter than those of radial plates. Internal anatomy as in Figure 15G. Polian vesicle single, sac-like. Stone canal free, extremely short, terminating in an oval, poorly calcified madreporic body. Stomach distinct, intestine with a single loop. Respiratory trees poorly developed, without branches, each terminating in a slightly curved distal end. Cloaca elongate. Longitudinal muscles unpaired, retractors well developed, originating from anterior third of longitudinal muscles. Gonad as two tufts of tubules, four apparently mature tubules on right and three on left of mesentery. Spicules of body wall comprise multilocular, imbricating, simple thecal plates; dorsal plates (Figure 15A) 305–457µm (mean 376 µm); ventral plates 248–405 µm (mean 323 µm), of one layer of calcareous material. Plates of oral and anal siphons usually without knobs, while superficial plates of the body wall usually with knobs, up to 12 knobs per dorsal plate and up to six per ventral plate, only occurring at free margin of plate; margin of plate irregular; holes large (14–27 in dorsal and 10–24 in ventral plates). Podial deposits include smaller plates with fewer holes and other much smaller deposits in the form of perforated rods and plates (ventrally 86–224 µm, mean 144 µm) (Figure 15C); endplates reduced (ca. 150 µm) with no regular arrangement of large and small holes (Figure 15B). Tentacles supported by perforated plates (38–110 µm, mean 61 µm) (Figure 15E) and rods (Figure 15D) (105–176 µm, mean 147 µm). Introvert with similar deposits (Figure 15F). Paratypes range in size from 5–8 mm, all with retracted tentacles; body form, colouration and podial distribution (3–8 podia per row dorsally and 4–10 per row ventrally) as in holotype. Dissected paratype (male) also with eight tentacles (five large and three small). Internal anatomy as in the holotype; gonad with two pairs of unbranched tubules, each pair comprising three mature and one immature tubule. Spicules as in holotype. Distribution Type locality only. Remarks This species is clearly referable to the family Vaneyllidae as it corresponds well with the diagnosis of the family as amended by Thandar (1998). It is here assigned to the genus Psolidothuria, which was erected to accommodate only P. octodactyla Thandar, from HOLOTHUROIDEA


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considerable depths (2524–3255 m.) off the Cape Peninsula. In its body form, number and shape of tentacles, the structure of the calcareous ring and the form of the body wall, podial and tentacle deposits it comes very close to the latter species. However, it differs from P. octodactyla in its maximum size (8 mm cf. 28 mm), the distribution and nature of the podia, the absence of anal papillae, the form of the respiratory trees, the larger number of holes (up to 27 cf. 17) to the thecal plates, the smaller size of the holes (up to 50 µm cf. up to 100 µm) and the larger number of knobs on such plates. Further, the new species occurs in much shallower waters of the east coast.

FIGURE 15. Psolidothuria yasmeena sp. nov. Holotype. SAM-A279134. A. scales from midlateral body wall; B. end-plate from ventro-lateral podium; C. plates and rod from ventro-lateral podia; D. rods from tentacle stalk; E. plates from tentacle tip; F. plates from introvert; G. internal anatomy; H. largest tentacle; I. calcareous ring. (CR—calcareous ring; pv—Polian vesicle). A scale a; B–F scale b; H–I scale c.

Acknowledgements I wish to thank Ms Liz Hoensen of the South African Museum and Drs Rick Kilburn and Dai Herbert of the Natal Museum for entrusting their valuable collections to me and Dr Nadia Ameziane Cominardi of MNHNP and Mr Pieter Stiewe of the Hamburg Museum for the loan of type materials. Gratitude is also owed to the ZMUA, for working space and access to Sluiter’ s (1901) types. I am also greatly indebted to Drs Francis Rowe and Mark O’ Loughlin who made invaluable criticisms and amendments to the manuscript. Financial 46


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support from the South African National Research Foundation (NRF) and the former University of Durban-Westville is gratefully acknowledged.

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Théel, H. (1886a) Holothuroidea. Part 2. Reports of the scientific Results of the Voyage of the H.M.S. ‘Challenger’, (Zoology), 39, 1–290. Théel, H. (1886b) Report on the Results of Dredging, under the Supervision of Alexander Agassiz, in the gulf of Mexico (1877–78), in the Caribbean Sea (1879–80), and along the Eastern Coast of the United States during the Summer of 1880, by the U.S. Coast Survey Steamer ‘Blake’. Bulletin of the Museum of Comparative Zoology, 13, 1–21. Vaney, C. (1905) Holothuries recueillis par M.Ch. Gravier sur la côte française des Somalis. Bulletin du Museum national d’Histoire naturelle de Paris, 11, 186–191. Vaney, C. (1908a) Les Holothuries recueillies par l’ Expédition Antarctiques Ecossaise. Zoologische Anzeiger, 33, 290–299. Vaney, C. (1908b) Les Holothuries de l’ Expedition antarctique nationale Ecossaise. Transactions of the Royal Society of Edinburgh, 46, 405–441.

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