New or notable records of brittle stars (Echinodermata: Ophiuroidea) from South Africa

 

 

Jennifer M. Olbers^I; Yves Samyn^II; Charles L. Griffiths^III

^IEzemvelo KZN Wildlife, Private Bag X3, Congella, Durban, 4052 South Africa E-mail: jennifer.olbers@kznwildlife.com
^IIRoyal Belgian Institute of Natural Sciences, Rue Vautier 29, Brussels, Belgium E-mail: yves.samyn@naturalsciences.be
^IIIDepartment of Biological Sciences, University of Cape Town, Rondebosch, 7700 South Africa E-mail: charles.griffiths@uct.ac.za

 

 

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ABSTRACT

Ophiuroid research in South Africa has not kept pace with global taxonomie research with the last major taxonomie review of the group being published in 1976. This paper documents all new records of Ophiuroidea from South Africa since (and including) 1977. These records originate from specimens housed in five zoological collections, from photographic records and from reports published in the non-taxonomic literature. A short review of the history of ophiuroid taxonomy in South Africa is also given and for each new record, key references, distribution, ecology, additional notes and, where possible, photographs, are presented. This has resulted in an additional 24 species being recorded within the mainland Exclusive Economic Zone of South Africa, elevating the total known number of ophiuroid species reported in the region to 137.

Keywords: taxonomy, nomenclature, new record.

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INTRODUCTION

The coastline of mainland South Africa is c. 3650 km in length, with an Exclusive Economic Zone (EEZ) of 1 068 659 km² (Griffiths et al. 2010). The greatest depth recorded within the mainland EEZ is c. 5700 m. The continental shelf is narrow along the east coast, wider on the west coast and reaches a maximum width ofc. 260 km offthe Agulhas Bank in the south. Marine systematic research in South Africa began during the mid-1750s to late 1800s (H.L. Clark 1923) and initially took the form ofa number oflarge expeditions, such as the Challenger and the Gazelle expeditions that collected within South African waters. The majority ofthese specimens were taken to European museums, where they were described and often well illustrated (Linder & Griffiths 1999; Griffiths et al. 2010). In the 20th century, further efforts to document the southern African echinoderm fauna were undertaken by other large expeditions, such as the Valdivia and some Antarctic expeditions (H.L. Clark 1923). Today, South Africa is well-known for its extraordinary biodiversity and is considered to be the third most biologically diverse country in the world (World Conservation Monitoring Centre 1992). Griffiths et al. (2010) reported that 12 914 marine species were known from South Africa, with 33% of these endemic to the region. Complementing this, over 291 000 records, including marine invertebrates, molluscs, fish and algae are curated at various coastal museums within South Africa alone (Griffiths et al. 2010).

The current state of knowledge for ophiuroids in South Africa is a result of contributions from a number of authors since the late 1700s (Fig. 1). The first record of an ophiuroid from South Africa was that by Retzius (1783) who reported Asterias euryale ( = Astrocladus euryale) from the Cape of Good Hope, followed by Müller & Troschel (1842) and Ljungman (1867) who added two and five additional species to the South African fauna, respectively. As a result of the Challenger expedition, which sampled seven stations within South African waters (excluding the Prince Edward and Marion Islands), Lyman (1878; 1882), reported four and 17 new ophiuroids, respectively. Later, Bell (1888) described one additional new ophiuroid from the same collection. Bell (1905) further added five new ophiuroids in one of his papers on the Echinoderma of South Africa. Döderlein (1910) wrote the first consolidated account of South African echinoderms, reporting 84 species ofechinoderm, including 29 ophiuroids.

 

 

More than a decade later, H.L Clark (1923) reported a total of 57 ophiuroid species as being known for South Africa, including six new species which were largely derived from the Pieter Faure expedition. Mortensen (1925) added two more species to the fauna from a collection sent to him from the Durban Museum (Astroschema capense (= Astromorpha capensis) and Ophiactis savignyi (Müller & Troschel, 1842)), the former being new to science. Hertz (1927a,b) added four new species to the South African fauna, but two of these were soon synonymized by Mortensen (1933a) in his significant contribution to the Ophiuroidea and Asteroidea of South Africa. Mortensen (1933a) recorded 36 new ophiuroid species from material collected mostly off the Pickle and the John. C. Meikle, bringing the total number of ophiuroids known for South Africa to 82 species. Mortensen (1936) reported on collections from the Discoveryexpedition and added two new species from South Africa. A.M. Clark (1952) described an additional three species collected during the University of Cape Town (UCT) Ecological Surveys and from the Africana. Later, A.M. Clark (1974) summarized records from 22 years ofcollections undertaken during the UCT Ecological Surveys and the Anton Bruun expedition that had accumulated since A.M. Clark's (1952) report by describing three new species and adding four new records to South Africa. Clark & Courtman-Stock (1976), now the standard current monograph on the southern African echinoderm fauna (excluding Holothuroidea), reported on 115 species of Ophiuroidea. However, only 101 of these species were found within the political borders of South Africa. Shortly afterwards, A.M. Clark (1977) reported on a number of deep-water species collected by the Meiring Naude, which added ten new ophiuroid species to the South African fauna. Madsen (1977) reported Ophiernus quadrispinus Koehler 1907 from off Cape point, a new record for South Africa. Following this, no taxonomic work was undertaken for 35 years until recently when Olbers & Samyn (2012) reported Ophiocomabrevipes Peters, 1851, O. dentata Müller & Troschel, 1842, O. doederleini de Loriol, 1899 and O. pusilla (Brock, 1888) as new records for South Africa. Later that year, Milne (2012) reported Ophiactis picteti (de Loriol, 1893), Macrophiothrix demessa (Lyman, 1862) and M. propinqua (Lyman, 1862) occurring at Sodwana Bay. These two reports raised the total number of ophiuroids reported in the published literature for South Africa to 119.

In addition to these published reports, additional data and unidentified material have continued to accumulate. Bolton et al. (2001) reported that between 1999 and 2001, 51 species of echinoderm were added to the KwaZulu-Natal (KZN) checklist. The voucher specimens supporting these records have been deposited in the collections of the Royal Museum for Central Africa in Tervuren, Belgium (RMCA) and although Samyn & Thandar (2003) used these records to conduct a preliminary biogeographical analysis, no species list was ever formally published. In addition, ophiuroid samples have continued to accumulate in the Iziko South African Museum (SAM) but have remained unidentified and mostly uncatalogued for a period of ~35 years.

The aim of the present contribution is to gather all data regarding additions to the ophiuroid fauna of South Africa subsequent to the previous monograph by Clark & Courtman-Stock (1976), and to list and document these in a single publication. These new records include both those published in papers subsequent to 1976 (as listed above), the identified but unpublished records of RMCA, photographic records and those newly identified by the authors from collections in the SAM.

 

MATERIALS AND METHODS

The ophiuroid species reported on in this account originate from a number of sources. The majority of the records are from unidentified specimens deposited in the SAM collection, while others originate from the Ezemvelo KZN Wildlife (EKZNW) Echinoderm collection, housed in Durban. Also reported on are the South African specimens housed in the RMCA in Tervuren, Belgium, the Smithsonian National Museum of Natural History, Washington DC (USNM), and the Australian Museum (AM) in Sydney. Additional records were obtained from photographic evidence sourced from the South African National Biodiversity Institute (SANBI) iSpot programme, the Animal Demography Unit EchinoMAP programme and published literature, as cited in the taxonomic account below.

Specimens are preserved either in 70% ethanol or dry. The RMCA collections are preserved dry, while those held by SAM and EKZNW are mostly preserved in ethanol. The specimens were collected using a variety of methods, including trawling, dredging, grab sampling, SCUBA diving and by hand on the shore. The depths at which the specimens were collected ranged from 0-2948 m.

Specimens were primarily identified using the keys and descriptions of Lyman (1878), Clark, A.M. (1952), Clark & Rowe (1971), Clark, A.M. (1974), Clark & Courtman-Stock (1976), Cherbonnier & Guille (1978), Mortensen (1925) and Mortensen (1933a). Additional useful literature is also cited for each species.

Taxa are arranged according to their currently known classification as given by Stöhr et al. (2014) in the World Ophiuroid Database linked to the World Register ofMarine Species (WoRMS). Species are presented under the binomen as considered valid by Stöhr et al. (2014). A comprehensive diagnosis is given and where possible, type material, distribution, ecology and additional notes are also reported. In addition, where specimens were available, these were photographed.

 

RESULTS

Species accumulation over time

The increasing number of ophiuroid species reported for the region since 1783 is presented in Fig. 1. For more than 60 years, only a single species was known for the region. The number of species slowly increased until 1923, when there were a number of reports by largely three European researchers whose work focused on South African echino-derm collections over approximately the following half century, culminating in the monograph of Clark & Courtman-Stock (1976). From that time until the start of the present study, no ophiuroid taxonomy has been undertaken in South Africa.

This report documents additions to the ophiuroid fauna since Clark & Courtman-Stock (1976), including A.M. Clark (1977) and Madsen (1977), raising the total by 24 species across 11 families and 19 genera. Therefore, the current total number of known species from South Africa is 137.

Of the 28 species reported here, 24 are new to South Africa while four are for noting (Amphilimna cribriformis A.M. Clark, 1974; Ophionephthys lowelli A.M. Clark, 1974; Ophiernus quadrispinus Koehler, 1908; Amphioplus (Lymanella) depressus (Ljungman, 1867)). Six species are represented by only a single record, while 11 have been reported five or more times. Twelve species were recorded in only shallow water (<30 m), four in deep water (31-500 m), five in very deep water (>501 m) while seven species traversed both shallow and deep depths. Most of the additions are Indo-Pacific species whose ranges have extended southwards into subtropical eastern regions of South Africa and were derived from the collection deposited in the RMCA and unidentified material in the SAM. Table 1 lists all species with their global and local distribution and depth ranges within South Africa.

 

TAXONOMIC ACCOUNT

Class OPHIUROIDEA Gray, 1840

Family ASTEROSCHEMATIDAE Verrill, 1899

Asteroschema salix Lyman, 1879

Plate 1A, B

Asteroschema salix Lyman, 1879: 66-67, pl. 17, figs 466-469; Baker, 1980: 23-24; McKnight, 2000: 21, 22. pl. 6, fig. 7.

Material/Records

SAM A28143, -31.0000°, 30.4500°, off Glenmore, depth 900 m, 12 May 1977, Meiring Naude, station number SM134, beam trawl, identified by M. Okanishi and J.M. Olbers.

Diagnosis

(See Baker 1980)

D.D. up to 10 mm. Disc round, indented interradially, lateral interradial surface almost vertical, body surface covered with epidermal plates with rounded granules. Radial shields elongated, narrow, raised, covered in plates, converging and almost meeting at centre of disc. Oral shields absent, adoral shields indistinct. Jaws covered by minute granules. Teeth seven, broad, triangular, lowermost appearing to be paired. Genital slits short, wide. Arms five, slender, coiling, narrow, higher than wide. No arm spines from first pair of tentacle pores to segment 15, then two arm spines, one slightly smaller. Arm spines short, innermost longest and cigar-shaped, finely serrated. Colour in life: pink (McKnight 2000).

Global distribution New Zealand (McKnight 2000), South Africa.

Ecology

Depth range: 341-1800 m (Lyman 1879; Baker 1980;McKnight 2000).

Habitat: No details recorded.

Remarks

Single specimen recorded off KZN south coast, previously only known from waters off New Zealand and is a noteworthy extension into the Indian Ocean. According to Baker (1980), type locality is west of Raoul Island, Kermadecs, depth 1152 m. Holotype housed in the Natural History Museum, London (BMNH 82.12.23.271B).

Family GORGONOCEPHALIDAE Ljungman, 1867

Astroboa nuda (Lyman, 1874)

Astrophyton nudum Lyman, 1874: 251-252, pl. 6, fig. 4-5.

Astroboa nuda Döderlein, 1911: 86-88; Mortensen, 1940: 67; Tsurnamal & Marder, 1966: 9-17, fig. 1-4; Clark & Courtman-Stock, 1976: 100, 108, 130-131; Cherbonnier& Guille, 1978: 17-18, pl. 1, figs 3-4; Baker, 1980: 60, fig. 22; Guille & Vadon, 1985: 62.

Astrophyton elegans Koehler, 1905: 123-125, pl. 13, fig. 2, pl. 18, fig. 1.

Astroboa nigra Döderlein, 1911: 83-86, pl. 9. figs 9, 9a.

Astroboa nuda var. elegans: Döderlein, 1927: 45.

Astroboa nuda var. nigra: Döderlein, 1927: 44; Balinsky, 1957: 2-3.

Material/Records Data Record Sink et al., 2006-6, -27.4817°, 32.7117°, Sodwana Bay, Wright Canyon, depth 70-120 m, ROV, November 2000, Determined by Gordon Paterson. Data Record Sink et al., 2006-55, -27.5263°, 32.7198°, Sodwana Bay, Jesser Canyon, depth 100-110 m, ROV, November 2000, determined by Gordon Paterson. Data Record SANBI, iSpot, -27.5355°, 32.6799°, Sodwana Bay Canyons, depth 69 m, 1 March 2014, determined by K. Sink.

Diagnosis

(See Clark & Courtman-Stock 1976; Baker 1980) D.D. up to 92 mm. Disc depressed interradially and centrally, interradial and radial areas naked towards centre of the disc but with an increasing presence of tiny tubercles towards disc margin. Radial shields, narrow, paved densely with low granules giving smooth appearance, raised at disc margin, slightly broader on distal side terminating in oval slightly concave plate, converging to centre of disc. Oral papillae short, narrow, no continuous fringe in distal notches. Teeth three to five, thicker than oral papillae but elongated. Genital slits small, wide. Genital papillae present on inner edge. Ventral interradial areas densely covered with tiny tubercles. Madreporite, one. Arms higher than wide basally, branching, first fork close to disc base, four to eight segments between forks with up to 28 forks along arm. Arms covered in small, smooth, polygonal plates, girdle bands present on arms from after 2nd fork but continuous before 3rd branch, girdle hooklets with secondary tooth. Arm spines absent before 15th fork, spines 3-4 with distal spines becoming hooklets with two hooks. Colour in life: black, white or yellow.

Global distribution

Indo-Pacific, Australia, Western Indian Ocean, Red Sea, Persian Gulf, East Indies, China and south Japan, Philippines, Madagascar, Mozambique (Kalk 1958; Richmond 2002; Rowe & Gates 1995; Clark & Rowe 1971; Tsurnamal & Marder 1966; Balinsky 1957; Clark & Courtman-Stock 1976; Macnae & Kalk 1958; Cherbonnier & Guille 1978), South Africa (Sink et al. 2006).

Ecology

Depth range: 0.5-120 m (Tsurnamal & Marder 1966; Sink et al. 2006).

Habitat: Found on coral reefs, both within deep crevices and on open reef.

Remarks

Previously known in Mozambique and hence not surprisingly recorded in South Africa. According to Rowe & Gates (1995), type locality is the Philippines, with the holotype being held at the Museum of Comparative Zoology (MCZ 2911).

Astroglymma cf. sculptum (Döderlein, 1896)

Plate 1C, D

Astrophyton sculptum Döderlein, 1896:299, pl. 18, figs 29a, b;

Baker, 1980: 66, 74, figs 19, 28, 31.

Astroglymna sculptum: (misspelling) Rowe & Gates, 1995: 365.

Gorgonocephalusrobillardide Loriol, 1899: 31-34, pl. 3, fig. 3. Astrodactylus robillardi: Döderlein, 1911: 96-98. Astroglymma sculptum Döderlein, 1927: 47-50, pl. 1, figs 3, 4; pl. 5, fig. 13; Koehler, 1930: 15, pl. 2, figs 10-12; Guille & Vadon, 1985: 62.

Astroglymma robillardi: Mortensen, 1933b: 34, pl. 3, figs 1,2; pl. 4, fig. 1.

Material/Records

SAM A74047, -27.7668°, 32.6500°, NE Gypsea Hill, depth 84-90 m, 9 June 1990, Meiring Naude, station number ZK22, Natal Museum Dredging Programme. USNM 1072476,-29.4500°, 31.5100°, east of Durban, Anton Bruun, station number 394B, depth 68-70 m, 25 September 1964, determined by A.N Baker.

Diagnosis (See Baker 1980)

D.D. up to 50 mm. Disc deeply excavated interradially. Radial shields long, slender, widelyseparated distallyalmost touching proximally, almost reaching centre of disc. Disc and radial shields covered in minute conical tubercles, ventral interradial area may bear long spinelets. Five madreporites present in angle of ventral interradial area. Oral shields smooth, adoral shields not distinct, deep pits bordering jaws. Oral papillae unequal, small, mostly spiniform. Teeth small, spatulate. Genital slits short, D-shaped. Genital papillae blunt-tipped on outer edge. Arms branching, first fork just beyond disc, forking at least 20 times along arm. Dorsal arms covered in low polygonal plates. Girdle bands narrow, present from arm bases, girdle hooklets with secondary tooth. Arm spines present from sixth fork as two stumps, becoming three with one or two terminal points, distally becoming hooklets with terminal point and smaller secondary tooth. Ventral arms covered with smaller flat polygonal plates, the ventral arms have ladder-like pits on first 2-3 forks.

Global distribution

Indo-west Pacific, Australia, India, Mauritius, Malaysian Archipelago, China Sea (Baker 1980; Imaoka et al. 1991; Rowe & Gates 1995), South Africa.

Ecology

Depth range: 7-300 m.

Habitat: On continental shelf and slope (Rowe & Gates 1995).

Remarks

Baker (1980) synonymized Astroglymma robillardi with A. sculptum, agreeing with Mortensen (1933b) that there were no reliable differences between sculptum and robillardi and that a specimen being found in the Indian Ocean gave no reason to have separate species. He did note that Mortensen's A. var. spinosum may have merit but without comparative material it should remain a variety.

In this study, the specimen of Astroglymma described from off Durban had some noticeable differences; however, not those that Mortensen used to differentiate A. robillardi from A. var. spinosum. Mortensen (1933b) used the following to differentiate sculptum from spinosum:i)spinosum disc has thick stumps ending in some hyaline thorns with similar stumps on radial shields except they are close set in comparison to sculptum. Ventrally, few stumps present.

The specimen at hand (SAM A74047) had fine tubercles and were close together giving a smooth appearance, while ventrally, the disc covering was similar to the dorsal disc; ii) in A. var. spinosum there were no distinct grooves on the oral frame and no ladder pits on proximal part of arm. In SAM A74047 these pits were present; iii) the arm spines in A. var. spinosum begin before the 1st fork and the primary hooks are larger and different in shape (A. var. spinosum hooks are hunched over) to sculptum. In SAM A74047, there are very few hooks on the arm spines and they begin from the 4th fork.

In addition to not being similar to A. var. spinosum, SAM A74047 also has a number of differences from A. sculptum: i) arm spines start from the 4th fork and not the 6th as in sculptum; ii) distal arm spines have hooks but very few with secondary hooks; iii) girdle hooklets only have a terminal hook, therefore no secondary hooks are present on girdle belts as described by de Loriol (1899), Baker (1980) and Mortensen (1933b); and iv) there are 8-10 forks on SAM A74047 which is much fewer than those for sculptum (up to 20 forks). Therefore, it is noted that SAM A74047, does not strictly conform to sculptum, robillardi or var. spinosum.

The second record for South Africa is held at the Smithsonian Institution, National Museum of Natural History in Washington, D.C. (USNM), (USNM 1072476). The type locality of this species is Amboina, Indonesia.

Family OPHIOMYXIDAE Ljungman, 1867

Ophiomyxaaustralis Lütken, 1869

Plate 1E, F

Ophiomyxa australis Lütken, 1869: 45, 98, 99; Lyman, 1882: 246; Koehler, 1907: 341, Benham, 1909: 101; H.L. Clark, 1915: 168, pl. 1, figs 1-2; H.L. Clark, 1916: 77; Matsumoto, 1917: 19-21, fig. 3, pl. 1, figs 4-7; H.L. Clark, 1938: 201, pl. 13, figs 1-21; H.L. Clark, 1939: 36-37; H.L. Clark, 1946: 170-171; Madsen, 1967: 141; Clark & Rowe, 1971: 78, 92-93, pl. 13, figs 3, 4; Devaney, 1974: 1 15-1 16; Cherbonnier & Guille, 1978: 18-19, pl. 3, figs 1, 2; Sloan etal., 1979: 99, figs 5, 6; Irimura, 1982: 2-4, fig. 1; Guille & Vadon, 1985: 62; Sastry, 1991: 375-376; Rowe & Gates, 1995: 406; Mbongwa, 2013: 15. Ophiomyxa brevispina von Martens, 1870: 249-50; de Loriol, 1893b: 425-426; Döderlein, 1896: 298, pl. 17, fig. 27; Koehler, 1905: 119; H.L. Clark, 1915: 170, pl. 1, figs 1, 2; Koehler, 1930: 48.

Ophiomyxa robillardi de Loriol, 1893a: 53-54, pl. 25, fig. 5. Ophiomyxa brevispina var. irregularis Koehler, 1898:111-112. Ophiomyxa irregularis Koehler, 1905: 119-120, pl. 12, fig. 1; Koehler, 1922a: 17-20, pl. 2, fig. 18, pl. 5, figs 1, 2, pl. 6, fig. 4, pl. 92, fig. 2; Koehler,1930: 48.

Material/Records

RMCAMT2305, -27.5227°, 32.6919°, Sodwana Bay, 2-mile Reef, no depth, 10 February 2001, collected by Y. Samyn, determined by Y. Samyn and M. Garcia. RMCA MT2274, -27.5227°, 32.6919°, Sodwana Bay, 2-mile Reef, depth 13 m, 9 August 1999, collected by Y. Samyn, determined by Y. Samyn and M. Garcia. RMCAMT2361, -27.5227°, 32.6919°, Sodwana Bay, depth 30 m, 14 August 1999, collected by Y. Samyn, determined by Y. Samyn and M. Garcia. SAM A28127, -27.5166°, 32.6833°, Sodwana Bay, depth 15 m, 24 July 1976, SCUBA, J.L.B. Smith Institute, determined by J.M. Olbers. SAM A28133, -27.5166°, 32.6833°, Sodwana Bay, depth 15 m, 25 July 1976, SCUBA, J.L.B.

Smith Institute, determined by J.M. Olbers. SAM A28140 and SAM A28142, -27.5166°, 32.6833°, Sodwana Bay, depth 14 m, 1 August 1976, SCUBA, J.L.B. Smith Institute, determined by J.M. Olbers. EKZNW SB_4_JMO_2010, -27.8667°, 32.6000°, Leadsman Shoal, depth 11 m, SCUBA, 13 October 2010, determined by J.M. Olbers. SAM A74042, -27.1000°, 32.8841°, off Dog Point, depth 74 m, 7 June 1990, Dredge, Meiring Naude, station number ZC10, Natal Museum Dredging Programme, determined by J.M. Olbers. SAM A74043, -27.1844°, 32.8350°, off Rocktail Bay, depth 75 m, 4 June 1987, Dredge, Meiring Naude, station number ZD2, Natal Museum Dredging Programme, determined by J.M. Olbers.

Diagnosis (See Cherbonnier & Guille 1978)

D.D. up to 23 mm. Disc pentagonal, covered with thick opaque smooth skin. Radial shields short, narrow, separated by width of arm base; disc margin scales overlap. Genital slits bordered by plates similar to ones on disc margin, long, narrow. Oral shields oval, triangular, covered by thick skin, longer than wide, abutting genital slit. Oral papillae three, broad, serrated, flattened and transparent on edges. Teeth similar. Arms five, covered in thick naked skin. Arm spines up to seven, one on segment one, then two and four on first free arm segments. Arm spines slender, serrated and rugose at tip, some becoming curved or slightly hooked. Dorsal arm plates irregular, fragmented, becoming less fragmented distally. Ventral arm plates distinctly broader than long, deep notch on distal side, not contiguous distally. Tentacle scales absent. Colour in life: blood-red dorsally and ven-trally, arms lightly banded with yellow.

Global distribution

East Africa and Islands, Madagascar, Kenya, Mascarene Basin, Red Sea, Seychelles, Somalia, Tanzania, Western Indian Ocean, New Zealand (Stöhr et al. 2014), Indo-west Pacific (Rowe & Gates 1995), South Africa (Mbongwa 2013).

Ecology

Depth range: 11-75 m.

Habitat: In sand, grey ooze, coral, crannies in coral, stones and gravel, mud, sandstone rubble, gorgonians (Lyman 1882; H.L. Clark 1938).

Remarks

According to Rowe & Gates (1995), the type localityis Bass Strait (as '...inter Australian et Tasmaniam') with the syntypes being held at the Natural History Museum of Denmark (ZMUC 474) (Tom Schi0tte, pers. comm.).

Family OPHIURIDAE Lyman, 1865

Subfamily Ophioleucinae Matsumoto, 1915

Ophiernus quadrispinus Koehler, 1908

Plate 2A, B

Ophiernus quadrispinus Koehler, 1908a: 533, 601-602; pl. 10, figs 102, 103; Koehler, 1908b: 142,146; Madsen, 1977: 120-121, fig. 7; Billett et al., 2013: 20-25.

Material/Records

SAMA22018, offSaldanhaBay,-33.8116°, 16.5000°, depth 2730 m, 27 August 1959, trawl, Africana II, station number A193, determined by A.M. Clark. SAM A22015, -34.6166°, 17.0500°, off Cape Town, depth 2875-2948 m, 8 December 1959, trawl, Africana II, station number A315, determined by A.M. Clark. Data Record Madsen, 1977, -34.4044°, 17.7437°, off Cape Point, depth 1700-1900 m, Fisheries Survey.

Diagnosis

(See Madsen 1977)

D.D. up to 7 mm. Disc pentagonal, covered in scales, dorsally and ventrally, scales slightly bigger abutting radial shields and genital slits. Sparse granules on disc margin extending onto margins of radial shields. Radial Shields, large, oval, longer than wide, separated by disc scales. Oral shields arrow-shaped, naked. Adoral shields not distinct, extending up to first ventral arm plate, may or may not be contiguous proximally. Jaws long. Oral papillae five to six, including two smaller papillae in series with scales around second oral pore. Teeth three to four, tapering to blunt point. Genital slits as long as interradial area, genital papillae absent. Ventral arm plates bell-shaped, first plate sunken, contiguous proximally becoming reduced and separated distally. Dorsal arm plates wider than long, distal edge straight proximally, becoming convex distally. Lateral plates increasing in size distally, hosting arm spines. Arm spines four, delicate, cylindrical, pointed, shorter than segment length, decreasing distally. Arm spines placed on mid-plate proximally, moving dorsally distally, upper bristle-like arm spine absent. Arms moderately long (all specimens broken), dorsal arm with slight keel. Tentacle scales two, sometimes one, varying in shape from pointed to round, unequal in size.

Global distribution

South Georgia and Crozet Island, Southern Ocean, Southern Atlantic, near the South Orkneys (Madsen 1977; Billett et al. 2013), South Africa (Madsen 1977).

Ecology

Depth range: 1700-3250 m (Madsen 1977).

Habitat: No information available.

Remarks

Although these specimens were trawled in 1959 and identified by A.M. Clark (date of determination unknown), they were not included in the Clark & Courtman-Stock (1976) monograph for unknown reasons. Later, Madsen (1977) described it as a new record for South Africa.

Granules on disc margin are similar to O. vallincola Lyman, 1878 but fewer in number. Similarly, granules extend onto margins of radial shields, but again fewer in number. Dorsal arm plates wider than long but slightly wider than in O. vallincola. Type locality is Southern Ocean (Koehler 1908a). In addition, the two similar species, O. seminudus and O. quadrispinus both lack the bristle-like supplementary spines on the lateral plate whereas these are present on O. vallincola.

Subfamily Ophiurinae Lyman, 1865

Amphiophiura sculptilis (Lyman, 1878)

Ophioglypha sculptilis Lyman, 1878: 84-85, pl. 4, figs 115, 116; Lyman, 1882: 37.

Ophioglypha variabilis Lyman, 1878: 85-86, pl. 4, figs 113, 114; Lyman, 1882: 37.

Ophiura sculptilis: Ludwig, 1901: 925; H.L. Clark, 1911: 77.

Ophioglypha remota Koehler, 1904a: 54, pl. 9, figs 1-3.

Amphiophiura sculptilis Koehler, 1914: 24; Koehler, 1922a: 364; H.L. Clark, 1915: 313; Matsumoto, 1915: 77; Hertz, 1927b: 74; H.L. Clark, 1939: 108; Madsen, 1951: 114; Litvinova, 1971: 299, pl. 3, figs2, 4, 5; Vadon &Guille, 1984: 588, 592-593, pl. 5,1-4; Guille&Vadon,1986:169; Manso, 2010: 196.

Material/Records

Data Record Guille & Vadon, 1986, -29.8133°, 34.5450° off Durban, depth 2608 m, 21 August 1979, dredge, Safari I (Marion-Dufresne), station number DS1, determined by A. Guille and A. Vadon.

Diagnosis

(See Lyman 1878; Vadon & Guille 1984)

D.D. up to 15 mm. Dorsal disc thick, scales thin and flat, large round central plate, five distinct plates separated by small irregular scales. Radial shields distinct, D-shaped, contiguous distally, tapering proximally with wedge of scales between them, large scale present on dorsal interradial area. Ventral interradial areas scaled, but dominated bylarge oral shield. Oral shield pentagonal, distal edge rounded, slightly longer than wide, covering most of the ventral disc surface. Adoral shields relatively broad, contiguous. Oral papillae five, broad, closely set, apical papillae blunt. Genital slits moderately long, genital papillae present, squarish becoming spiniform, forming arm combs dorsally. Dorsal arm plates fan-shaped, rounded distal edge, contiguous. Lateral arm plates broad, meeting ventrally. Ventral arm plates squat, bell-shaped, constricted by large tentacle pore, distal edge longer than proximal edge, wider than long, distal edge straight becoming rounded, not contiguous. Arm spines up to six, blunt. Tentacle pores large, tentacle scales up to five within disc, two on remaining arm.

Global distribution

Antarctic Ocean, Northern Atlantic, Southern Atlantic, Zanzibar, Oman, Réunion, Indonesia, Japan, South America, Brazil, Bay of Bengal (Koehler 1914; Koehler 1922a; H.L. Clark 1939; Vadon & Guille 1984), South Africa (Guille & Vadon 1986).

Ecology

Depth range: 300-4320 m (Vadon & Guille 1984).

Habitat: Grey sand, globigerina ooze, grey mud (Koehler 1914; Koehler 1922a).

Remarks

Single record from South Africa, collected during French expedition Safari I on Marion-Dufresne, reported by Guille & Vadon (1986). Type locality is off Japan.

Family AMPHIURIDAE Ljungman, 1867

Amphilimna cribriformis A.M. Clark, 1974

Plate 2C, D

Amphilimna cribriformis A.M. Clark, 1974: 442-444, figs 1a-d; Thomas, 1975: 131, 132, 137; Clark & Courtman-Stock, 1976: 122, 165, 166, figs 182, 183; Liao, 1989: 342.

Material/Records

SAM A22784 (disintegrated holotype), NE of Durban, depth 118 m, -29.5670°, 31.6500°, dredge, sandy green-brown mud, 9 September 1964, station number NAD40V; SAM A22787 (paratype), NE of Durban, depth 86 m, -29.4833°, 31.7500°, dredge, mud, 9 September 1964, station number NAD52E; SAM A22785 (paratype), one specimen in poor condition, off Zimbali, depth 115 m, -29.5670°, 31.6500°, dredge, mud, no station data or collection date; SAM A22788 (paratype), NE of Tongaat, depth 150 m, -29.5833°, 31.6333°, dredge, sandy green brown mud, 9 September 1964, station number NAD35W; SAM A22789 (paratype), off Zimbali, depth 118 m, -29.5677°, 31.6519°, dredge, sandy green brown mud, 9 September 1964, station number NAD43G; SAM A22790 (paratype), off La Mercy, depth200 m,-29.6333°, 31.6019°, dredge, mud, 8 September 1964, station number NAD33B; SAM A22791 (paratype), off Sheffield Beach, depth 86 m, -29.4833°, 31.7500°, dredge, mud, 9 September 1964, station number NAD55C.

Diagnosis

(See A.M. Clark 1974; Clark & Courtman-Stock 1976)

D.D. up to 6.5 mm. Disc pentagonal, indented radially, uniformly white, both dorsally and ventrally due to preservation. Dorsal and ventral disc covered in medium-sized fine disc scales with scattered, tapering, sharp spinelets, no change in spinelet, scale density or size on disc margin. Radial shields long, narrow, spines may be absent. Genital plates large, lie at angle in which they appear to be overlapping, each plate hosting two stout spines at their dorsal end. Oral shields triangular with rounded angles, as long as wide, widest distally. Adoral shields restricted to lateral edge of oral shield, triangular with inner margin curved, not contiguous. Jaws slightly elongated, two to four asymmetrical apical oral papillae, three spinose distal papillae, two distal-most being on edge of adoral shield. Teeth single, broad with small elongated oral tentacle scale either side. Arms long and thin, first two to four dorsal arm plates short, compressed or rudimentary, narrow. First free arm plate fan-shaped with convex distal edge, as long as wide, narrowly contiguous, plates translucent, porous and brittle with underlying structure visible. First ventral arm plate appearing triangular, adjacent to adoral plates, second arm shield with straight distal edge, broader between tentacle pores. Ventral arm plates thereafter with slight convex edge, becoming concave distally, narrowing adjacent to tentacle pores, longer than broad. Arm spines six, with first seven to nine arm plates hosting flattened, webbed arm spines, forming a wing-like flange which excludes lowermost spine. Beyond disc, arm spines free, flattened, becoming round and tapering distally. Tentacle scales two on segments one to ten, outer scale small, inner scale spinose resembling an arm spine, becoming reduced and eventually completely lost, single tentacle scales after segment ten.

Global distribution

Mozambique, South Africa (A.M. Clark 1974; Clark & Courtman-Stock 1976).

Ecology

Depth range: 86-200 m.

Habitat: Sandy mud, continental shelf (A.M. Clark 1974; Thomas 1975; Clark & Courtman-Stock 1976).

Remarks

Upon examination, it was found the holotype has disintegrated, it is suggested that if a neotype were to be erected, the paratype SAM A22787, collected 13 km from original type locality, would be the most appropriate specimen because this individual, one of nine paratypes, is most probably in the best condition.

In 1899, Verrill placed Amphilimna into the family Amphiuridae, which was supported by H.L Clark (1915) and Koehler (1922a). Later in 1967, Thomas placed Amphilimna into the Ophiacanthidae, which was supported by A.M. Clark (1974), Clark & Courtman-Stock (1976) and Liao (1989). Paterson (1985) then placed it into a subfamily Ophiotominae (family Ophiacanthidae). In 2010, Martynov proposed to place Amphilimna back into the Amphiuridae as suggested by Verrill (1899), despite having an atypical dental plate. This genus has been dubbed an aberrant genus between Amphiuridae and Ophiacanthidae.

Amphioplus (Lymanella) depressus (Ljungman, 1867)

Plate 2E, F

Amphipholis depressa Ljungman, 1867: 312. Ophiophragmus affinis Duncan, 1887: 89-90, pl. 8, figs 4-6. Amphiura relicta Koehler, 1898: 69, pl. 4, figs 37, 38; Koehler, 1900: 4, pl. 16 figs. 15, 16.

Amphioplus relictus: H.L. Clark, 1915: 256; H.L. Clark, 1938: 251.

Amphioplus depressus: H.L. Clark, 1915: 254; H.L. Clark, 1946: 205; James, 1970: 142-144, fig. 1g-k.

Amphipholis hastata Ljungman, 1867: 313.

Amphioplus hastatus: H.L. Clark, 1915: 257; H.L. Clark, 1923: 331; Koehler, 1927: 6; H.L. Clark, 1939: 75-76; Day & Morgans,1956: 308;A.M.Clark,1967: 47;Vine,1986:195.

Amphioplus (Lymanella) hastatus: A.M. Clark, 1970: 51, 54-55, fig. 9p, q; Clark & Rowe, 1971: 80, 102, fig. 24a; Hughes & Gamble, 1977: 355; Cherbonnier & Guille, 1978: 81, 83-86, figs 36, 37; Sloan et al., 1979: 101; Richmond, 2002: 326.

Amphioplus (Lymanella) depressus: A.M. Clark, 1970: 54; Clark & Rowe, 1971: 102; Gibbs et al., 1976: 117-118; Baker 1979: 46.

Material/Records

DBN52H, -29.8660°, 31.0270°, Durban Bay, depth 0 m, 18 July 1950, collected by hand, UCT Ecological Survey Collection, determined by unknown. SAM A74078, -34.0817°, 23.0126°, Knysna, depth 0 m, 7 July 1960, collected by hand, UCT Ecological Survey Collection, determined by T. O'Hara. Data Record Mbongwa, 2013, -27.7320°, 32.6264°, Sodwana Bay, depth unknown, 27 February 2013, determined by J.M. Olbers andN.A. Mbongwa.

Diagnosis

(See A.M. Clark 1970; Clark & Rowe 1971)

D.D. up to 10 mm. Primary rosette may or may not be distinct. Disc scales moderate in size, overlapping, central scales may be larger than peripheral scales. Disc margin sharp, sometimes with small projections or spines. Radial shields contiguous for half their lengths, may be half disc radius or less. Oral shields narrow, diamond-shaped, longer than wide, adoral shields triangular, contiguous. Oral papillae four, arranged in a continuous row forming a straight line, third papilla slightly enlarged. Arm length approximately 6-7 times disc diameter. Dorsal arm plates oval, wider than long, distal margin convex, contiguous. Ventral arm plates pentagonal, flat distally, narrowly contiguous. Arm spines, up to three, pointed, about as long as segment. Tentacle scales, two, large covering pore.

Global distribution

Arabian Sea, Persian Gulf, Bay ofBengal, Red Sea, Madagascar, Australia, Mozambique, Indonesia, Philippines, Australia, Fiji and Japan (Cherbonnier & Guille 1978; Baker 1979; Clark & Rowe 1971; Rowe & Gates 1995), South Africa.

Ecology

Depth range: 0-82 m.

Habitat: Associated with seagrass (Syringodium iso-etifolium and Cymodocea serrulata), mud and sand (Cherbonnier & Guille 1978), mud (James 1970), brown mud and detritus.

Remarks

Mortensen (1940) stated that the marginal spines are not diagnostic for hastatus, given that specimens from the Persian Gulf are inconsistent. Later H.L. Clark (1946) stated that it was doubtful that hastatus could be distinguished from A.(Lymanella) depressus because they have relatively broad radial shields, no distinct median distal angle on their dorsal arm plates and often the marginal scales are more or less specialized. In addition, he also mentioned that the prominent primary rosette is not a reliable character because ofthe likelihood ofloss and regeneration; this was further supported by A.M. Clark (1970). Clark & Rowe (1971) distinguished hastatus from depressus using the distinctness of the primary rosette, the size of the radial shields and whether there were spines on the disc margin, all of which were considered unlikely to be distinguishing characters by Mortensen (1940), H.L Clark (1946), A.M. Clark (1970) and Baker (1979). Therefore, based onthe material at hand and these characters it is believed that hastatus is a synonym of depressus.

A.M. Clark found that the South African records of Amphioplus (Lymanella) hastatus reported by Day & Morgans (1956) and by Day (1974) are in fact misidentifica-tions of Amphioplus (Lymanella) integer (Clark & Courtman-Stock 1976). According to Rowe & Gates (1995) the type locality is Mozambique (Ljungman 1867).

Ophionephthys lowelli A.M. Clark, 1974

Plate 3A, B

Ophionephthys lowelli Clark, 1974: 462-464, figs 10a-e; Clark & Courtman-Stock, 1976: 103, 116-117, 159-160; Mbongwa, 2013: 15.

Material/Records

SAM A22782 (disintegrated holotype), off East London, depth 55 m, -33.0500°, 27.9000°, dredge, brown sand and shell, 17 July 1959, station number SCD82P; SAM A22781 (paratype), NE of East London, depth 55 m, -32.5505°, 28.6352°, dredge, sand and mud, 16 July 1959, station number SCD74S; SAM A74075, Sodwana Bay, depth 0 m, -27.5396°, 32.6804°, by hand, collected by Rebecca Milne, among algae, 15 October 2010, identified by J.M. Olbers; Unaccessioned (UKZN), Sodwana Bay, no depth, -27.7320°, 32.6264°, SCUBA, 27 February 2013, identified by J.M. Olbers.

Diagnosis

(See Clark 1974; Clark & Courtman-Stock 1976)

D.D. up to 8 mm. Disc round, in all specimens at hand the dorsal disc 'lid' is missing. Oral shields variable, as long as wide or wider, triangular with broadly rounded angles, widest proximally or rhombic with proximal lobe being flattened. Adoral shields triangular, widely separated inter-radially, with broad distal lobe contiguous with lateral arm shield. Jaws slightly sunken, with two large, broad infradental oral papillae appearing in preserved specimens to be apical papillae. Two spiniform, rugose-tipped oral papillae, one shorter than the other and both attached to oral plate and in series with infradental papillae. Oral tentacle scale distinct, short and sharp situated close to teeth. No genital papillae, genital slits small and indistinct. Arms long, approximately ten times disc length, first seven to nine dorsal arm plates rudimentary, showing underlying structure, plates becoming whole, square or slightly longer than broad, with rounded edges slightly convex on distal side and concave on proximal side, broadly contiguous. Ventral arm plates similar in shape, convex distally, overlapping each other, longer than broad. Arm spines four or five, lowest one thick, blunt, approximately segment length, remaining spines slightly shorter and tapering but blunt, covering not smooth, slightly rough. Tentacle scale single, oval, longer than broad, moderate in length, approximately half segment length.

Global distribution

South Africa (endemic).

Ecology

Depth range: 0-55 m.

Habitat: Found in brown sand, shell and mud and coral sand (A.M. Clark 1974; Clark & Courtman-Stock 1976).

Remarks

During this study, the holotype was borrowed for examination but found to have disintegrated in the jar. Ifa neotype were required to be erected, it is suggested that one of the seven paratype specimens (SAM A22781) be selected because these were collected 90 km from the original type locality and appear to be in reasonable condition.

A.M. Clark (1974) mentioned that her figure ofthe dorsal disc was reconstructed because the upper side was probably covered with a continuous extremely fine scaling, towards the periphery where it turns brown when partially dried. No specimens available for examination here (paratypes and one new specimen) had any dorsal disc resemblance to the reconstructed dorsal disc in A.M. Clark (1974), i.e. they were all missing their dorsal disc 'lids'. The single fresh specimen available for examination was uniformly white, both dor-sally and ventrally, with no distinct markings or colouration.

Family OPHIACTIDAE Matsumoto, 1915

Ophiactis cf. picteti (de Loriol, 1893)

Plate 3C, D

Ophiocnida picteti de Loriol, 1893b: 405-407, pl. 13, fig. 2. Ophiactis picteti: H.L. Clark, 1915: 267; Clark &Rowe, 1971: 82,104; Cherbonnier&Guille, 1978: 123-125, fig. 56; Sloan et al., 1979: 101-102; Humphreys, 1981: 10, 21; Milne, 2012: 155.

Material/Records

RMCA MT2275, -30.9620°, 30.3050°, Trafalgar, KZN, depth25 m, 21 August 1999, SCUBA, collected by Y. Samyn, determined by M. Garcia. SAMA28136,-27.5166°, 32.6833°, Sodwana Bay, depth 15 m, 28 July 1976, SCUBA, J.L.B. Smith Institute, determined by J.M. Olbers. SAM A74053, SAM A74056, SAM A74060,SAM A74070, SAM A74071,SAM A74072 and SAM A74073, -27.5230°, 32.6920°, Sodwana Bay, depth 7.5 m, 15 October 2010, SCUBA, ACEP II, determined by J.M. Olbers. SAM A74063 and SAM A74065, -27.5230°, 32.6920°, Sodwana Bay, depth 12.5 m, 15 October 2010, SCUBA, ACEP II, determined by J.M. Olbers. SAM A74057, -27.5230°, 32.6920°, Sodwana Bay, depth 22 m, 15 October 2010, SCUBA, ACEP II, determined by J.M. Olbers. SAM A74069, -27.5230°, 32.6920°, Sodwana Bay, depth unknown, 15 October 2010, SCUBA, ACEP II, determined by J.M. Olbers.

Diagnosis

(See Cherbonnier & Guille 1978)

D.D. up to 6 mm. Disc round, dorsally covered with overlapping scales, many conical small spinelets mainly in interradial areas and on margin. Ventral interradial areas with finer scales, scattered conical spines. Arms five, simple. Radial shields elongated, narrow triangular, length at least two-thirds disc radius, each pair separated by four enlarged scales, distally approximating or contiguous, light patch on distal part of each radial shield. Genital slits ending at edge of disc, no distinct scales, genital papillae absent. Oral shields spearhead-shaped or oval, slightly wider than long, may be truncated on distal side; adoral shields contiguous interradially, single apical papillae, two to three distal oral papillae. Up to seven arm spines (usually six), short, longest less than twice segment length, tapering to blunt tips, three uppermost ones stout and conical and rugose, remaining spines elongated and decreasing in size toward ventral side. Dorsal arm plates oval, becoming elliptical, wider than long, distal edge convex, broadly contiguous. Ventral arm plates hexagonal, edges rounded in proximal part ofarm, becoming flat-truncated on both sides, slightly wider than long. Single tentacle scale, large, round. Colour in life: disc and arms brown with white, marbled, arms banded sometimes with dark spots, ventrally arms white, spinelets white.

Global distribution

East Indies, Indo-Malayan Region, Western Indian Ocean, Australia, Madagascar, Kenya and Tanzania (Clark & Rowe 1971; Rowe & Gates 1995; Cherbonnier & Guille 1978; Humphreys 1981), South Africa (Milne 2012).

Ecology

Depth range: 0-50 m. Habitat: Coral patches, coral reef flats (Humphreys 1981).

Remarks

In South Africa, as Sloan et al. (1979) suggested, O. picteti has also been misidentified as O. hemiteles in some works. Milne (2012) reported O. hemiteles collected in Sodwana Bay (initially identified by J.M. Olbers), but these were re-examined again by J.M. Olbers and were found to be O. picteti based on the presence of the narrow median distal lobe (Sloan et al. 1979) and the blunt arm spines.

Clark & Rowe (1971) used skin-covered oral shields as a character but in the specimens at hand, which were dry on examination, these do not show the oral shield obscured. In addition, Cherbonnier & Guille (1978) mentioned that the adoral shields are small but in the specimens at hand, the adorals could be adequately seen. Although the arms are broken on all specimens at hand, it is noted that de Loriol (1893) also suggested that the disc diameter and arm length ratio was approximate in the descriptions, which is D.D.:A.L. = 1:6. In addition, Sloan et al. (1979) stated that the Indian Ocean specimens have a narrow median distal lobe on the oral shields which is true of the specimens at hand.

According to Rowe & Gates (1995) the type locality is Amboina, Indonesia, and was thought to be held at the Natural History Museum in Genève but upon investigation, this was not the case (Jean Mariaux, pers. comm).

Family OPHIOCOMIDAE Ljungman, 1867

Ophiocomella sexradia (Duncan, 1887)

Plate 3E, F

Ophiocnida sexradia Duncan, 1887: 92-93, pl. 8, fig. 10, 11; Koehler, 1905: 33.

Ophiocoma parva H.L. Clark, 1915: 292, pl. 14, figs 8, 9; H.L. Clark, 1921: 132, pl. 13, fig. 4; H.L. Clark, 1938: 331-332; A.H. Clark, 1939: 5-7, pl. 1, figs 1, 2; H.L. Clark, 1946: 247; Balinsky, 1957: 27; Kalk, 1958: 207, 216, 237; Macnae & Kalk,1969: 104, 106, 130; Clark & Rowe, 1971: fig. 38d.

Amphilimna sexradia: H.L Clark, 1915: 259.

Amphilimna sexradiata: Koehler, 1927: 3.

Ophiocomellaclippertoni: A.H. Clark, 1939: 5-7 (as O.parva); A.H. Clark, 1952: 296.

Ophiocomella schultzi A.H. Clark, 1941: 481-483; Clark & Rowe, 1971, figs 38c, e.

Ophiomastix sexradiata A.H. Clark, 1952: 297-298; Clark &

Rowe, 1971: 86, 118, figs 38a, b. Ophiocomella sexradia: Clark & Rowe, 1971: 86-87, 118, figs 38c-f; Devaney, 1974: 162-164; Clark & CourtmanStock, 1976: 105, 122, 175; Hughes & Gamble, 1977: 355; Cherbonnier&Guille,1978: 178-179, pl. 12, figs 5, 6; Sloan

et al., 1979: 109; Vine, 1986: 195; Sastry, 1991: 374, 382, pl. 4, fig. 20; Rowe & Gates, 1995: 389; Richmond, 2002: 326; Putchakarn & Sonchaeng, 2004: 423; Stöhr et al., 2008: 547, 555-556; Mbongwa, 2013: 16.

Material/Records

EKZNW RR_9_JMO_2010, -29.9861°, 30.9645°, Reunion Rocks, intertidal, 27 February 2010, collected and determined by J.M. Olbers. EKZNW RR_7_JMO_2010, -29.9861°, 30.9645°, Reunion Rocks, intertidal, 30 January 2010, collected and determined by J.M. Olbers. EKZNW RR_4_ JMO_2010, -29.9861°, 30.9645°, Reunion Rocks, intertidal, 24 September 2010, collected and determined by J.M. Olbers.

Diagnosis

(See Clark & Courtman-Stock 1976; Cherbonnier & Guille 1978)

D.D. up to 6 mm. Disc covered with short blunt spines, densities may differ. Radial shields not distinct. Oral shields variable, round, rhombic, spearhead-shaped or hexagonal. Adoral shields not contiguous. Dental papillae four to six, usually in series. Oral papillae three. Teeth blunt and wide. Genital slits narrow and elongated. Arms six, rarely three or seven. Dorsal arm plates fan-shaped, as wide as long. Ventral arm plates squarish, distal edge rounded, proximal edge truncated. Arm spines, up to four, sometimes five, tapering to blunt tip or may be square tipped, one segment length. Tentacle scale, one, oval, first pair of pores may have two. Fissiparous. Colour in life: disc dark brownish or green, arms banded with brown, green or red.

Global distribution

Indo-Pacific, Mozambique, China, India, south Japan, Australia, Tasman Sea, Reunion, Rodrigues, Hawaiian Islands (Richmond 2002; Clark & Rowe 1971; Rowe & Gates 1995; Rowe & Richmond 2004; Sastry 1991), South Africa (Mbongwa 2013).

Ecology

Depth range: 0-33 m (Koehler 1905; Rowe & Gates 1995). Habitat: Associated with sponges, coral bases and sea grass beds, algae.

Remarks

In KZN, this species appears to be associated with the six-armed species Ophiactis savignyi found in and among rocky shore algae scrapings.

According to Rowe & Gates (1995), the type locality is Mergui Archipelago, Burma (Myanmar).

Ophiomastix koehleri Devaney, 1977

Plate 4A, B

Ophiomastix koehleri Devaney, 1977: 274-283, fig. 1-4; Cherbonnier&Guille, 1978: 186-188, pl. 11, figs 1, 2; Sloan et al., 1979: 92, 109, fig. 16; Humphreys,1981: 10, 25.

Material/Records

RMCA MT2146, -30.2637°, 30.8264°, Aliwal Shoal, depth 15 m, August 1999, collected by Y. Samyn and E. Vanden Berghe, determined by Y. Samyn. RMCA MT2161,

-27.4472°, 32.7167°, Sodwana Bay, 7-mile Reef, depth 18 m, February 2001, collected by Y. Samyn, determined

by Y. Samyn and M. Garcia. SAM A28130, -27.5166°,

32.6833°, Sodwana Bay, depth 15 m, 25 July 1976, SCUBA, J.L.B. Smith Institute, determined by J.M. Olbers. SAM A28120, -27.5166°, 32.6833°, Sodwana Bay, depth 14 m, 23 July 1976, SCUBA, J.L.B. Smith Institute, determined by J.M. Olbers.

Diagnosis

(See Devaney 1977; Cherbonnier & Guille 1978)

D.D. up to 23 mm. Disc round and puffy, dorsally disc covered uniformly by short, rounded granules, ventrally disc with similar granules but not extending up to oral shields, leaving a broken wide V-shaped interradial area with scales, dark brown, variegated with whitish grey. Oral shields round with dark patch on each surrounded by white on margin, adorals small, not contiguous. Genital slits large, almost reaching disc margin, genital papillae present, extending to oral shields. Dorsal arm plates fan-shaped, much wider than long, convex distally, most often a thin white line bordering the plates, narrowly contiguous. Ventral arm plates fan-shaped with convex distal edges, brown with small grey patch surrounded by white margin. Arm spines three to four on each side of same or mostly adjacent segments, often alternating, uppermost spine markedly longer, cigar-shaped, clavate distally and more or less bifurcate at tip, broadly banded with bands becoming more obvious distally, up to five times segment length, remaining spines cigar-shaped with blunt tip, greyish bands not always around full circumference of spine, two to three times segment length, shortest being 1.5 times segment length. Tentacle scales two, becoming one after approximately one third of arm length, oval, similar in size. Colour in life: uniformly dark purple, brown, black with white edges, dorsal arm plates off-white with large irregular purple patches, giving arm banded appearance. Upper arm spines pale or purple mottled, clavate, remaining arm spines purple and white annulations, tentacle scales banded, oral shields with large dark purple blotches.

Global distribution

Madagascar, Zanzibar, Kenya, Aldabra, Comoros (Cherbonnier & Guille 1978), South Africa.

Ecology Depth range: 0-18 m.

Habitat: Under Porites coral colonies, over sandy gravel in lagoonal seagrass bed (Sloan et al. 1979).

Remarks

According to Devaney (1977) the type locality is Zanzibar and the holotype is held at the Natural History Museum in London (BMNH 1965-6-1-451). The granules on the ventral interradial area do not extend to oral shields but the spines originating from the genital slits do extend up to oral shields.

Ophiomastix venosa Peters, 1851

Plate 4C, D

Ophiomastix venosa Peters, 1851: 464-465; Lütken, 1869: 44; Lyman, 1882: 175; Koehler, 1904b: 73-74, figs 28, 29; H.L. Clark, 1915: 296; H.L. Clark,1921: 134, 138; H.LClark, 1923: 349; Balinsky, 1957: 27-28; Kalk, 1958: 237; Macnae & Kalk, 1969: 130; Clark & Rowe, 1971: 88, 120; Clark & Courtman-Stock, 1976: 105, 122, 176-177, fig. 191; Devaney, 1978: 279, 350-353, figs 41, 42; Cherbonnier & Guille, 1978: 190-192, pl. 14, figs 1-2, fig. 63; Sloan et al., 1979: 109-111; Tortonese, 1980: 117, 128, fig. 12; Humphreys, 1981: 10, 25.

Material/Records

RMCAMT2353, -27.5227°, 32.7129°, Sodwana Bay, depth 21 m, 7 November 2003, collected and determined by Y.

Samyn. SAM A28128, -27.5166°, 32.6833°, Sodwana Bay,

depth 15 m, 25 July 1976, SCUBA, J.L.B. Smith Institute, determined by J.M. Olbers. AM J.10517, -31.9839°, 29.1525°, Coffee Bay, depth unknown, collection date 1974.

Diagnosis

(See Clark & Courtman-Stock, 1976; Devaney, 1978; Cher-bonnier & Guille 1978)

D.D. up to 31 mm. Disc round and puffy, dorsal disc scales fine, light brown, pair ofradiating dark brown lines outlined in white starting from base of each arm and meandering in random pattern. Ventral disc scales lighter brown and coarser in proximal interradial areas, some ovate imbricated scales delimiting periphery of disc. Granules sparsely scattered on both dorsal and ventral sides of disc, with scattered cylindrical spines towards margin of dorsal disc. Radial shields visible, but not distinct. Genital slits large, reach margin of disc, genital papillae absent. Oral shields slightly wider than long, adoral shields triangular, largely separated. Arm spines two to four, alternating in number, cigar-shaped, but tapering with darker longitudinal line; on every two to three segments, upper arm spine is enlarged and has clavate, cloven or digitate tip, c. 3.5-4 times segment length, longitudinal line absent on largest spines, other arm spines approximately two times segment length. Dorsal arm plates broadly fan-shaped, wider than long, broadly contiguous, becoming slightly longer than wide, narrowly contiguous. Ventral arm plates pentagonal but truncated, distal side straight or convex, lateral sides may be concave. Tentacle scales two basally, distally one, ovate. Colour in life: disc light brown with radiating lines on disc, radial shields with black petaloid pattern (Humphreys 1981), arm spines with longitudinal dark stripe, dorsal arm plates brownish with a darker faded line down length of arm, ventrally uniformly light brown.

Global distribution

Aldabra, Kenya, Mascarene Basin, Mozambique, Rodriguez, Seychelles, Somalia, Tanzania, Philippines, Madagascar, Comoros, Bay of Bengal, tropical Indo-Pacific (Clark & Rowe 1971; Cherbonnier & Guille 1978; Devaney 1978; Tortonese 1980), South Africa.

Ecology Depth range: 0-21 m.

Habitat: Found in shallow lagoons, often on sand and rubble, algal carpet, under boulders, coral heads, and Porites in lagoonal seagrass beds (Sloan et al. 1979), may be in same habitat with Ophiocoma scolopendrina (de Lamarck, 1816) (Balinsky 1957).

Remarks

Clark & Courtman-Stock (1976) recorded this species in Mozambique. Type locality is Mozambique.

Family OPHIOTRICHIDAE Ljungman, 1867

Macrophiothrix demessa (Lyman, 1862)

Plate 4E, F

Ophiothrix demessa Lyman, 1862: 82; 1865: 172-173; Marktanner-Turneretscher, 1887: 310; Brock, 1888: 513; Koehler, 1905: 91-92, pl. 9, figs 5, 6; H.L. Clark, 1915: 270; 1921: 109; 1939: 83.

Ophiothrix mauritiensis de Loriol, 1893a: 38, pl. 24, fig. 5. Ophiothrixcoronata Koehler, 1905: 91, pl. 9, figs 8, 9; Koehler, 1922a: 217-218, pl. 40, fig. 5, pl. 41, figs 1-4, pl. 98, fig. 1; Koehler, 1930: 137; Vine, 1986: 195.

Amphiophiothrix demessa: H.L. Clark, 1946: 217; Endean, 1957: 243; Fell, 1960: 24.

Macrophiothrix mossambica Balinsky, 1957: 18, fig. 7, pl. 3, figs 11-12.

Macrophiothrix demessa: A.M. Clark, 1968: 289-291, figs 3e, f, 4h, 5h, 7e; Clark & Rowe, 1971: 82, 114, pl. 37f; Devaney, 1974: 139-140; Clark & Courtman-Stock, 1976: 111, 138, fig. 114; Sloan etal., 1979: 102-103; Cherbonnier & Guille, 1978:151-152, pl. 4,figs 1,2; fig. 61: 7-9; Sastry, 1991: 374, 377, pl. 3, fig 16; Milne, 2012: 155.

Material/Records

RMCAMT2156, -27.5227°, 32.6919°, Sodwana Bay, 2-mile Reef, depth 13 m, 4 September 1999, collected by Y. Samyn and E. Vanden Berghe, determined by M. Garcia. RMCA MT2154, -30.2637°, 30.8264°, Aliwal Shoal, depth 16 m, 15 August 1999, collected by Y. Samyn and E. Vanden Berghe, determined by Y. Samyn and M. Garcia. RMCA MT2166, -26.9335°, 32.8871°, Bhanga Nek, depth 20 m, 14 August 1999, collected by Y. Samyn, determined by M. Garcia and Y. Samyn. SAM A74052, -27.5396°, 32.6804° Sodwana Bay, depth unknown, 15 October 2010, collected by R. Milne, ACEP II, determined by J.M. Olbers. SAM A28145, -29.5500°, 31.4167°, Ballito, depth 60-64 m, 7 June 1988, Meiring Naude, station number XX131, Natal Museum Dredge, determined by J.M. Olbers. EKZNW LR_3_ME_2008, -27.7167°, 32.6501°, NE of Liefeldt's Rocks, depth 50 m, 8 June 1988, Meiring Naude, Natal Museum Dredge, determined by J.M. Olbers.

Diagnosis

(See Clark & Courtman-Stock 1976; Cherbonnier & Guille 1978).

D.D. up to 12 mm. Disc puffy, covered dorsally with long thornystumps, two to six terminal points, ventrallycovered with similar stumps, but more scattered and typicallysingle terminal points. Radial shields triangular, two-thirds length of disc radius, covered with shorter and less numerous stumps than those on disc. Jaws elongated, oral shields broadly triangular, much wider than long. Adoral shields not contiguous. Genital slits halfway to disc, genital plate large, adjacent to slit, with disc spinelets not continuing to edge of genital slit. Dorsal arm plates broadly fan-shaped, about twice as wide as long or wider, broadly in contact, armed with small rugose granules or sparse stumps. Ventral arm plates wide, fan-shaped, but with rounded proximal edges, as long as wide, sometimes slightly longer than wide, contiguous. Arm spines up to 14, thorny over entire length, glassy, longest spine at least three times segment length, shortest ventrally. Tentacle scale one, triangular. Colour in life: dorsal side ofdisc greyish with more or less conspicuous dark pink spots, ventrally lighter and fewer spots. Arms banded purple, pink or red with white dorsally and lighter ventrally, with two to three arm segments between bands.

Global distribution

Tropical Indo-west Pacific, Australia, Hawaiian Islands, Mozambique, Mauritius, Zanzibar, Red Sea, India, Seychelles, Maldive Islands, Philippines, China Sea (Rowe & Gates 1995; Sastry 1991; Clark & Rowe 1971; Hoggett 1991), South Africa (Milne 2012).

Ecology

Depth range: 0-128 m.

Habitat: Concealed in coral, deep rocky crevices, under stones with coarse sand and with Lithothamnion (Hoggett 1991).

Remarks

Recorded from Mozambique in Clark & Courtman-Stock (1976). According to Rowe & Gates (1995) the type locality is in the Hawaiian Islands (recorded as Sandwich Islands). Specimens are held at the Museum ofComparative Zoology (holotype: MCZ 2278; paratypes: MCZ 2279, MCZ 2280, MCZ 2281 and MCZ 4095).

Macrophiothrix propinqua (Lyman, 1862)

Plate 5A, B

Ophiothrix propinqua Lyman, 1862: 83-84; 1874: 234; Ljungman, 1867: 333; Marktanner-Turneretscher, 1887: 308; Brock, 1888: 510; Koehler, 1898: 98-100, pl. 3, figs 20-22; Koehler, 1905: 81; H.L. Clark, 1915: 277; H.L. Clark, 1921: 113; Koehler, 1922a: 256-257, pl. 38, figs 1-2, pl. 101, fig. 4; Murakami, 1943a: 207-208.

Ophiothrix triloba von Martens, 1870: 260-261.

Ophiothrix bedoti de Loriol, 1893b: 420-422, pl. 15, fig. 1.

Ophiothrix schmidti Djakonov, 1930: 237-239, pl. 12: 1, 2. Ophiotrichoides propinqua: H.L. Clark, 1946: 232; Balinsky, 1957: 21; Endean, 1957: 244.

Macrophiothrix schmidti: A.M. Clark, 1966: 649.

Ophiothrix (Keystonea) propinqua: A.M. Clark, 1966: 648; A.M. Clark, 1968: 283, fig. 2e; Clark & Rowe, 1971: 86-87, 107; Clark & Courtman-Stock, 1976: 102, 111, 145; Gibbs et al., 1976: 127; Cherbonnier & Guille, 1978: 149, pl. 5, figs 1, 2, fig. 61: 11, 12; Sloan etal., 1979: 103; Guille & Wolff, 1984: 6.

Ophiothrix (Placophiothrix) westwardi Devaney, 1974: 143148, figs 8-14.

Macrophiothrix propinqua: A.M. Clark, 1980: 537; Guille & Vadon, 1985: 62; Hoggett, 1991: 1130-1133, fig. 28, 29; Sastry, 1991: 378, pl. 3, fig 18; Milne, 2012: 155.

Material/Records

RMCAMT2216, -27.5227°, 32.6919°, Sodwana Bay, depth unknown, 10 April 1999, collected by Y. Samyn and E. Vanden Berghe, determined by Y. Samyn and M. Garcia.

RMCA MT2261, -30.2637°, 30.8264°, Aliwal Shoal, depth 20 m, 14 August 1999, collected by Y. Samyn and E. Vanden Berghe, determined by Y. Samyn and M. Garcia. SAM A28123, SAM A28124, and SAM A28126, -27.5166°, 32.6833°, Sodwana Bay, depth 15 m, 24 July 1976, SCUBA, J.L.B. Smith Institute, determined by J.M. Olbers. SAM A28134, -27.5166°, 32.6833°, Sodwana Bay, depth 15 m, 25 July 1976, SCUBA, J.L.B. Smith Institute, determined by J.M. Olbers. EKZNW SB_1_ME_2008, -27.5227°, 32.6919°, Sodwana Bay, depth 10-15 m, 12 September 1987, Meiring Naude, Natal Museum Dredging Programme, collected by D. Herbert, R. Broker and M. Mander, determined by J.M. Olbers. SAM A74051, -27.5396°, 32.6804°, Sodwana Bay, depth unknown, 15 October 2010, ACEP II, collected by R. Milne, determined by J.M. Olbers. SAM A74055, -27.5230°, 32.6920°, Sodwana Bay, depth 7.5 m, 15 October 2010, ACEP II, collected by R. Milne, determined by J.M. Olbers. SAM A74039, -26.9002°, 32.9180° SE of Kosi Bay, depth 50 m, 3 June 1990, Meiring Naude, station number ZA37, Natal Museum Dredging Programme, determined by J.M. Olbers. EKZNW SX_10_2014, -26.9266°, 32.8866°, Saxon Reef, depth 9 m, 4 December 2014, collected by J.M. Olbers, determined by J.M. Olbers.

Diagnosis

(See Clark & Courtman-Stock 1976; Cherbonnier & Guille 1978)

D.D. up to 12 mm. Disc round, puffy, scales clearly visible, with or without armament. Radial shields large, more than half disc radius, naked, disc scales in single or multiple rows between radial shields, some with tubercles or spines. Ventral interradial areas with tubercles, but somewhat sharper than dorsal ones. Oral shields diamond-shaped, wider than long, adoral shields sometimes touching. Genital slits narrow, reaching margin of disc, genital papillae absent, genital plate conspicuous. Dorsal arm plates fan-shaped to elliptical, much wider than long, especially distally, with proximal edge short, distal edge slightly trilobed, broadly contiguous for more than one third of their breadth, some proximal-most plates contain point on distal end. Ventral arm plates square and slightly wider than long, most often with slight distal notch. Arm spines up to eight, finely serrated over total length or serrations at apex of spine, orientated proximally, glassy, longest two times longer than segment length, shortest on ventral side. Tentacle scale, one, large, oval. Colour in life: pink, purple with patterns on disc, arms banded every three to four segments. Radial shields reddish, sometimes with blue patches, distal edge outlined with white.

Global distribution

Tropical Indo-west Pacific, India, Aldabra, Comoros, Madagascar, Kenya, Mascarene Basin, Mauritius, Mozambique, Red Sea, Seychelles, Somalia, Tanzania, Western Indian Ocean (Rowe & Gates 1995; Richmond 2002), South Africa (Milne 2012).

Ecology

Depth range: 0-80 m.

Habitat: Associated with coral, coral slabs, in crevices.

Remarks

In 1966, A.M. Clark designated propinqua to Ophiothrix (Keystonea), but in 1980, she transferred it to Macro-phiothrix because of their similarities in arm and internal jawstructures. M. propinqua is easilyrecognized from other Macrophiothrix species due to its smaller size, radial shields always naked except for a few stumps near edges and arm spines that are shorter than in other Macrophiothrix species.

According to Rowe & Gates (1995), the type locality is Kiribati (as Kingsmill Ils) with the holotype held at the Museum of Comparative Zoology (MCZ 2399).

Ophiothrix (Acanthophiothrix) purpurea von Martens, 1867

Plate 5C, D

Ophiothrix purpurea von Martens, 1867: 346; Döderlein, 1896: 296, pl. 14, fig. 12, pl. 17, figs 23-23a; Koehler, 1905: 102; H.L. Clark, 1915: 277; Koehler, 1922a: 261, pl. 58, figs 3-4, pl. 101, fig. 6; Vine, 1986: 195.

Ophiothrix fallax de Loriol, 1893a: 47-48, pl. 25, fig. 2.

Ophiothrix lorioli Döderlein, 1896: 297, pl. 14, figs 13a, b, pl. 17, figs 24a, b.

Placophiothrix purpurea: H.L. Clark, 1939: 86-87; Clark & Spencer-Davis, 1966: 599.

Ophiothrix (Acanthophiothrix) purpurea: Clark & Rowe, 1971: 86-87, 112, figs 35d, 36, pl. 15, figs 4, 11; Devaney, 1974: 141-142; Cherbonnier & Guille, 1978: 148-149, pl. 5, figs 5-6, fig. 61: 5-6; Sloan etal., 1979: 103; Rowe & Gates, 1995: 424-425; Humphreys, 1981: 23; Price & Rowe, 1996: 75; Putchakarn & Sonchaeng, 2004: 422; Guille & Vadon, 1985: 63.

Material/Records

RMCAMT2185, -27.5227°, 32.6919°, Sodwana Bay, depth 12 m, 8 April 1999, SCUBA, collected by Y. Samyn and E. Vanden Berghe, determined by M. Garcia. RMCA MT2211, -27.5227°, 32.6919°, Sodwana Bay, depth 16 m, 15 August 1999, SCUBA, collected by Y. Samyn and E. Vanden Berghe, determined by M. Garcia. RMCA MT2288, -27.5227°, 32.6919°, Bhanga Nek, depth 15 m, 10 September 1999, SCUBA, collected by Y. Samyn and E. Vanden Berghe, determined by M. Garcia. RMCA MT2289, -26.9335°, 32.8871°, Bhanga Nek, depth 29 m, 16 August 1999, SCUBA, collected by Y. Samyn, P Timm and E. Vanden Berghe, determined by Y. Samyn and M. Garcia. RMCA MT2357, -27.5227°, 32.6919°, Sodwana Bay, depth unknown, 19 February 2003, SCUBA, collected by Y. Samyn and E. Vanden Berghe, determined by M. Garcia. SAM A28122, RW76-14_1, -27.5166°, 32.6833°, Sodwana Bay, depth 15 m, 24 July 1976, SCUBA, J.L.B. Smith Institute, determined by J.M. Olbers. EKZNW SB_2_ME_2008, -27.5333°, 32.6910°, Sodwana Bay, depth 13 m, 6 July 1987, SCUBA, collector G. Williams, determined by J.M. Olbers. Data Record Echino- MAP 445, -27.5265°, 32.6822°, Sodwana Bay, Bikini Reef, depth unknown, 10 May 2013, photographed and determined by C. Griffiths and G. Jones.

Diagnosis

(See Clark & Rowe 1971; Cherbonnier & Guille 1978) D.D. up to 17 mm. Disc pentagonal, dorsally scaled with armament of scattered tubercles and spinelets (long and short) interradially and between radial shields, smaller spinelets on disc margin. Disc ventrally with small spinelets. Radial shields triangular, naked, large, about two-thirds disc radius, more or less conspicuous dark purple stripe along proximal edge, central area more or less variegated whitish and purple, proximal edge concave. Oral shields elliptical, with sharp point on proximal side, much wider than long. Adoral shields contiguous. Genital slits almost to margin, with genital plate from about halfway. Dorsal arm plates hexagonal, distal side convex, as long as wide or longer, consecutive plates in contact for less than one-third oftheir widths. Ventral arm plates somewhat fan-shaped, distal side concave, proximal edge convex becoming straight, as long as or longer than wide. Arm spines up to seven, mostly five, glassy, upper spines smooth becoming serrated, up to five times segment length, lower spines shorter and more serrated, lowest arm spine often very short with hooks. Tentacle pores large. Tentacle scale one, small, pointed. Colour in life: dorsally reds, pinks and whites, some with striking lines, arms have thin dark longitudinal line along length of arm both dorsally and ventrally, dorsal arm plates with some lateral whitish patches.

Global distribution

Aldabra, Madagascar, Mascarene Basin, Red Sea, Seychelles, Tanzania, Western Indian Ocean, Australia (Rowe & Gates 1995; Stöhr et al.2014), South Africa.

Ecology

Depth range: 5-508 m (Rowe & Gates 1995). Habitat: Epizoic onMillepora spp., soft corals, gorgonians and crinoids (Sloan et al. 1979; Price & Rowe 1996).

Remarks

Specimens were consistent with descriptions given by Cherbonnier & Guille (1978) and Clark & Rowe (1971). Most distinctive feature of this species is the dark longitudinal lines on both dorsal and ventral arms. Type locality is Amboina, Indonesia (Rowe & Gates 1995).

Ophiothrix (Ophiothrix) echinotecta Balinsky, 1957

Plate 5E, F

Ophiothrix (Ophiothrix) echinotecta Balinsky, 1957:16-17, fig. 6, pl. 3, figs 9-10; Kalk, 1958: 198; Macnae & Kalk, 1969: 99, 106, 129; Clark & Rowe, 1971: 84-85, 109; Clark & Courtman-Stock, 1976: 112, figs 112, 116, 101, 143; Tortonese 1980: 122; Humphreys, 1981: 23.

Ophiothrixechinoteta (Misspelling): Mbongwa, 2013: 16.

Material/Records

RMCA MT2257, -27.9335°, 32.8871°, Bhanga Nek, depth 20 m, 14 August 1999, SCUBA, collected by Y. Samyn and E. Vanden Berghe, determined by Y. Samyn and M. Garcia. SAM A28118, SAM A28125, SAM A28129, SAM A28131, SAM A28135, -27.5166°, 32.6833°, Sodwana Bay, depth 14-15 m, 23 July 1976, SCUBA, J.L.B. Smith Institute, determined by J.M. Olbers. SAM A28137, SAM A28138, SAM A28139, -27.2225°, 32.7966°, Island Rock, depth unknown, 28 July 1976, SCUBA, J.L.B. Smith Institute, determined by J.M. Olbers. SAM A28141, -27.5166°, 32.6833°, Sodwana Bay, depth 14 m, 1 August 1976, SCUBA, J.L.B. Smith Institute, determined by J.M. Olbers. SAM A28146, -29.5500°, 31.4167°, Ballito, depth 60-64 m, 7 June 1988, Dredge, Meiring Naude, Natal Museum Dredging Programme, determined by J.M. Olbers. SAM A28147, -27.5338°, 31.3188°, SE of Sheffield Beach, depth 50 m, 1 June 1990, Dredge, Meiring Naude, station number XX153, Natal Museum Dredging Programme, determined by J.M. Olbers. SAM A74049, -28.5167°, 32.4333°, off Cape St Lucia, depth 55 m, 12 June 1988, Meiring Naude, station number ZO6, dredge, Natal Museum Dredging Programme, determined by J.M. Olbers. Data Record Mbongwa, 2013, -29.9997°, 30.9450°, Isipingo, intertidal, 13 March 2013, collected by N.A. Mbongwa, determined by J.M. Olbers.

Diagnosis

(See Clark & Courtman-Stock 1976)

D.D. up to 8 mm. Disc round, covered with stumps with density on radial shields being slightly less, stumps bicuspid, tricuspid and multifid, stumps reaching ventral interradial areas. Radial shields length about half disc radius, some scattered short stumps, fewer than rest of disc. Oral shields diamond-shaped, twice as wide as long. Adoral shields narrow and contiguous. Genital slits reaching halfway up to margin with distinct genital plate adjacent to slit. Dorsal arm plates fan-shaped, single rugose, short stump on distal point on many plates, narrowly contiguous. Ventral arm plates slightly broader than long with straight or slightly concave distal edge. Arm spines up to ten, serrated for total length, glassy, the middle spine longest, approximately three times segment length, upper spines shorter with uppermost spine being short stumps, lowermost being transformed into a hook. Tentacle scale one, small, maybear one or two sharp points at tip. Colour in life: dorsal disc light and dark greens, greys and blues. Radial shields variegated white and green, slightly darker than remaining disc, dorsal arm plates sometimes with dark transverse line or chevron on distal side. Ventral side slightly lighter.

Global distribution

Madagascar, Kenya, Mozambique, Somalia, Tanzania (Balinsky 1957; Clark & Rowe 1971; Tortonese 1980; Humphreys 1981), South Africa (Mbongwa 2013).

Ecology

Depth range: 0-64 m.

Habitat: Found in rock hollows and under echinoids Echinometra mathaei and Stomopneustes variolaris (Balinsky 1957). May also be found on coarse sand, gravel, shell debris, stones and sponges.

Remarks

Probably the easiest South African Ophiothrix species to identify, due to a rugose stump present on manyofthe dorsal arm plates. Type locality is at Lighthouse Rocks, Inhaca Island, Mozambique. Type specimens are held at the Iziko South African Museum (holotype SAM A22355 and paratypes SAM A22356).

Ophiothrix (Ophiothrix) foveolata Marktanner-Turneretscher, 1887

Plate 6A, B

Ophiothrix foveolata Marktanner-Turneretscher, 1887: 313, pl. 13, figs 32, 33; Koehler, 1905: 76-77; H.L Clark, 1915: 280; Koehler, 1922a: 238-239, pl. 47, figs 4-7, pl. 98, fig. 6; Koehler, 1930: 140; Clark & Spencer Davies, 1966: 599; A.M. Clark, 1966: 647; Clark & Rowe, 1971: 84, 85, 110, pl. 15, fig. 3; Day, 1974: 94; Putchakarn & Sonchaeng, 2004: 423; Mbongwa, 2013: 16.

Ophiothrix insidiosa Koehler, 1898: 92-93, pl. 4, figs 34-36.

Ophiothrixpoecilodisca H.L. Clark, 1915: 276, pl. 13, fig. 5; H.L. Clark,1923: 341.

Placophiothrixfoveolata: Balinsky, 1957: 20, pl. 4, fig. 15; Kalk, 1958: 207, 214; Macnae & Kalk, 1962:111; Macnae & Kalk, 1969: 102, 106, 130.

Ophiothrix (Ophiothrix) foveolata: Day, 1969: 184; Clark & Courtman-Stock, 1976: 101, 112, 143-144, fig. 118.

Material/Records

RMCAMT2174, -27.4132°, 32.7268°, Sodwana Bay, 9-mile Reef, depth 18 m, 12 August 1999, SCUBA, collected by Y. Samyn and E. Vanden Berghe, determined by Y. Samyn and M. Garcia. EKZNW SB_1_JMO_2009, -27.5361°, 32.6851°, Sodwana Bay, depth 9 m, 8 December 2009, SCUBA, collected and determined by J.M. Olbers. SAM A28148, -30.0686°, 31.0508° Amanzimtoti, depth 300-305 m, 9 July 1985, dredge, Meiring Naude, station number XX66, Natal Museum Dredging Programme, determined by J.M. Olbers. SAM A74040, -26.8667°, 32.9168° NE of Kosi Bay, depth 51-53 m, 3 June 1990, dredge, Meiring Naude, station number ZA40, Natal Museum Dredging Programme, determined by J.M. Olbers. SAM A74044, -27.5188°, 32.7022°, off Sodwana Bay, depth 77 m, 2 June 1990, dredge, Meiring Naude, station number ZH18, Natal Museum Dredging Programme, determined by J.M. Olbers. SAM A74045, -27.5168°, 32.7166°, off Sodwana Bay, depth 70 m, 2 June 1990, dredge, Meiring Naude, station number ZH22, Natal Museum Dredging Programme, determined by J.M. Olbers. SAM A74046, -27.5502°, 32.7069°, SE of Sodwana Bay, depth 70 m, 2 June 1990, dredge, Meiring Naude, station number ZH23, Natal Museum Dredging Programme, determined by J.M. Olbers. SAM A74048, -27.7500°, 32.6502°, off Gypsea Hill, depth 110 m, 8 June 1988, dredge, Meiring Naude, station number ZK4, Natal Museum Dredging Programme.

Diagnosis

(See Clark & Courtman-Stock 1976)

D.D. up to 13 mm. Disc round, disc mostly naked covered in scales and some scattered small granules, though peripherally some scattered large trifid stumps, disc scales moderately large. Radial shields triangular, naked, more than two thirds disc radius, single row of scales between radial shields. Oral shields broad diamond-shaped, much wider than long. Adoral shields moderate in size, may or may not be contiguous. Genital slits halfway to disc margin, genital papillae absent, distinct genital plate. Dorsal arm plates fan-shaped, distal edge convex, consecutive plates in contact for less than halftheir width. Ventral arm plates square or rectangular, wider than long, distal edge concave, proximal edge slightly convex or straight. Arm spines up to eight, glassy, serrated, longest four to five times segment length, some spines with dark longitudinal bands and some with clavate tips. Tentacle scale one, small, tapering. Colour in life: ranging from orange, light brown to violet, young specimens may be bright red, radial shields whitish, patterned with dark purple lines and pinkish patches, adradial edges ofradial shields mayhave dark lines. Arms transversed with same striking dark lines as on disc.

Global distribution

Madagascar, Mozambique, Thailand (Cherbonnier & Guille 1978; Clark & Rowe 1971; A.M. Clark 1980; Putchakarn & Sonchaeng 2004; Stöhr et al. 2014), South Africa.

Ecology

Depth range: 9-305 m.

Habitat: Coral reefs, under dead coral blocks, rock crevices and in Cymodocea beds (Clark & Courtman-Stock 1976; Day 1969).

Remarks

Quite similar to M. propinqua, but radial shield size is a useful character to distinguish between species. According to Rowe & Gates (1995), the type locality is Aru Islands, Indonesia.

Family OPHIACANTHIDAE

Ljungman, 1867

Ophiotoma cf. alberti (Koehler, 1896)

Plate 6C, D

Ophiotrema alberti Koehler, 1896: 251; Koehler, 1906: 6; Koehler, 1907: 324; Koehler, 1908a: 612; Koehler, 1909: 196-198; Matsumoto, 1915: 62; Koehler, 1922a: 90; Mortensen, 1927: 183; Gage et al., 1983: 288; Paterson, 1985: 57-58, fig. 3; Smirnov et al., 2014: 197.

Ophiotoma alberti: Madsen, 1951:113; O'Hara & Stöhr, 2006: 75; Martynov, 2010: 18, 92, 97-103, 126, 131, figs 66A-E, fig. 67A, fig. 68, figs 6C, H, figs 13C, E, F, fig. 18L, fig. 28B.

Material/Records

SAM A22112, -33.8116°, 16.5000°, off Saldanha Bay, depth 2730 m, 27 August 1959, trawl, Africana II, station number A193, determined by A.M. Clark.

Diagnosis

(See Paterson 1985; Martynov 2010)

D.D. up to 17 mm. Disc with small scales, scattered spinelets also extending onto ventral interradial areas. Radial shields naked, sometimes visible, rounded pear-shaped. Oral papillae four or five, in continuous series with two scales in oral tentacle pores. Oral shield large, wider than long, nearly D-shape with rounded proximal edge and slight distal lobe. Adoral shields wing-like, proximal edge indented opposite second oral tentacle pore. Dorsal arm plates triangular to bell-shaped, twice as wide as long, nearly contiguous. Ventral arm plates rectangular with obtuse proximal angle and slight indent on distal edge, nearly contiguous on proximal arm. Arm spines four, glassy, smooth, slightly flattened, rounded tip, uppermost longest, up to two segments in length. Tentacle pores large. Tentacle scales needle-like or round, variable, small if not absent, number from zero to five.

Global distribution Rockall Trough, Bay of Biscay, Azores, northeastern Atlantic (Paterson 1985), South Africa (A.M. Clark, unpubl.).

Ecology

Depth range: 1862-4354 m (Smirnov et al. 2014).

Habitat: No information available.

Remarks

A single specimen is catalogued in the SAM collection, identified by A.M. Clark but for reasons unknown not included in the monograph by Clark & Courtman-Stock (1976). According to Paterson (1985), the type locality is North Atlantic Ocean and the type is held at the Natural History Museum in Paris (MNHN).

Ophiotoma cf. gracilis (Koehler, 1914)

Plate 6E, F

Ophiotrema gracilis Koehler, 1914: 112-114, pl. 12, figs 1,2; Paterson, 1985: 54, 58, fig. 23; Borrero-Pérez et al., 2008:181, fig. 7I.

Ophiotoma gracilis: Martynov, 2010: 98, 103, 141.

Material/Records

SAM A22103, -34.6166°, 17.0500°, off Cape Town, depth 2875-2948 m, 8 December 1959, trawl, Africana II, station number A315, determined by A.M. Clark.

Diagnosis

(See Paterson 1985; Martynov 2010)

D.D. up to 11 mm. Disc with small scales, scattered spinelets also extending onto ventral interradial areas, but not up to oral shields. Radial shields naked, elongated triangular, rounded distal margin, separated but diverging. Oral papillae five, conical, in a continuous series with two scales in oral tentacle pores, which are more elongated, almost spiniform. Oral shield large, much wider than long, spearhead-shaped, with distinct distal lobe. Adoral shields elongated, contiguous. Genital slits narrow. Dorsal arm plates triangular, distal edge convex, almost as long as wide, not contiguous. Ventral arm plates pentagonal, proximal edge concave and obtuse, lateral edges excavated by tentacle pores, much longer than wide, not contiguous. Arms relatively slender. Arm spines four, fine, smooth, pointed, uppermost longest, up to 1.5 segments in length. Tentacle pores large. Tentacle scales subequal, five or six, spinose.

Global distribution

Lesser Antilles and Colombia (Borrero-Pérez et al. 2008), South Africa (A.M Clark, unpubl.).

Ecology

Depth range: 490-2948 m (Borrero-Pérez et al. 2008; A.M. Clark, unpubl.).

Habitat: No information available.

Remarks

A single specimen is catalogued in the SAM collection and was identified by A.M. Clark (date unknown), but for unknown reasons not reported in the monograph by Clark & Courtman-Stock (1976). Prior to the study by Borrero-Pérez et al. (2008), the holotype was the only known specimen.

A number of differences occur between Ophiotoma alberti and O. gracilis.InO. gracilis the arms are more slender, arm spines finer, dorsal arm plates narrower, smaller and more widely separated, ventral arm plates longer, the oral tentacle pore scales are different in size and shape to the oral papillae and the oral shield is spearhead-shaped. The tentacle scales in alberti are smaller and while in gracilis they are spinose they are more obvious. On the specimen at hand (SAM A22103), the arm spines are missing.

According to Borrero-Pérez et al. (2008) the type locality is Lesser Antilles and is held at the Smithsonian Institution (USNM 32301) (Koehler 1914).

Family OPHIODERMATIDAE Ljungman, 1867

Ophiochaeta hirsuta Lütken, 1869

Plate 7A, B

Ophiochaeta hirsuta Lütken, 1869: 38, 71; H.L. Clark, 1915: 222; Clark & Rowe, 1971: 127, figs 44a, b; Gibbs et al., 1976: 129; Sloan etal., 1979: 115; Marsh et al., 1993: 62; Rowe & Gates, 1995: 398; Price & Rowe, 1996: 78.

Ophiochaeta boschmai A.H. Clark, 1964: 388-340, fig. 2.

Material/Records

RMCAMT2307, -27.5227°, 32.6919°, Sodwana Bay, depth unknown, 10 August 1999, SCUBA, collected by Y. Samyn, determined by Y. Samyn and M. Garcia. RMCA MT2293, -26.9335°, 32.8871°, Bhanga Nek, depth 29 m, 14 August 1999, SCUBA, collected by Y. Samyn, IP Timm andE. Vanden Berghe, determined by Y. Samyn and M. Garcia. RMCA MT2290, RMCA MT2291, -27.5227°, 32.6919°, Sodwana Bay, depth 8-24 m, 8-10 August 1999, SCUBA, collected by Y. Samyn and E. Vanden Berghe, determined by Y. Samyn and M. Garcia.

Diagnosis

(See Clark & Rowe 1971; Sloan et al. 1979) D.D. up to 7 mm. Disc pentagonal, completely covered in indented granules and long, thin spinelets both dorsally and ventrally, spinelets densest on ventral interradials close to oral shields. Oral shields and adoral shields may have granules, but few if present. Radial shields concealed by granulation and spinelets. Marginal plates covered by rounded and enlarged granules, disc spines dense on disc margin. Genital slits single, up to halfway to margin, genital papillae absent. Oral shields triangular, slightly longer than wide; no supplementary oral shields. Adoral shields large, triangular, not contiguous. Oral papillae five to six, pointed. Teeth three, lowermost bluntly pointed, second square and uppermost pointed. Dorsal arm plates triangular, distal edge straight, proximally narrowly contiguous, distally not contiguous. Ventral arm plates pentagonal, distal edge straight or somewhat convex. Arm spines up to 12, tapering, subequal, all shorter than one segment length. Tentacle scales oval, two proximally, but one along most of arm. Colour in life: grey to brown, mottled, arms banded with dark brown, light brown and white.

Global distribution

Indo-Malayan Region, Western Indian Ocean, Aldabra, Australia, South Pacific Islands, Red Sea (Richmond 2002; Rowe & Gates 1995; Clark & Rowe 1971), South Africa.

Ecology

Depth range: 0-26 m (Rowe & Gates 1995).

Habitat: Associated with Porites coral colonies or on sandy gravel in lagoonal sea grass beds (Sloan et. al. 1979).

Remarks

All specimens from South Africa are housed in the RMCA and are consistent with descriptions given by Clark & Rowe (1971). Type locality is Fiji Islands (H.L. Clark 1915). Rowe & Gates (1995) suggested the material may be in the Natural History Museum of Denmark, but upon investigation, this was found not to be the case (Tom Schi0tte, pers. comm.).

Ophioconis cupida Koehler, 1905

Plate 7C, D

Ophioconis cupida Koehler, 1905: 15-16, pl. 1, figs 19, 20; Clark & Rowe, 1971: 88-89, 127; Cherbonnier & Guille, 1978: 222-223, pl. 16, figs 3, 4; Vine, 1986: 195; Rowe & Gates, 1995: 399. Ophiurodon cupida: Matsumoto, 1915: 84; Matsumoto, 1917: 315.

Ophiurodon cupidum: Koehler, 1930: 278; H.L. Clark, 1939: 95-96; Murakami, 1943a: 213; H.L. Clark, 1946: 255.

Material/Records

SAM A74041, -26.8669°, 32.9167°, NE of Kosi Bay, depth 49 m, 3 June 1990, dredge, Meiring Naude, station number ZA41, Natal Museum Dredging Programme, determined by J.M. Olbers.

Diagnosis

(See Cherbonnier & Guille 1978)

D.D. up to 4 mm. Disc pentagonal, disc almost completely covered in granules both dorsally and ventrally. Radial shields concealed by granulation. Oral shields triangular, wider than long; supplementary oral shields present, but concealed by granulation. Adoral shields relatively large, triangular, not contiguous. Oral shields and adoral shields may have granules, but easily rubbed off. Oral papillae, five to six, pointed. Teeth three, lowermost wide, large, square, edges translucent. Genital slits single, almost up to disc margin, genital papillae absent. Dorsal arm plates fan-shaped with distal point, narrowly contiguous. Ventral arm plates pentagonal, distal edge pointed, narrowlycontig-uous. Arm spines up to eight, tapering, subequal, longest one slightly longer than segment length. Tentacle pores moderatelylarge. Tentacle scales one, elongated oval, translucent. Colour grey or white with large brown patches on radial areas, arms banded with brown.

Global distribution

Australia, Bay ofBengal, Comoros, Red Sea, Madagascar, Philippines, China, Japan, Pacific Islands (Cherbonnier & Guille 1978; Rowe & Gates 1995), South Africa.

Ecology

Depth range: 10-600 m (Rowe & Gates 1995). Habitat: Found amongst sand, kelp and stones.

Remarks

Easily recognizable within the family by the translucent teeth and tentacle scales. According to Rowe & Gates (1995), the type locality is Moluccas and Sulu Archipelago (Indonesia) with the syntypes held at the Naturalis (ZMA.ECH.O 2004, ZMA.ECH.O 2005 and ZMA.ECH.O 2035 (Joke Bleeker, pers. comm.).

Ophiodyscrita acosmeta H.L. Clark, 1938

Plate 7E, F

Ophiodyscrita acosmeta H.L. Clark, 1938: 356-357; H.L. Clark, 1946: 265; Clark & Rowe, 1971: 88-89, 128; Rowe & Gates, 1995: 399; Price & Rowe, 1996: 79; Lane etal., 2000: 483; Marsh & Morrison, 2004: 295,298, 302, 306, 312, 337.

Ophiocryptus pacificus Murakami, 1943b: 188-189, fig. 10.

Ophiostegastus compsus A.M.Clark, 1968: 317-321,fig. 10.

Material/Records

RMCAMT2183,-27.4472°, 32.7167°, Sodwana Bay, 7-mile Reef, depth 23 m, July 2000, SCUBA, collected by Y. Samyn and I. Tallon, determined by F.W.E. Rowe and J.M. Olbers. RMCA MT2184, -27.5227°, 32.6919°, Sodwana Bay, 2-mile Reef, depth 16 m, August 1999, SCUBA, collected by Y. Samyn and E. Vanden Berghe, determined by F.W.E. Rowe and J.M. Olbers.

Diagnosis

(See H.L. Clark 1938)

D.D. up to 8 mm. Disc round to pentagonal, covered with small, slightly-indented granules completelycoveringdorsal and ventral side. Granules on oral shields, adorals, supplementary oral shields and dorsal, ventral and lateral arm plates. Radial shields also concealed, some granules slightly enlarged over marginal area. Oral shields ovate to spearhead-shaped, wider than long. Oral papillae, six to seven, flattened. Genital slit up to two-thirds to margin of disc. Arms slightly flattened. Dorsal arm plates D-shaped, twice as wide as long, with rounded distal edge, with two more-or-less conspicuous whitish patches on distal edge of each plate, covered with granules similar to those on disc. Ventral arm plates more or less rhombic, becoming bell-shaped with distal edges round, not contiguous distally. Lateral plates slightly projecting. Arm spines up to eight, appressed, short, less than half segment length. Tentacle scales, ovate, up to three basally, inner one largest, two becoming one toward distal segments of arm. Colour in life: dorsally brown and grey, ventrally lighter, dorsal disc patchy light and dark. Arms banded with darker bands of three to five segments.

Global distribution

Japan, China and Australia (Clark & Rowe 1971), South Africa.

Ecology Depth range: 0-23 m. Habitat: No detail recorded.

Remarks

Type locality is Broome, Western Australia with the holo-type at the Museum of Comparative Zoology (MCZ 5294) (H.L. Clark 1938).

Ophiopeza spinosa (Ljungman, 1867)

Plate 8A, B

Ophiarachna spinosum Ljungman, 1867: 305. Ophiopeza dubiosa: A.M. Clark, 1968: 313.

Ophiopeza fallax: Lütken, 1869 (non Peters, 1851): 35.

Ophiopezaspinosa: Clark&Rowe,1971: 90-91,127, fig. 44e; Gibbs et al., 1976: 130; Cherbonnier & Guille, 1978: 227-228, pl. 17, figs 3, 4; Tortonese, 1980: 129; Humphreys, 1981: 10; Vine, 1986: 195; Vail & Rowe, 1989: 273-275, fig. 3; Marsh etal., 1993: 62; Rowe & Gates,1995: 400-401; Mbongwa, 2013: 16.

Pectinura spinosa: Lyman, 1874: 221.

Ophiopezella dubiosa de Loriol, 1893a: 7, pl. 23, fig. 2; H.L. Clark, 1909: 120; H.L Clark, 1915: 304.

Ophiopezellaluetken de Loriol, 1893b: 392-394, pl. 13, fig. 1.

Ophiopezella spinosa: H.L. Clark, 1909: 120; H.L. Clark, 1915: 304; H.L. Clark, 1921: 141; Koehler, 1922a: 338-339; H.L. Clark, 1946: 258.

Distichophis clarki Ely, 1942: 46-48, fig. 12.

Material/Records

(See Cherbonnier & Guille 1978)

RMCA MT2181, RMCA MT2254, -27.5227°, 32.6919°, Sodwana Bay, 2-mile Reef, depth 8-24 m, 9-10 August 1999, SCUBA, collected by Y. Samyn and I. Tallon, determined by Y. Samyn. RMCA MT2284, RMCA MT2296, -26.9335°, 32.8871°, Bhanga Nek, depth 20 m, 14 August 1999, SCUBA, collected by Y. Samyn and E. Vanden Berghe, determined by M. Garcia. SAM A28119, -27.5166°, 32.6833°, Sodwana Bay, depth 14 m, 23 July 1976, dredge, J.L.B. Smith Institute, determined by J.M. Olbers. EKZNW LS_1_ME_ 2008, -27.8736°, 32.6036°, Leadsman Shoal, depth 9-14 m, 14 August 1988, collected by D. Herbert, determined by J.M. Olbers. EKZNW SX_ME_2_2008, -26.9311°, 32.8905°, Saxon Reef, depth 13 m, 4 December 2008, collected and determined by J.M. Olbers.

Diagnosis

D.D. up to 11 mm. Disc pentagonal, covered with dense coat of small indented granules, closely packed up to oral shields, extending onto oral plates or jaws. Disc margin with series of plates in interradial areas. Radial shields concealed by granulation. Oral shields spearhead-shaped, supplementary oral shield not covered by granulation, adoral shields present, relatively large, not contiguous, not distinct, covered in granules. Oral papillae, six to seven, elliptical leaf-shaped, pointed but blunt. Teeth four. Genital slits reach halfway to disc margin, genital papillae absent. Dorsal arm plates broadly fan-shaped basally, wider than long, but rounded on distal edge, becoming more typical fan-shaped distally, not contiguous distally. Ventral arm plates bell-or fan-shaped, narrowly contiguous. Arm spines, up to 12, conical, short, half segment length, slightly longer basally, appressed. Tentacles scales two for most of arm length, inner one large, ovate, almost twice as long as outer one. Colour in life: disc grey to brown yellow, sometimes with patchy patterns, arms banded and marbled.

Global distribution

Somalia, Aldabra, Mascarene Basin, Western Indian Ocean, Kenya, Madagascar, Mozambique, Red Sea, Mauritius, Seychelles, Tanzania, Hawaii, southeast Polynesia (Rowe & Gates 1995; Tortonese 1980), South Africa.

Ecology

Depth range: 0-74 m (Rowe & Gates 1995).

Habitat: Found under rocks and coral rubble (Stöhr et al. 2014), amongst mixed algae and on coral reefs.

Remarks

All specimens were consistent with descriptions given by Clark & Rowe (1971) and Cherbonnier & Guille (1978). According to Rowe & Gates (1995) the type locality is Foa Island, Tonga.

Ophiarachna affinis Lütken, 1869

Plate 8C, D

Ophiarachna affinis Lütken, 1869: 34, 98; de Loriol, 1893b: 411-413; H.L. Clark, 1909: 128; Koehler, 1904b: 76-77; H.L. Clark, 1915: 299, pl. 18, figs 1, 2; Koehler, 1922a: 333-335, pl. 4, fig. 1; Koehler, 1930: 271-272, pl. 14, fig. 1; Clark & Rowe, 1971: 88-89, 123, fig. 42a; Devaney, 1974: 175-176; Sloan etal., 1979: 111, figs 17,18; Rowe & Gates, 1995: 395.

Ophiarachna clavigera Brock, 1888: 495-497.

Material/Records

RMCAMT2344, -27.5367°, 32.6900°, Sodwana Bay, depth 31 m, 6 June 2003, SCUBA, collected by Y. Samyn, determined by Y. Samyn and M. Garcia. RMCA MT2140, -27.5227°, 32.6919°, Sodwana Bay, depth 15 m, 10 August 1999, SCUBA, collected by Y. Samyn and E. Vanden Berghe, determined by M. Garcia. SAM A28121, SAM A28132, -27.5166°, 32.6833°, Sodwana Bay, depth 15 m, 23 July 1976, dredge, J.L.B. Smith Institute, determined by J.M. Olbers.

Diagnosis

(See H.L. Clark 1915; Devaney 1974)

D.D. up to 28 mm. Disc round, somewhat puffy, densely covered in round granules both dorsally and ventrally, granules extending onto oral plates. Radial shields not distinct. Oral shields naked, spearhead-shaped with marbled patterns, single supplementary plate, naked, half width of oral shield. Adoral shields small, not contiguous. Oral papillae five to six, shape varies, broad, elliptical leaf-shaped, middle papillae more slender than proximal and distal ones. Oral tentacle scales three, deep in mouth. Teeth five, lowest tooth square becoming pointed. Genital slits long and reach edge of disc margin, genital papillae absent, but disc granules up to edge ofslit. Arm spines up to five, lowermost spine flattened and blunt, others flattened, but conical or tapering, twice segment length, basally lowermost arm spines may reach three to four times segment length. Dorsal arm plates rectangular, with slight distal concave notch on distal side, twice as wide as long proximally, becoming equal distally. Ventral arm plates, square to fan-shaped prox-imally, becoming longer than wide distally, distal edges slightly convex, plates have thin lighter margin along whole arm length. Tentacle scales, two, oval, outer one somewhat rectangular. Colour in life: disc brown and grey with widely-spaced spots both dorsally and ventrally, arms banded with broad dark and light brown bands of four to nine segments, with four longitudinal dark lines down the length of arm. Arm spines annulated with grey and brown.

Global distribution

Indonesia, Aldabra, Australia, Fiji, Samoa, Mozambique, Seychelles, Red Sea in East Indies, Philippines and South Pacific Islands (Clark & Rowe 1971; Richmond 2002; H.L. Clark 1909; Rowe & Gates 1995), South Africa.

Ecology

Depth range: 0-31 m.

Habitat: Found under Porites colonies over sandy gravel, under boulders over sand and rubble and among coral rubble (Devaney 1974; Sloan et al. 1979).

Remarks

Apart from the different oral configuration, this species is similar in colouration to Ophiocoma (Breviturma) doederleini but distinguished by the presence offour longitudinal dark lines along arms.

According to Rowe & Gates (1995) the type locality is Fiji Islands, with the holotype held at the Natural History Museum of Denmark (ZMUC 470) (Tom Schi0tte, pers. comm.)

Ophiarachnella gorgonia (Müller & Troschel, 1842)

Plate 8E, F

Ophiarachna gorgonia Müller & Troschel, 1842:105. Pectinura gorgonia: Lütken, 1869: 15; Lyman, 1882: 15; Koehler, 1898: 58, pl. 2, figs 1, 2.

Pectinura marmorata Lyman, 1874: 222, pl. 5, figs 1-7.

Pectinura venusta de Loriol, 1893a: 16-19, pl. 23, fig. 3.

Ophiarachnella marmorata: H.L Clark, 1915: 305.

Ophiarachnella gorgonia: H.L. Clark, 1909: 123-124; Matsumoto, 1917: 323-324, pl. 6, fig. 7; H.L. Clark, 1921: 141-142, pl. 12, fig. 5, pl. 35, figs 4, 5; Koehler, 1922a: 339-340; H.L. Clark, 1946: 260-261; A.M. Clark, 1965: 66; Clark & Rowe, 1971: 88, 125, fig. 42b, pl. 20, fig. 2; Cherbonnie r& Guille, 1978: 217-218, pl. 15, figs 5, 6; Sloan et al., 1979: 111; Tortonese, 1980: 129; Humphreys, 1981: 10; Irimura, 1982: 66, 67, fig. 39, pl. 13, fig. 6; Guille & Vadon, 1985: 64; Marsh et al., 1993: 62; Rowe & Gates, 1995: 396; Putchakarn & Sonchaeng, 2004: 423.

Material/Records

RMCAMT2139, -27.5227°, 32.6919°, Sodwana Bay, depth 8-24 m, 10 August 1999, SCUBA, collected by Y. Samyn and E. Vanden Berghe, determined by M. Garcia. RMCA MT2256,RMCAMT2266,-30.2637°, 30.8264°, Aliwal Shoal, depth 14-18 m, 3 August 1999, SCUBA, collected by Y. Samyn and E. Vanden Berghe, determined by M. Garcia. RMCA MT2329, Sodwana Bay, depth unknown, 21 July 2000, SCUBA, collected by Y. Samyn and I. Tallon, determined by M. Garcia. RMCA MT2322, RMCA MT2327, -27.5227°, 32.6919°, Sodwana Bay, 2-mile Reef, depth unknown, 12-24 July 2000, SCUBA, collected by Y. Samyn and I. Tallon, determined by M. Garcia. RMCA MT2144, -26.9335°, 32.8871°, Bhanga Nek, depth 20 m, 14 August 1999, SCUBA, collected by Y. Samyn and E. Vanden Berghe, determined by M. Garcia. RMCA MT2328, -27.4472°, 32.7167°, Sodwana Bay, 7-mile Reef, depth unknown, 22 July 2000, SCUBA, collected by Y. Samyn and I. Tallon, determined by M. Garcia.

Diagnosis

(See Cherbonnier & Guille 1978)

D.D. up to 19 mm. Disc rounded with slight indentations on both sides at base of arms, covered in rounded granules dorsal and ventrally, peripheral granules slightly elongated. Radial shields naked, ovate, longer than wide, relatively small. Granules on ventral interradial areas closely packed up to oral shields and onto oral plates or jaws. Oral shields naked, pentagonal, large, supplementary oral shields distal to each oral shield, D-shaped, often equal to length of oral shield. Adoral shields small, not contiguous, triangular. Oral papillae oval and flattened, distalmost broadest. Teeth four to five, lowermost square, others pointed. Genital slits reach disc margin, genital papillae absent, but disc granulation to slit edge. Arms triangular in cross-section for more than half arm length. Arm spines up to 11, appressed to arm, tapering, approximately half segment length. Dorsal arm plates elliptical proximally, twice as wide as long, broadly in contact, becoming fan-shaped and narrowly in contact in distal parts. Distal edge on proximal most segments sometimes scalloped. Ventral arm plates hexagonal, distal edge convex, becoming flattened distally, wider than long proxi-mally, but longer than wide distally. Tentacle scales two for most of the arm length, inner one oval and long, outer one rectangular and slightly pointed. Colour in life: disc green, brown and white with patches both dorsally and ventrally, radial shields may be mottled white, arms banded dark green and white, ventrally uniformly white, with white patches on interradial areas. Arm spines similar in coloration to arm segments.

Global distribution

China, Japan, Australia, East Indies, Philippines, South Pacific Islands, Ceylon, Bay of Bengal, Thailand, Somalia, Tanzania, Kenya, Madagascar, Mozambique, Mascarene Islands, islands of the Western Indian Ocean, Red Sea, Mauritius (Clark & Rowe 1971; Rowe & Gates 1995), South Africa.

Ecology

Depth range: 0-39 m.

Habitat: Under Porites colonies over gravel, beneath encrusting coral colonies, rubble and among algae (Humphreys 1981; Sloan et al. 1979).

Remarks

Some disc scales exposed, but may be a result ofgranules being rubbed off. All South African records held at the Royal Museum for Central Africa.

Hoareau et al. (2013) found three clades within gorgonia, two from the Western Indian Ocean. Based on the colour morphology, it is believed that this species is from Hoareau's lineage number two. However, sequencing the South African O. gorgonia specimens would be required to confirm this.

Ophiarachnella septemspinosa (Müller & Troschel, 1842)

Plate 9A, B

Ophiarachna septemspinosa Müller & Troschel, 1842: 105-106.

Pectinura septemspinosa Lütken, 1869: 15; Lyman, 1882: 17; de Loriol, 1893b: 395, pl. 13, fig. 2; Koehler, 1905: 9.

Pectinura rigida Lyman, 1874: 224. Ophiarachna armata Troschel, 1879: 137-138.

Ophiarachnella septemspinosa: H.L. Clark, 1909: 126; Koehler, 1930: 273; H.L Clark, 1938: 349-350;1946: 263-264; Cherbonnier & Guille, 1978: 218-219, pl. 16, figs 1, 2; Humphreys, 1981: 10; Guille & Vadon, 1985: 64; Marsh et al., 1993: 62; Gates & Rowe, 1995: 397; Mbongwa, 2013: 16.

Material/Records

RMCA MT2137, RMC AMT2143, RMCA MT2138, RMCA

MT2334, -27.5367°, 32.6900°, Sodwana Bay, depth 8-30 m, 10-15 August 1999, SCUBA, collected by Y. Samyn and E. Vanden Berghe, determined by M. Garcia and Y. Samyn.

RMCA MT2295, RMCA MT2245, RMCA MT2244, -30.2637°, 30.8264°, Aliwal Shoal, depth 15-44 m, 5-13 August 1999, SCUBA, collected by Y. Samyn and E. Vanden Berghe, determined by M. Garcia and Y. Samyn. RMCA MT2141, -26.9335°, 32.8871°, Bhanga Nek, depth 16 m, 15 August 1999, SCUBA, collected by Y. Samyn and E. Vanden Berghe, determined by M. Garcia. RMCA MT2350, -27.5367°, 32.6900°, Sodwana Bay, depth 31 m, 7 November 2003, SCUBA, collected by Y. Samyn, determined by Y. Samyn and J.M. Olbers. RMCA MT2348, -30.8332°, 30.4343°, Protea Banks, depth 38 m, 13 November 2003, SCUBA, collected by Y. Samyn, determined by Y. Samyn and J.M. Olbers. EKZNW LSS_4_EKZNW, -27.8667°, 32.6000°, Leadsman Shoal, depth 11 m, 13 October 2010, SCUBA, collected by J. Wood and J. Dives, determined by J.M. Olbers. EKZNW AS_1_JMO_2008, -30.2669°, 30.8001°, Aliwal Shoal, depth 17 m, 14 June 2008, SCUBA, collected byJ. Wood and J. Dives, determined byJ.M. Olbers.

Diagnosis

(See Cherbonnier & Guille 1978)

D.D. up to 38 mm. Disc round, flat, densely covered in granules both dorsally and ventrally, extending onto jaws. Radial shields naked, contrasting in colour with disc, small, circular. Oral papillae three to four, elliptical, slightly pointed. Teeth broad, but not square. Oral shields naked, oval but truncated distally by large supplementary oral shield, as wide as oral shield, some specimens have marbled oral shields. Adoral shields small, not contiguous. Genital slits long and reaching edge of disc margin, genital plate distinct and slightly higher than interradial area. Dorsal arm plates elliptical rectangular, more than twice as long as wide, rounded lateral angles, proximal edges straight, distal margins may be scalloped. Ventral arm plates hexagonal, convex distally, somewhat concave proximally, wider than long, becoming longer toward distal end of arm, tentacle pore indenting lateral edges. Arm spines up to nine, conical or tapering, same length as segment with exception of lowermost arm spine which is twice as long as segment, cigar-shaped, flattened and tip square. Tentacle scales two, oval, outer one somewhat broader than inner, becoming one distally. Colour in life: uniformly grey, red, yellow or greenish, ventrally lighter, arms banded.

Global distribution

Aldabra, Mascarene Basin, Western Indian Ocean, Kenya, Madagascar, Mozambique, Red Sea, Mauritius, Seychelles, Tanzania, China and South Japan, Australia, East Indies, Philippine Islands, Maldives (Rowe & Gates 1995; Clark & Rowe 1971; Cherbonnier & Guille 1978; Richmond 2002), South Africa (Mbongwa 2013).

Ecology

Depth range: 0-55 m.

Habitat: Found under boulders over sand, under coral (Millepora spp.) colonies, in sand channels and rubble areas of reef flats (Humphreys 1981; Sloan et al. 1979).

Remarks

Easily recognizable by the small radial shields and striking colours. The South African specimens are 'very red' in comparison to the Australian red specimens and may represent a cryptic species complex (Tim O'Hara, pers. comm.).

According to Rowe & Gates (1995) the type localityis the Moluccas, Indonesia. Type material is housed in the Naturalis (ZMA.ECH.O 7084 and RMNH.ECH. 3566, Joke Bleeker, pers. comm.).

Family OPHIOLEPIDIDAE Ljungman, 1867

Ophioplocusimbricatus (Müller & Troschel, 1842)

Plate 9C, D

Ophiolepis imbricata Müller & Troschel, 1842: 93-94.

Ophioplocus tessellatus Lyman, 1862: 76-77.

Ophioplocus imbricatus: Lyman, 1865: 69-70; Lyman, 1882: 20, pl. 35, figs 10-12; Studer, 1882: 7; de Loriol, 1893a: 12-13; Bell, 1898: 849; Bell, 1909: 11; Koehler, 1922a: 435-436; Koehler, 1922b: 48, pl. 84, fig. 12; Clark, H.L. 1915: 344; Clark, 1921: 143, pl. 12, fig. 8, pl. 35, figs 1-3; Clark, 1938: 365-366; Clark, 1946: 275-276; Clark & Rowe, 1971: 90-91, 128; Cherbonnier & Guille, 1978: 239-242, fig. 77a-f; Humphreys, 1981: 11; Guille & Vadon, 1985: 64; Rowe, 1989: 287; Putchakarn & Sonchaeng, 2004: 423; Stöhret al., 2008: 547, 553.

Ophioplocus imbricata: Rowe & Gates, 1995: 435.

Material/Records

RMCAMT2306, -27.6180°, 32.6880°, Sodwana Bay, depth 10 m, July 2000, SCUBA, collectedbyY. Samyn andI. Tallon.

Diagnosis

(See Clark & Rowe 1971; Cherbonnier & Guille 1978)

D.D. up to 11 mm. Disc round, disc scales slightly imbricated and distinct, with central plate present, scales naked. Radial shields small, elongated oval, widely separated, naked. Genital slits small, short, quarter length of interradial area, genital papillae present. Oral shields triangular, moderately large, much wider than long, rounded angles, widest distally. Adoral shields relativelywide, maybe contiguous or slightly separated. Jaws slightly sunken, four to five oral papillae, distalmost being broadest, remaining papillae elliptical leaf-shaped, apical papillae bluntly pointed. Teeth four, rounded. Oral tentacle scale inside oral slit. No arm combs. Dorsal arm plates fragmented along entire length of arm, with lateral arm plates becoming more prominent distally. Ventral arm plates slightly wider than long, rectangular and contiguous, becoming triangular and non-contiguous distally, distal edge rounded throughout. Arm spines three, stout, thick, conical, mostly appressed to arms, no longer than one segment length, becoming shorter distally, occasionally lowermost longest. Tentacle scales, two, ovate or similar to spines in shape, rarely three. Colour in life: dark green or grey with irregular patterns and patches on dorsal disc conforming to interradial areas, arms banded, ventrally brown but pale.

Global distribution

Somalia, Aldabra, Mascarene Basin, Kenya, Madagascar, Mozambique, New Zealand, Red Sea, Mauritius, Seychelles, Tanzania, Australia, Reunion, Andaman Sea (Putchakarn & Sonchaeng, 2004; Stöhr et al., 2008; Rowe & Gates, 1995; Stöhr et al., 2014), South Africa.

Ecology

Depth range: 0-197 m (Koehler 1922a). Habitat: Found with grey sand and mud (Koehler 1922a), angiosperm beds, bases of coral or patch reefs (Humphreys 1981).

Remarks

Both Müller & Troschel (1842) and Rowe & Gates (1995) reported the type localities as Mauritius and Timor (Indonesia) which is surprising, as these locations are extremely far apart. A broken non-type specimen was located at the Naturalis which was recorded from Indonesia (Joke Bleeker, pers. comm.).

H.L. Clark (1938) reported examining a number of specimens with colours ranging from grey to those with the dorsal disc tinted with orange and orange or red-orange ventrally, particularly near the oral shields, while others were light brown dorsally and orange-red ventrally.

 

DISCUSSION

Although no species new to science were found during this study, 24 species new to South Africa are documented. This represents more than a 15% addition to the ophiuroid fauna of the region, representing a significant addition to the fauna.

The large number of records from the east coast suggests under-sampling, compared to the south and west coasts, which have historically been more thoroughly sampled (Griffiths et al. 2010). Deep-water sampling is also required, especially on the east coast, and abyssal sampling is virtually non-existent in the whole region (Griffiths et al. 2010). Of the 145 new data records, 137 were from KwaZulu-Natal, four from the Western Cape, four from the Eastern Cape and no records from the Northern Cape.

 

ACKNOWLEDGEMENTS

Financial support for this project came from the Flemish Community (Bilateral (International) Scientific and Technological Cooperation, project numbers BIL98/84 and BIL01/46), Fund for Scientific Research Flanders to Y. Samyn and from a grant to C.L. Griffiths through the National Research Foundation SEAChange Programme. Elizabeth Hoenson (Iziko South African Museum) is acknowledged for patiently assisting in the collection and in sourcing historical records. We are grateful to Jean Harris, Bridget Elliot, John Dives and the Dive Unit at Ezemvelo KZN Wildlife for invaluable logistical assistance. Peter Timm of Triton Diving and Mark Addison of Blue Wilderness are also thanked for leading the dive operations. We are especially thankful to Frank Rowe, who has given invaluable support in identifications and comments during the review process. Didier Van den Spiegel (RMCA) kindly supplied some photographs, reviewed and corrected the earliest drafts and hosted the first author at RMCA on three occasions. Various museum staff and associates are also acknowledged for their assistance with photographs, collection information and literature, Sabine Stöhr (SMNH), Dave Pawson (USNM), Tim O'Hara (NMV), Marc Eléaume (MNHN), Joke Bleeker (Naturalis), Carsten Lüter (ZMB) Jean Mariaux (GMNH) and Tom Schi0tte (ZMUC). The authors are indebted to the reviewers, Tim O'Hara, Frank Rowe and Sabine Stöhr for comments which greatly improved this paper.

 

ABBREVIATIONS

A.L. Arm length.

AM Australian Museum, Sydney, Australia.

BMNH Natural History Museum, London, United Kingdom. D.D. Disc diameter.

EKZNW Ezemvelo KZN Wildlife, South Africa. GMNH Muséum d'Histoire Naturelle (Natural History Museum), Genève, Switzerland.

KZN KwaZulu-Natal.

MCZ Museum of Comparative Zoology, Harvard University, Massachusetts, U.S.A.

MNHN Muséum national d'Histoire Naturelle (Natural History Museum, Paris, France).

Naturalis Naturalis Biodiversity Centre, Leiden, Netherlands (incorporating ZMA Zoölogisch Museum Amsterdam (ZMA) and Rijksmuseum van Natuurlijke Historie (RMNH)).

NMV Museum Victoria, Australia.

RMCA Royal Museum for Central Africa, Tervuren, Belgium.

SAM Iziko South African Museum, Cape Town, South Africa.

UCT University of Cape Town, South Africa.

UKZN University of KwaZulu-Natal, South Africa.

USNM Smithsonian Institution, National Museum of Natural History, Washington, D.C., U.S.A.

ZMB Museum für Naturkunde an der Universität Humboldt zu Berlin, Berlin, Germany.

ZMUC Natural History Museum of Denmark, Copenhagen, Denmark.

 

REFERENCES

BAKER, A.N. 1979. Some Ophiuroidea from the Tasman Sea and adjacent waters. New Zealand Journal of Zoology 6: 21-51.         [ Links ]

BAKER, A.N. 1980. EuryalinidOphiuroidea(Echinodermata)from Australia, New Zealand and the south-west Pacific Ocean. New Zealand Journal of Zoology 7: 11-83.         [ Links ]

BALINSKY, J.B. 1957. The Ophiuroidea ofInhaca Island. Annals of the Natal Museum 14(1): 1-33.         [ Links ]

BELL, F.J. 1888. Descriptions of four new species of ophiuroids. Proceedings of the Scientific Meetings of the Zoological Society of London 1888(2): 281-284.         [ Links ]

BELL, F.J. 1898. On the Actinogonidiate Echinoderms collected by Mr J. Stanley Gardiner at Funafuti and Rotuma. Proceedings of the Zoological Society of London 1898: 849-850.         [ Links ]

BELL, F.J. 1905. On the Echinoderma found off the coast of South Africa: Part 3. Ophiuroidea. Marine Investigations in South Africa 3: 254-260.         [ Links ]

BELL, F.J. 1909. Report on the Echinoderma (other than Holothurians) collected by Mr. J. Stanley Gardiner in the western parts of the Indian Ocean. Transactions of the Linnaean Society of London, Zoology 13(1): 17-22.         [ Links ]

BENHAM, W.B. 1909. Scientific results of the New Zealand Government trawling expedition 1907. Echinoderma. Records of the Canterbury Museum 1: 83-116.         [ Links ]

BILLETT, D.S.M., BETT, B.J., EVANS, R., CROSS, I., TYLER, P.A. & WOLFF, G.A. 2013. The reproductive ecology of deep-sea ophiuroids around the Crozet plateau, southern Indian Ocean, under contrasting productivity regimes. Deep-sea Research II 92: 18-26.         [ Links ]

BORRERO-PEREZ, G.H., BENAVIDES-SERRATO, M., SOLANO, O. & NAVAS, G.R. 2008. Brittle-stars (Echinodermata: Ophiuroidea) from the continental shelf and upper slope of the Colombian Caribbean. Revista de Biologia Tropical 56 (Suppl. 3): 169-204.         [ Links ]

BOLTON, J.J., COPPEJANS, E., ANDERSON, R.J., DE CLERCK, O., SAMYN, Y., LELIAERT, F. & THANDAR, A.S. 2001. Biodiversity of seaweeds and echinoderms in the Western Indian Ocean. South African Journal of Science 97: 453-454.         [ Links ]

BROCK, J. 1888. Die Ophiuriden-fauna des indischen Archipels. Zeitschrift für wissenschaftliche Zoologie 47(3): 465-539.         [ Links ]

CHERBONNIER G. & GUILLE, A. 1978. Echinodermes: Ophiurides. Faune de Madagascar 48. 272 pp. Paris: Editions du C.N.R.S.         [ Links ]

CLARK, A.H. 1939. Echinoderms (other than holothurians) collected on the presidential cruise of 1938. Smithsonian Miscellaneous Collections 98 (11): 1-18.         [ Links ]

CLARK, A.H. 1941. A new brittlestar of the genus Ophiocomella from Canton Island. Journal of the Washington Academy of Sciences 31: 481-83.         [ Links ]

CLARK, A.H. 1952. Echinoderms from the Marshall Islands. Proceedings of the United States National Museum 102(3302): 265-303.         [ Links ]

CLARK, A.H. 1964. Description of two new species of Ophiuroidea collected during the Snellius expedition. Zoologische Mededelingen Leiden 39: 385-390.         [ Links ]

CLARK, A.M. 1952. Some Echinoderms from South Africa. Transactions of the Royal Society of South Africa 33: 193-221.         [ Links ]

CLARK, A.M. 1965. Japanese and other ophiuroids from the collections of the Munich Museum. Bulletin of the British Museum (Natural History) Zoology 13(2): 39-71.         [ Links ]

CLARK, A.M. 1966. Notes on the family Ophiotrichidae (Ophiuroidea). Journal of Natural History13(9): 637-655.         [ Links ]

CLARK, A.M. 1967. Echinoderms from the Red Sea, Part 2 (Crinoids, Ophiuroids, Echinoids and more Asteroids). Bulletin of the Sea Fisheries Research Station, Israel 41: 26-58.         [ Links ]

CLARK, A.M. 1968. Notes on some tropical Indo-Pacific Ophiotrichids and Ophiodermatids. Bulletin of the British Museum (Natural History) Zoology16(7): 277-322.         [ Links ]

CLARK, A.M. 1970. Notes on the family Amphiuridae (Ophiuroi-dea). Bulletin ofthe British Museum (Natural History) Zoology 19(1): 3-81.         [ Links ]

CLARK A.M. 1974. Notes on some echinoderms from South Africa. Bulletin ofthe British Museum (Natural History) Zoology26(6): 421-487.         [ Links ]

CLARK, A.M. 1977. The South African Museum's Meiring Naude Cruises, Part 4. Echinoderms. Annals of the South African Museum 73(6): 133-147.         [ Links ]

CLARK, A.M. 1980. Some Ophiuroidea from the Seychelles Islands and Inhaca, Mozambique. Revue de Zoologie Africaine 94(3): 533-558.         [ Links ]

CLARK, A.M. & COURTMAN-STOCK, J. 1976. The Echinoderms of Southern Africa. 277 pp. British Museum, London.         [ Links ]

CLARK, A.M. & ROWE, F.W.E. 1971. Monograph of Shallow-water Indo-west Pacific Echinoderms. 238 pp. London: British Museum.         [ Links ]

CLARK, A.M. & SPENCER DAVIS, P. 1966. Echinoderms of the Maldive Islands. Annals and Magazine of Natural History 13(8): 597-612.         [ Links ]

CLARK, H.L. 1909. Notes on some Australian and Indo-Pacific echinoderms. Bulletin of the Museum of Comparative Zoology 52(7): 109-135.         [ Links ]

CLARK, H.L. 1911. North Pacific ophiurans in the collection of the United States National Museum. Bulletin of the United States National Museum 75: 1-336.         [ Links ]

CLARK, H.L. 1915. Catalogue of recent ophiurans, based on the collection ofthe Museum ofComparative Zoology. Memoirs of the Museum of Comparative Zoology at Harvard College 25(4): 164-376.         [ Links ]

CLARK, H.L. 1916. Report on the sea lilies, starfishes, brittle stars and sea urchins obtained by the F.I.S. Endeavour on the coasts of Queensland, New South Wales, Tasmania, Victoria, South Australiaand Western Australia. EndeavourResearch 4: 1-123.         [ Links ]

CLARK, H.L. 1921. The echinoderm fauna of the Torres Strait: its composition and its origin. Publications ofthe Carnegie Institute, Washington 10: 1-233.         [ Links ]

CLARK, H.L. 1923. The echinoderm fauna ofSouth Africa. Annals of the South African Museum 13(7): 221-38.         [ Links ]

CLARK, H.L. 1938. Echinoderms from Australia, an account of collections made in 1929 and 1932. Memoirs ofthe Museum of Comparative Zoology at Harvard College 55: 1-596.         [ Links ]

CLARK, H.L. 1939. The John Murray Expedition. Scientific Reports: Ophiuroidea. 6(2): 29-136. London: British Museum (Natural History).         [ Links ]

CLARK, H.L. 1946. The echinoderm fauna of Australia, its composition and its origin. Publications of the Carnegie Institute, Washington 566: 1-565.         [ Links ]

DAY, J.H. & MORGANS J.F.C. 1956. The ecology of South African estuaries 7. The biology of Durban Bay. Annals of the Natal Museum 13: 259-312.         [ Links ]

DAY, J.H. 1969. A Guide to Marine Life on South African Shores. 300 pp. Cape Town: Balkema.         [ Links ]

DAY, J.H. 1974. The ecology of Morrumbene Estuary, Moçambique. Transactions of the Royal Society of South Africa 41: 43-97.         [ Links ]

DE LAMARCK, J.B. 1816. Les Stellérides. Histoire Naturelle des Animaux sans vertèbres 2: 528-568.         [ Links ]

DE LORIOL, P. 1893a. Catalogue raisonné des Echinodermes recueillis par M. V de Robillard à l'île Maurice III. Ophiurides et Astrophytides. Memoires de la Société de Physique et d'Histoire Naturelle de Genève 32(3): 1-63.         [ Links ]

DE LORIOL, P. 1893b. Echinodermes de la Baie d'Amboine. Revue Suisse de Zoologie 1: 359-426.         [ Links ]

DE LORIOL, P. 1899. Notes pour servir a l'histoire des Echinodermes. VII. Mémoires de la Société de Physique et d'Historie Naturelle de Genève 33: 1-34.         [ Links ]

DEVANEY, D.M. 1974. Shallow water asterozoans of south eastern Polynesia. II. Ophiuroidea. Micronesica 10(1): 105-204.         [ Links ]

DEVANEY, D.M. 1977. Ophiomastix koehleri, a new ophiocomid brittle star (Echinodermata: Ophiuroidea) from the Western Indian Ocean. Proceedings of the Biological Society of Washington 90: 274-283.         [ Links ]

DEVANEY, D.M. 1978. A review of the genus Ophiomastix (Ophiuroidea: Ophiocomidae). Micronesica 14(2): 273-359.         [ Links ]

DJAKONOV, A.M. 1930. Echiniden, Ophiuriden und Asteriden gesammelt von Prof. P.J. Schmidt bei den Riu-Kiu Inseln im Jahre 1926-1927. Zoologische Jahrbücher 59: 233-252.         [ Links ]

DÖDERLEIN, L. 1896. Bericht über die von Herrn Professor Semon bei Amboina und Thursday Island gesammelten Ophiuroidea. Denkschriften medizinisch-naturwissenschaftliche Gesellschaft Jena 8: 279-300.         [ Links ]

DÖDERLEIN, L. 1910. Asteroidea, Ophiuroidea, Echinodea. In: Forschungsreise im westlichen und zentralen Südafrika, Schultz, L. (ed.), 4(1) Denkschriften medizinisch-naturwissenschaftliche Gesselschaft Jena 16: 245-258.         [ Links ]

DÖDERLEIN, L. 1911. Japanische und andere Euryalae. 147 pp. Verlag der Bayerischen Akademie der Wissenschaften in Kommission des Verlags R. Oldenbourg.         [ Links ]

DÖDERLEIN, L. 1927. Indopacifische Euryalae. München, Verlag der Bayerischen Akademie der Wissenschaften in Kommission des Verlags R. Oldenbourg. 105 pp.         [ Links ]

DUNCAN, P.M. 1887. On the Ophiuridae of the Mergui Archipelago, collected for the trustees of the Indian Museum, Calcutta, by Dr John Anderson, F.R.S. Superintendent of the Museum. Linnaean Society Journal of Zoology 21: 85-106.         [ Links ]

ELY, C.A. 1942. Shallow-water Asteroidea and Ophiuroidea of Hawaii. Bernice P. Bishop Museum Bulletin 176: 1-63.         [ Links ]

ENDEAN, R. 1957. The biogeography of Queensland's shallow water echinoderm fauna (excluding Crinoidea), with a rearrangement of the faunistic provinces of tropical Australia. Australian Journal of Marine and Freshwater Research 8(3): 233-273.         [ Links ]

FELL, H. B. 1960. Synoptic keys to the genera of Ophiuroidea. Zoology Publications from Victoria University of Wellington 26: 1-44.         [ Links ]

GAGE, J.D., PEARSON, M., CLARK, A.M., PATERSON, G.L.J. & TYLER P.A. 1983. Echinoderms of the Rockall Trough and adjacent areas. 1. Crinoidea, Asteroidea and Ophiuroidea. Bulletin of the British Museum of Natural History, Zoology. London 43(5): 263-308.         [ Links ]

GIBBS, P.E., CLARK, A.M. & CLARK, C.M. 1976. Echinoderms from the northern region of the Great Barrier Reef, Australia. Bulletin of the British Museum (Natural History) Zoology 30(4): 101-144.         [ Links ]

GRAY, J. E. 1840. A synopsis of the genera and species of the class Hypostoma (Asterias Linnaeus). Annals of the Magazine of Natural History 6: 175-184.         [ Links ]

GRIFFITHS, C.L., ROBINSON, T.B., LANGE, L. & MEAD, A. 2010. Marine biodiversity in South Africa: an evaluation of current states of knowledge. PLOS ONE 5(8): e12008. DOI: 10.1371/journal. pone.0012008.         [ Links ]

GUILLE, A & VADON, C. 1985. Les Ophiures littorales de Nouvelle-Calédonie. Bulletin Muséum national d'Histoire naturelle, Paris 7(1): 61-72.         [ Links ]

GUILLE, A. & VADON, C. 1986. Les Ophiurides In: Guide des etoiles de mer, oursins at autres echinoderms du lagon de Nouvelle-Caledonie. 238 pp. GUILLE, A., LABOUTE, P. & MENOU J.L. (eds). Paris: Orstom.         [ Links ]

GUILLE, A. & WOLFF, W.J. 1984. Résultats biologiques de l'expedition Snellius. Echinodermata: Ophiuroidea. Zoologische Verhandelingen 213: 3-39.         [ Links ]

HERTZ, M. 1927a. Die Ophiuroiden der Deutschen Süd polar Expedition. Deutsche Süd polar-Expedition 1901-1903, 19. Zoologie 11: 1-54.         [ Links ]

HERTZ, M. 1927b. Die Ophiuriden der Deutschen Tiefsee-Expedition. I. Chilophiurida Matsumoto (Ophiolepididae, Ophioleucidae, Ophiodermatidae, Ophiocomidae). Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf dem Dampfer 'Valdwia' (1898-1899) 22: 59-122.         [ Links ]

HOGGETT, A.K. 1991. The genus Macrophiothrix (Ophiuroidea: Ophiotrichidae). Invertebrate Taxonomy 4: 1077-1146.         [ Links ]

HUGHES, R.N. & GAMBLE, J.C. 1977. A quantitative survey of the biota of intertidal soft substrata on Aldabra Atoll, Indian Ocean. Philosophical Transactions ofthe Royal Society B 279: 327-355.         [ Links ]

HUMPREYS, W.F. 1981. The Echinoderms of Kenya's Marine Parks and Adjacent Regions. 39 pp. Tervuren, Belgium: Royal Museum of Central Africa.         [ Links ]

IMAOKA, T., IRIMURA, S., OKUTANI, T., OGURO, C., OJI, T. & KANAZAWA, K. 1991. Echinoderms from Continental Shelf and Slope around Japan, Vol. 2. 184 pp. Tokyo: Japan Fisheries Resource Conservation Association.         [ Links ]

IRIMURA, S. 1982. The Brittle-stars of Sagami Bay, Japan.95pp. Japan: Biological Laboratory Imperial Household.         [ Links ]

JAMES, D.B. 1970. Studies on Indian echinoderms. 4. On the brittle-stars Amphioplus gravelyi sp. nov. and Amphioplus depressus (Ljungman) from the Indian coasts. Journal of the Marine Biological Association of India 12: 139-145.         [ Links ]

KALK, M. 1958. Ecological studies on the shores of Mozambique. 1. The fauna of intertidal rocks at Inhaca Island, Delagoa Bay. Annals of the Natal Museum 14: 189-242.         [ Links ]

KOEHLER, R. 1896. Note preliminaire sur les Ophiures des premières campagnes de la 'Princess Alice'. Memoires de la Societé Zoologique de France 9: 241-253.         [ Links ]

KOEHLER, R. 1898. Echinoderms recueillis parl'Investigatordans l'Ocean Indien, II les ophiures littorales. Bulletin scientifique de la France et de la Belgique 31: 55-126.         [ Links ]

KOEHLER, R. 1900. Echinoderma of the Indian Museum. Illustrations of the shallow-water Ophiuroidea collected by the R.I.M.S. Investigator, 1-4, pls 15-21. Calcutta, India.         [ Links ]

KOEHLER, R. 1904a. Ophiures de l'expédition du Siboga. Part 1. Ophiures de mer profonde. Siboga Expedition Monographs 45a: 1-176.         [ Links ]

KOEHLER, R. 1904b. Ophiures nouvelles ou peu connues. Memoirs Société Zoologique de France 17: 54-119.         [ Links ]

KOEHLER, R. 1905. Ophiures de l'expédition du Siboga. Part 2. Ophiures littorales. Siboga Expedition Monographs 45b: 1-142.         [ Links ]

KOEHLER, R. 1906. Description des Ophiures nouvelles recueilles par le Travailleur et Talisman pendant les campagnes de 1880, 1881, 1882 et 1883. Memoires de la Société zoologique de France 19: 5-35.         [ Links ]

KOEHLER, R. 1907. Revision de la collection des ophiures du Museum d'Historie Naturelle de Paris. Bulletin biologique de la France et de la Belgique 41: 279-351.         [ Links ]

KOEHLER, R. 1908a. Astéries, Ophiures et Echinides de l'Expedition Antarctique National Ecossaise. Transactions of the Royal Society of Edinburgh 46(3): 529-649.         [ Links ]

KOEHLER, R. 1908b. Astéries, Ophiures et Échinides recueillis dans les mers australes par la Scotia (1902-1905). Zoologischer Anzeiger 32(6): 140-147.         [ Links ]

KOEHLER, R. 1909. Echinodermes provenant de campagnes du yacht Princesse-Alice. Retsultats Campagnes Scientifiques accomplies sur son yacht par Albert I, Prince Soverain de Monaco 34: 1-317.         [ Links ]

KOEHLER, R. 1914. A Contribution to the Study of Ophiurans of the United States National Museum. 173 pp. Washington DC: Smith sonian Institution, United States National Museum.         [ Links ]

KOEHLER, R. 1922a. Ophiurans of the Philippine seas and adjacent waters. Bulletin of the United States National Museum 100(5): 1-597.         [ Links ]

KOEHLER, R. 1922b. Echinodermata: Ophiuroidea. Scientific Reports of the Australian Antarctic Expedition 1911-1914 under the leadership of Sir Douglas Mawson 8(2): 1-98.         [ Links ]

KOEHLER, R. 1927. Ophiures recueillies aux iles Gilbert, Marshall et Fiji. Meddelanden från Göteborgs musei zoologiska avdelning 33(3): 1-13.         [ Links ]

KOEHLER, R. 1930. Ophiures recueillies par le Docteur Th. Mortensen dans les Mers d'Australie et dans l'Archipel Malais. Papers from Dr. Th. Mortensen's Pacific Expedition 1914-16. LIV. Videnskabelige Meddeleser fra Dansk naturhistorisk Forening i Kjobenhavn 89: 1-295.         [ Links ]

LANE, D.J.W., MARSH, L.M., VAN DEN SPIEGEL, D. & ROWE, F.W.E. 2000. Echinoderm fauna of the South China Sea: an inventory and analysis of distribution patterns. The Raffles Bulletin of Zoology, Supplement 8: 459-493.         [ Links ]

LIAO, Y. 1989. Two new species of the genus Amphilimna (Echinodermata: Ophiuroidea) from southern China. Chinese Journal of Oceanology and Limnology 77(4): 339-344.         [ Links ]

LINDER, H.P. & GRIFFITHS, C.L. 2000 (1999). Introduction and overview. In: GRIFFITHS, C.L. & LINDER, H.P. (eds) Ahistory of Systematic Biology in South Africa. Transactions ofthe Royal Society of South Africa 54(1): 1-4.         [ Links ]

LITVINOVA, N.M., 1971. [The Brittle-stars of the genus Amphiophiura of the Pacific and Indian Oceans collected by soviet expeditions on the Vityaz and Akademik Kurchakov]. Transactions of the P.P. Shirshov Institute of Oceanology 92: 298-316 [In Russian].         [ Links ]

LJUNGMAN, A. 1867. Ophiuroidea viventia huc usque cognita. Ofversigt af Kongl. Vetenskaps-Akademiens Forhandlingar 23(9): 303-336.         [ Links ]

LUDWIG, H. 1901. Echinodermen: Ophiuroidea. In: Dr H.G. Bronn's Klassen und Ordnungen des Thier-reichs, wissen-schalftlich dargesteldt in Wort und Bild: pp. 745-966.         [ Links ]

LÜTKEN, C.F. 1869. Additamenta ad historiam Ophiuridarum. Tredie Afdelning. Det kongelige danske Videnskabernes Selskaps Skrifter. 5 Raekke. Naturvidenskabelig og mathematisk Afdelning 8: 20-109.         [ Links ]

LYMAN, T. 1862. Descriptions of new Ophiuridae. Proceedings of the Boston Society of Natural History 8: 75-86.         [ Links ]

LYMAN, T. 1865. Ophiuridae and Astrophytidae. Illustrated Catalogue of the Museum of Comparative Zoology, Harvard University 1 : 1-200.         [ Links ]

LYMAN, T. 1874. Ophiuridae and Astrophytidae, new and old. Bulletin ofthe Museum ofComparative Zoology, Harvard College 3(10): 221-272.         [ Links ]

LYMAN, T. 1878. Ophiuridae and Astrophytidae of the exploring Voyage of H.M.S. 'Challenger', under Prof. Sir Wyville Thomson, F.R.S. Bulletin of the Museum of Comparative Zoology 5(7): 65-168.         [ Links ]

LYMAN, T. 1879. Ophiuridae and Astrophytidae of the exploring Voyage of H.M.S. Challenger, under Prof. Sir Wyville Thomson, F.R.S. Bulletin of the Museum of Comparative Zoology 6(2): 17-83.         [ Links ]

LYMAN, T. 1882. Challenger Reports: Ophiuroidea. 386 pp. H.M.S. Challenger Reports. WYVILLE-THOMSON, C. (ed.). London: published by Order of her Majesty's Government.         [ Links ]

MACNAE, W. & KALK, M. 1958. A Natural History of Inhaca Island, Mozambique, 1st Edition. 163 pp. Johannesburg: Witwatersrand University Press.         [ Links ]

MACNAE, W. & KALK, M.1962. The fauna and flora ofsand flats at Inhaca Island, Mozambique. Journal of Animal Ecology 31: 93-128.         [ Links ]

MACNAE, W. & KALK, M. 1969. A Natural History of Inhaca Island, Mozambique, revised edition. 113 pp. Witwaters rand University Press, Johannesburg.         [ Links ]

MADSEN, F.J. 1951. Reports ofthe Swedish Deep-sea Expedition 1947-1948: Ophiuroidea. 173 pp. Göteborg: Elanders Boktryckeri aktiebolag.         [ Links ]

MADSEN, F.J. 1967. Ophiuroidea. In: BANZ Antarctic Research Expedition 1929-1931, under the command of Sir Douglas Mawson (BANZARE Reports series B). P.M. THOMAS (ed.) 9(3): 121-145. Adelaide: Mawson Institute for Antarctic Research, University of Adelaide.         [ Links ]

MADSEN, F.J. 1977. The Ophioleucidae. Galathea Report 14: 109-122.         [ Links ]

MANSO, C.L.C. 2010. Deep-water Ophiuroidea (Echinodermata) from off Chile in the Eastern South Pacific. Biota Neotropica 10(2): 185-199.         [ Links ]

MARKTANNER-TURNERETSCHER, G. 1887. Beschreibung neuer Ophiuriden und Bemerkungen zu bekannten. Annalen des Naturhistorischen Museums in Wien, Naturhistorisches Museum (Österreich) 2: 291-316.         [ Links ]

MARSH, L.M., VAIL, L., HOGGETT, A.K. & ROWE, F.W.E. 1993. Echinoderms of Ashmore Reef and Cartier Island, northern- western Australia. Records of the Western Australian Museum Supplement 44: 53-65.         [ Links ]

MARSH, L.M. & MORRISON, S.M. 2004. Echinoderms of the Dampier Archipelago, Western Australia. Records ofthe Western Australian Museum Supplement 66: 293-342.         [ Links ]

MARTYNOV, A. 2010. Reasessment of the classification of the Ophiuroidea (Echinodermata), based on morphological characters. I. General character evaluation and delineation ofthe families Ophiomyxidae and Ophiacanthidae. Zootaxa 2697: 1-154.         [ Links ]

MATSUMOTO, H. 1915. A new classification of the Ophiuroidea: with descriptions of new genera and species. Proceedings ofthe Academy of Natural Sciences of Philadelphia 67(1): 43-92.         [ Links ]

MATSUMOTO, H. 1917. A monograph of Japanese Ophiuroidea, arranged accordingto a new classification. Journal of the College of Science, Imperial University, Tokyo 38(2): 1-408.         [ Links ]

MBONGWA, N.A. 2013. Phylogenetic marker identification in brittle stars (Echinodermata: Ophiuroidea) from the east coast of South Africa. B.Sc. Hons project, School of Life Sciences, University of KwaZulu-Natal.         [ Links ]

McKNIGHT, D.G. 2000. The Marine Fauna of New Zealand: Basket-stars and Snake-stars (Echinodermata: Ophiuroidea: Euryalinida). 79 pp. Wellington: National Institute of Water and Atmospheric Research.         [ Links ]

MILNE, R. 2012. Macro invertebrates associated with macrophytes in Sodwana Bay with further consideration of amphipod taxonomy. M.Sc. dissertation, University of Cape Town, South Africa.         [ Links ]

MORTENSEN, T. 1925. On some echinoderms from South Africa. Annals and Magazine of Natural History 9(16): 146-154.         [ Links ]

MORTENSEN, T. 1927. Handbook of the Echinoderms of the British Isles. 471 pp. Humphrey Milford, Oxford University Press.         [ Links ]

MORTENSEN, T. 1933a. Echinoderms of South Africa (Asteroidea and Ophiuroidea). Papers from Dr. Th. Mortensen's Pacific Expedition 1914. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjobenhavn 93: 215-490.         [ Links ]

MORTENSEN, T. 1933b. Studies of Indo-Pacific Euryalids. Videnskabelige Meddelelser fra Dansk Naturhistorisk Foreningi Kjobenhavn 96: 1-75.         [ Links ]

MORTENSEN, T. 1936. Discovery Reports: Echinoidea and Ophiuroidea. 167 pp. Cambridge University Press.         [ Links ]

MORTENSEN, T. 1940. Echinoderms from the Iranian Gulf. Asteroidea, Ophiuroidea and Echinoidea. Danish Scientific Expeditions in Iran 2: 55-110.         [ Links ]

MÜLLER, J.H. & TROSCHEL, F.H. 1842. System der Asteriden. Braunschweig, 134 pp.         [ Links ]

MURAKAMI, S. 1943a. Report on the ophiurans of Yaeyama, Ryukyu. Journal ofthe Department ofAgriculture, Kyusyu Imperial University 75: 205-222.         [ Links ]

MURAKAMI, S. 1943b. Report on the ophiurans of Palao Caroline Islands. Journal of the Department of Agriculture, Kyusyu Imperial University 7(4): 159-203.         [ Links ]

O'HARA, T. & STÖHR, S. 2006. Deep water Ophiuroidea (Echino- dermata) of New Caledonia: Ophiacanthidae and Hemieuryalidae. Mémoire du Muséum national d´ Histoire naturelle 193: 33-141.         [ Links ]

OLBERS, J.M. & SAMYN, Y. 2012. The Ophiocoma species (Ophiurida: Ophiuroidea) of South Africa. Western Indian Ocean Journal of Marine Science 10(2): 137-154.         [ Links ]

PATERSON, G.L.J. 1985. The deep sea Ophiuroidea of the North Atlantic Ocean. Bulletin of the British Museum (Natural History) Zoology 49(1): 1-162.         [ Links ]

PETERS, W. 1851. Übersicht der an der Küste von Mossambique eingesammelten Ophiuren, unter denen sich zwei neue Gattungen befinden. Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königlich Preussischen Akademie der Wissenschaften zu Berlin 1851: 463-66.         [ Links ]

PRICE, A.R.G. & ROWE, F.W.E. 1996. Indian Ocean echinoderms collected during the Sinbad Voyage (1980-1981): 3. Ophiuroidea and Echinoidea. Bulletin of the Natural History Museum of London (Zoology) 62(2): 71-82.         [ Links ]

PUTCHAKARN, S. & SONCHAENG, P. 2004. Echinoderm fauna of Thailand. Science Asia 30: 417-428.         [ Links ]

RETZIUS, A.J. 1783. Anmärkningar v.d. Asteriae genus. Kungliga Svenska Vetenskapsakademiens handlingar 4: 230-248.         [ Links ]

RICHMOND, M. D. 2002. A Field Guide to the seashores of Eastern Africa and the Western Indian Ocean. 461 pp. Sida/SAREC-UDSM        [ Links ]

ROWE, F.W.E. 1989. Nine new deep-water species of Echino-dermata from Norfolk Island and Wanganella Bank, northeastern Tasman Sea, with a checklist of the echinoderm fauna. Proceedings of the Linnaean Society of New South Wales 111(4): 257-291.         [ Links ]

ROWE, F.W.E. & GATES, J. 1995. Echinodermata. Zoological Catalogue of Australia 33. 505 pp. Melbourne: CSIRO.         [ Links ]

ROWE, F.W.E. & RICHMOND, M.D. 2004. A preliminary account of the shallow water echinoderms of Rodrigues, Mauritius, Western Indian Ocean. Journal of Natural History 38: 3273-3314.         [ Links ]

SAMYN, Y. & THANDAR, A. S. 2003. Towards an understanding of the shallow-water echinoderm biodiversity of KwaZulu-Natal, Republic of South Africa. In: FÉRAL, J.-P. & DAVID, B. (eds), Echinoderm Research. Lisse, Netherlands: Swets & Zeitlinger, 2001: 41-47.         [ Links ]

SASTRY, D.R.K. 1991. Echinodermata: Asteroidea, Ophiuroidea and Echinoidea, In: GHOSH, A.K. (ed), Fauna ofLakshadweep. State Fauna series, 2: pp. 363-397. Calcutta: Zoological Survey of India.         [ Links ]

SINK, K.J., BOSHOFF, W., SAMAAI, T., TIMM, P.G., KERWATH, S.E. 2006. Observations of the habitats and biodiversity of the submarine canyons at Sodwana Bay. South African Journal of Marine Science 102: 466-74.         [ Links ]

SLOAN, N.A., CLARK, A.M. & TAYLOR, J.D. 1979. The echino-derms of Aldabra and their habitats. Bulletin of the British Museum (Natural History) Zoology 37(2): 81-128.         [ Links ]

SMIRNOV, I.S., PIEPENBURG, D., AHEARN, C.G. & JUTER-ZENKA, K.V. 2014. Deep-sea fauna of European seas: annotated species check-list of benthic invertebrates living deeper than 2000 m in the seas bordering Europe. Ophiuroidea. Invertebrate Zoology 11(1): 192-209.         [ Links ]

STÖHR, S., CONAND, C. & BOISSIN, E. 2008. Brittle stars (Echinodermata: Ophiuroidea) from La Réunion and the systematic position of Ophiocanops Koehler, 1922. Zoological Journal of the Linnaean Society 153: 545-560.         [ Links ]

STÖHR, S., O'HARA, T. & THUY, B. (eds) 2014. World Ophiuroi dea database. Accessed at http://www.marinespecies.org/ophiu-roidea on 14 October 2014.         [ Links ]

STUDER, T. 1882. Übersicht über die Ophiuriden, welche während der Reise S.M.S. Gazelle um die Erde 1874-76 gesammelt wurden. Abhandlungen der Koeniglichen Akademie der Wissen schaften in Berlin 34: 1-37.         [ Links ]

THOMAS, L.P. 1967. The systematic position of Amphilimna (Echinodermata; Ophiuroidea). Proceedings of the Biological Society of Washington 80: 123-130.         [ Links ]

THOMAS, L.P. 1975. The Ophiacanthid genus Amphilimna (Ophiuroidea: Echinodermata). Proceedings of the Biological Society of Washington 88: 127-139.         [ Links ]

TORTONESE, E. 1980. Researches on the coast of Somalia. Littoral Echinodermata. Monitore Zoologico Italiano 13(5): 99-139.         [ Links ]

TROSCHEL, F.H. 1879. Eine neue Art der Ophiuridengattung Ophiarachna. Sitzungsberichte der niederrheinischen Gesellschaft in Bonn 1879: 135-138.         [ Links ]

TSURNAMAL, M. & MARDER, J. 1966. Contributions to the knowledge of the Red Sea, No. 33. Observations on the basket star Astroboa (Lyman) on coral reefs at Eilat (Gulf of Aqaba). Israel Journal of Zoology 15: 9-17.         [ Links ]

VADON, C. & GUILLE, A. 1984. Les Ophiuridae (Ophiuroidea, Echinodermata) de la campagne MD 32 du Marion-Dufresne autour de l'ile de La Reunion. Bulletin Muséum national d´ Histoire naturelle, Paris 4(6): 583-615.         [ Links ]

VAIL, L. & ROWE, F.W.E. 1989. Status of the genera Ophiopeza and Ophiopsammus (Echinodermata: Ophiuroidea) in Australian waters, with the description ofa new species. Proceedings of the Linnaean Society of New South Wales 110(3): 267-288.         [ Links ]

VERRILL, A.E. 1899. Report on the Ophiuroidea collected by the Bahama Expedition in 1893. Bulletin from the Laboratories of Natural History of the State University of Iowa 5(1): 1-86.         [ Links ]

VINE, P. 1986. Red Sea Invertebrates. 224 pp. Immel Publishing, London.         [ Links ]

VON MARTENS, E. 1867. Ueber ostasiatische Echinodermen. Archiv für Naturgeschichte 33: 106-120.         [ Links ]

VON MARTENS, E. 1870. Die Ophiuriden des indischen Oceans. Archiv für Naturgeschichte 36: 244-262.         [ Links ]

WORLD CONSERVATION MONITORING CENTRE, 1992. Development of a national biodiversity index: a discussion paper. Cambridge: World Conservation Press.         [ Links ]