Rissoides barnardi ( Manning, 1975a )

Rissoides barnardi ( Manning, 1975a)

( Fig. 29 View FIGURE 29 )

Squilla desmarestii .— Barnard, 1950: 842, fig. 1a.

Meiosquilla desmarestii .— Manning, 1969a: 13.

Meiosquilla barnardi Manning, 1975a: 363–366 , fig. 1 [ type locality KwaZulu-Natal, South Africa]

Rissoides barnardi .— Manning & Lewinsohn, 1982: 353 (list).

Material examined. KZN: SAMC-A079426 1 ♂ (TL 37.5 mm), off Port Edward , 31°06’48.0”S 30°17’48.0”E, 8 Jul 1985, 120– 125 m, coll. G. C. Williams. GoogleMaps

Diagnosis. Cornea bilobed, set almost transversely on stalk. Ocular scales subtruncate. A1 peduncle subequal to, or slightly longer than, carapace. Rostral plate cordiform, apex rounded. Carapace smooth, lacking carinae except for short, reflected MG and posterior LT carinae. Raptorial claw dactylus with 5 teeth; propodus not distinctly tapering distally. Mandibular palp absent. 4 epipods present. TS5 lateral process with slender, oblique, laterally-projecting lobe, rounded laterally. TS6–8 with IM carina; TS6 and 7 lateral processes broadly rounded posteriorly; TS8 sternal keel slender with rounded apex. AS 1–5 with IM, LT and MG carina; AS 6 with distinct SM, IM and LT carinae. Abdominal carinae spined as follows: SM6 , IM 5–6 , LT 5–6, MG 4–5 . Telson stout with primary teeth bases swollen in male holotype; SM teeth with movable apices; IM and LT teeth slightly inwardly curved, apices sharp; denticles pointed, SM 6–10 , IM 10–15 , LT 1. Uropodal exopod proximal segment with 4–5 movable spines on outer margin, distalmost not extending beyond midway distal segment; protopod with inner margin crenulate .

Colour in alcohol. Completely faded. Cornea dark grey-brown.

Colour in life unknown.

Measurements. ♂ (n = 1) TL 37.5 mm. CI 386. A1 peduncle 1.11CL. A2 scale 0.51CL. PLDI 308. The present specimen represents the largest specimen recorded for the species.

Distribution and habitat. Only previously known from South African waters around Durban and now Port Edward, KZN. Associated with sandy and muddy substrata, depth 120–200 m (present study).

Remarks. Originally recorded from South Africa as the Mediterranean species Rissoides desmarestii Risso, 1816 ( Barnard 1950; Manning 1969a), R. barnardi is distinguished from R. desmarestii by the form of the propodus of the raptorial claw. In R. barnardi , the propodus does not become distinctly narrower distally, as it does for R. desmarestii . When compared to R. desmarestii , R. barnardi also appears to mature at a smaller size with Manning’s (1975a) male holotype exhibiting swellings at the bases of the telson’s primary teeth at TL 30 mm. This secondary sexual characteristic is known to only occur in R. desmarestii specimens at least 50 mm in total length ( Manning 1975a).

Although listed in the museum catalogue, the original holotypic and paratypic material of R. barnardi could not be located after searching the Iziko stomatopod collection. The present specimen was found among unidentified specimens in the museum collection and is the largest specimen recorded for the species. The male specimen agrees well with early accounts given by Barnard (1950) and Manning (1969a) and with the description of R. barnardi (as Meiosquilla barnardi ) by Manning (1975a).

DISCUSSION

The historical accumulation of known mantis shrimp species in South Africa is shown in Fig. 30 View FIGURE 30 and shows a very slow rate of accumulation over the first century with only five species reported. Barnard (1950) presented the first major review of the group and listed 13 species from South Africa based on his own observations as well as scattered records by early crustacean taxonomists ( Krauss 1843; Hansen 1895; Stebbing 1902, 1908, 1910, 1917). Further species additions by Manning (1969a, 1975a) and later Kensley & Buxton (1984) increased this total to 17, although not all records were correctly attributed or can be corroborated herein.Although limited to a single collection site, the most recent study by Ahyong (2005) added significantly to this total, bringing the known number of mantis shrimp species to 26, including one species described as new to science. The present study adds five more new species records and thus brings the known number of mantis shrimp species to 31. Apart from Barnard (1950), however, there has also been no holistic treatment of the group from South African waters and the present study is the first to provide a fully illustrated guide to the known stomatopod fauna of South Africa within a single account.

Stomatopods are notoriously difficult to collect due to their agile and defensive nature. They also occur in a wide range of habitat types from soft sediments to coral reefs, where they inhabit burrows and crevices. Therefore , a diverse range of sampling methods is needed to accurately estimate their regional diversity. Collection methods normally employed for this group include trawling, dredging, spearing, trapping, and pumping, as well as manual excavation of burrows and noosing of individuals (Ahyong et al. 2017). Historically , in South Africa, unspecified bulk sampling of Crustacea from the late 19 th century to the present has resulted in most of the squilloids and lysiosquilloids in museum collections. Meanwhile , the introduction of targeted sampling of stomatopods by hand and the utilization of SCUBA diving in the 1960s and 1970s allowed for the collection of the more inconspicuous gonodactyloids. For example, in 1976, R. Winterbottom of the J. L. B. Smith Institute undertook the targeted collection of coral reef related stomatopods from Sodwana Bay, which were later examined and reported on by Ahyong (2005) and now make up a large proportion of the gonodactyloid specimens at the Iziko South African Museum. The large number of new records found in that limited collection is indicative of the potential that such collection methods have in uncovering further new records form other sites .

Several records of species previously reported from within South African waters could not be authenticated from existing material, as the corresponding specimens are either lost from the Iziko collection or could not be traced. Four species previously documented from South Africa, Raoulserenea komaii Moosa, 1991 , Heterosquilloides insignis Kemp, 1911 , Oratosquilla mauritiana Kemp, 1913 and Quollastria gonypetes Kemp, 1911 , lacked material for examination herein. Of these, the record of R. komaii is the only contemporary species record ( Ahyong 2005) and while the specimen is extant in the Iziko collection, the sample was unfortunately damaged to an extent that it is no longer taxonomically useful. Although included herein, it is suggested that the historical records of two unverified species; O. mauritiana and Q. gonypetes , require sampling of new material for confirmation of species occurrence in South Africa.

Several species previously suspected to occur in South Africa are confirmed to be present. Barnard’s (1950) record of Gonodactylus demanii and all specimens in the Iziko collection previously identified as G. demanii are now considered to represent Gonodactylellus lanchesteri , which was first reported from South Africa by Ahyong (2005). First suggested by Barnard (1950) and reiterated by Manning (1969a), the presence of Odontodactylus scyllarus is confirmed for South Africa for the first time. However, four species listed to occur in Mozambican waters by Barnard (1950, 1958) and Manning (1969a) are still not yet recorded in South Africa. These are Alima neptuni Linnaeus, 1768 (as Squilla hierogliphica ), Gonodactylaceus falcatus Forskål, 1775 (as Gonodactylus glabrous ), Keppelius hystricotelson ( Barnard, 1958) (as Lysiosquilla hystricotelson ) and Lenisquilla lata Brooks, 1886 (as Squilloides lata ).

The South African Stomatopoda represented in the Iziko collection includes a widespread tropical component as well as species associated with temperate waters. Twenty-one species recorded here have comparatively widespread distributions in the Indo-West Pacific ( Bathysquilla crasisspinosa , Gonodactylellus lanchesteri , Gonodactylus botti , Gonodactylus chiragra , Gonodactylus smithii , Odontodactylus hansenii , O. scyllarus , Chorisquilla spinosissima , Raoulserenea komaii , R. ornata , R. oxyrhyncha , Lysiosquilla maculata , L. tredecimdentata , Heterosquilloides insignis , Clorida albolitura , C. latrellei , Harpiosquilla harpax , Kempella mikado , Miyakella holoschista , M. nepa and Quollastria gonypetes ), while most of the remaining species are restricted to the Western Indian Ocean ( Gonodactylellus crosnieri , Gonodactylolus paulus , Gonodactylus cf. chiragra , Mesacturoides fimbriatus , Natosquilla investigatoris and Oratosquilla mauritiania ). In general, the species that occur in the subtropical parts of the country also occur in Mozambique and Madagascar, a trend visible across other taxonomic groups. Fifteen stomatopod species from South Africa are shared with Madagascar, while only 10 are shared with Mozambique, of which G. botti and the undescribed species of Clorida were only recently recorded for the first time in Mozambique (Brokensha et al. 2023). This is probably indicative of the sparse collecting efforts undertaken in Mozambique.

Of the South African Stomatopoda , only two species; Lysiosquilla capensis and Pterygosquilla capensis , occur in cold-temperate waters. The essentially temperate water South African stomatopod species are the endemics L. capensis and P. capensis . Last studied by Manning (1969a), the large collection of P. capensis specimens at Iziko South African Museum allowed for a more detailed investigation of the species. The high variation found in the diagnostic characters of both the species and the genus revealed that Pterygosquilla needs a global revision beyond the scope of this paper. It is possible that the observed variation is due to the presence of a yet unrecognised species and that P. capensis is part of a species complex. The species is frequently caught in the demersal fishery survey trawls off the south and west coasts and any future study should make use of this accessibly to fresh material for genetic analysis. The present study found the diagnostic characters used by Manning (1969a) and Ahyong (2012) to be stable, but to properly consider the findings herein, P. gracilipes from Chile should be investigated. Pterygosquilla capensis is the most abundant stomatopod species on the west coast and is considered the only species known to occupy the continental waters of South Africa south of the Namibian border to St. Helena Bay. However, the present study extends the species range further east, to within the mixing zone on the south coast.

Meanwhile, most tropical squilloid genera widespread in the Indo-West Pacific recorded here were found to occur along a short stretch of the east coast between Durban and St. Lucia Bay ( Clorida sp. , C. latrellei , Harpiosquilla harpax , Kempella mikado , Miyakella holoschista , Quollastria gonypetes , and Rissoides barnardi ) with M. nepa being the only squillid species to range south from KwaZulu-Natal into the Eastern Cape. This area holds South Africa’s only commercial crustacean trawl fishery ( Sink et al. 2019), likely explaining the high abundance of species records and museum specimens collected from this area. Two lysiosquillids ( Lysiosquilla maculata and L. tredecimdentata ) and a bathysquilloid ( Bathysquilla crasisspinosa ) also share this stretch of coastline. These mantis shrimp taxa are known to favour rather specific habitat types (Ahyong et al. 2017) with lysiosquillids and squillids generally inhabiting level mud and sand habitats into which they burrow. Historically, earlier sampling efforts would target certain areas and habitats, especially False Bay and Table Bay in the Western Cape, Algoa Bay in the Eastern Cape, and Durban Bay in KwaZulu-Natal ( Griffiths 1999), nearly exclusively using trawlers on soft substrates, but leaving those stomatopod species with deep burrows and/or reef-associated lifestyles largely unsampled and overlooked.As gonodactyloids are generally associated with coral or rocky reef systems, such as the ones just south of Mozambique, the limited historical sampling resulted in them being vastly underrepresented until the rise of SCUBA diving and targeted collection in under-sampled habitats in the 1970s.

The gonodactyloids recorded herein are restricted to the stretch of coastline influenced by warm-temperate to subtropical waters from just south of Durban to the Mozambican border. These species are typically associated with coral reef and rubble habitats and include four of the five new species records for South Africa ( Gonodactylellus crosnieri , Gonodactylus cf. chiragra , Odontodactylus hansenii and O. scyllarus ). The majority of gonodactylids represented herein are collected from the subtropical reef systems in the northernmost region of the east coast, just south of Mozambique. Most of these records are reported on by Ahyong (2005) for stomatopods from Sodwana Bay and its immediate surroundings. Other coral associated stomatopods from this area are the protosquillid Chorisquilla spinosissima , the takuid Mesacturoides fimbriatus and the four pseudosquillids. Meanwhile, the southernmost specimen of Gonodactylus chiragra corresponds to localities associated with the reef systems of the KwaZulu-Natal Bight off St. Lucia. Excluding G. chiragra , species of Odontodactylus and Gonodactylus are exclusively collected from north of Cape Vidal in the warm coastal waters known for its thriving soft coral communities ( Sink et al. 2019).

Two species from the east coast are currently considered South African endemics and both are restricted to the warm-temperate and subtropical waters of KwaZulu-Natal. Pseudosquillisma kensleyi inhabits the subtropical coral reefs in the far north, while Rissoides barnardi is rather associated with the deeper habitats of sand or mud substrates between Port Edward and Durban. However, both P. kensleyi and R. barnardi are reported herein from singular preserved specimens, leaving their true distribution poorly known.

Overall, the taxonomic accounts of the 31 stomatopod species presented in this study aim to collate current knowledge of the South African stomatopod fauna and to greatly simplify accurate identification of known regional species. However, despite the coverage of the present work, numerous species remain poorly sampled, leaving many unanswered questions of distribution and taxonomy to be investigated. The mobile and inconspicuous nature of the group also suggests that their abundance and diversity is still greatly underrepresented in the existing collections. Habitat-specific sampling targeting of mantis shrimps would undoubtfully find many new species records and perhaps new species to science in the region.

ACKNOWLEDGEMENTS

This study is based on research supported by the National Research Foundation of South Africa (Grant Numbers 128324, and 138888) through the Integrated Biodiversity Information Programme and Extension Support Master’s Scholarship at the University of Cape Town. Our sincerest thanks to the staff at the Iziko South African Museum, with special acknowledgement to Wayne Florence, Albé Bosman, and Romano Adams for access to and assistance with specimens.

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