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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 338

On the Zoogeography of Southern African Decapod Crustacea, with a Distributional Checklist of the Species

Brian Kensley

SMITHSONIAN INSTITUTION PRESS

City of Washington

1981

ABSTRACT

Kensley, Brian. On the Zoogeography of Southern African Decapod Crustacea, with a Distributional Checklist of the Species. Smithsonian Contributions to Zoology, number 338, 64 pages, 4 figures, 4 tables, 1981.—Decapod crustacean research in southern Africa is reviewed. The terrestrial and freshwater, pelagic, and benthic decapods are discussed separately. The Atlantic, Indo-Pacific, and endemic components of the benthic fauna are discussed, related to neighboring islands, seamounts, and shoals, and compared with other southern hemisphere faunas. A checklist for about 700 species, with vertical and geographical distribution information, is provided.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Kensley, Brian Frederick.

On the zoogeography of Southern African decapod crustacea.

(Smithsonian contributions to zoology ; no. 338)

Bibliography: p.

1. Decapoda (Crustacea)—Africa, Southern—Geographical distribution. 2. Crustacea—Geographical distribution. 3. Crustacea—Africa, Southern—Geographical distribution. I. Title. II. Series.

QL1.S54 no. 338 [QL444.M33] 591s 81-607972 [595.3′840968] AACR2

On the Zoogeography of Southern African Decapod Crustacea, with a Distributional Checklist of the Species

Brian Kensley

____________

Brian Kensley, Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

Introduction

Since Barnard's (1950) invaluable monographic treatment, no single comprehensive work on southern African Decapoda has appeared. The decapod fauna has, however, received considerable attention, and many new records have been noted. While much taxonomic work remains to be done, and while many southern African areas have been poorly collected, it was nevertheless thought useful to review the group as a whole, drawing such zoogeographic conclusions as are possible from the available data. To this end, a species list has been compiled to give some idea, albeit incomplete, of the total fauna known to date and to enable comparisons with other areas.

ACKNOWLEDGMENTS.—My sincere thanks are due to the trustees and director and staff of the South African Museum, Cape Town, for assistance and hospitality during my visits in 1978 and 1979; to the Zoology Department of the University of Cape Town for the use of collections and data; to Dr. F. A. Chace, Jr., and Dr. R. B. Manning of the Smithsonian Institution, and Dr. I. Pérez-Farfante and Dr. A. B. Williams of the Systematics Laboratory, National Marine Fisheries Service, National Oceanic and Atmospheric Administration, for reading the manuscript and for providing many useful comments and criticisms, and for additional data; and to Mrs. Cynthia Brown for assistance with maps and figures.

Brief Historical Review of Decapod Collecting and Research in Southern Africa

Probably the earliest serious collector of southern African decapods was Sir Andrew Smith, founder of the South African Museum, who, on his return to England in 1837, gave his collection of crabs to W. S. MacLeay. This resulted in the earliest report on southern African decapods, “The Annulosa of South Africa” in Smith's Zoology of South Africa of 1838. Several of MacLeay's types are now in the Australian Museum, Sydney.

Dr. Ferdinand Krauss spent the years 1838–1840 collecting around the South African coast and published Die Südafrikanischen Crustaceen in 1843. Several expedition vessels subsequently collected in southern African waters, including the Challenger, Gazelle, Valdivia, and Gauss, as well as the United States North Pacific Exploring Expedition. Ortmann (1896) based his zoogeographiic discussions on decapods and included this accumulated knowledge in his pioneering work.

The next important addition to knowledge came from the work of the Cape Colony research vessel Pieter Faure from 1897–1907. These collections were first reported on by the Rev. T. R. R. Stebbing, and later by K. H. Barnard, both in the Marine Investigations of South Africa, and in the Annals of the South African Museum. The first checklist of South African Crustacea was produced by Stebbing (1910) in the Annals of the South African Museum. The Union government vessel Pickle made several collections during the 1920s, the material being examined by Stebbing, Calman, and Barnard. The culmination of all this work was K. H. Barnard's Descriptive Catalogue of South African Decapod Crustacea published in 1950, in which about 500 species were reviewed.

The present list contains about 700 species. The extra species and records have been added by several workers. Barnard (1954, 1955, 1958) included several new records as a result of the collecting done by the universities of Cape Town and the Witwatersrand. Several contributions have since been made to the southern African decapod literature, including Forest (1954) on hermit crabs, Hayashi and Miyake (1968) on stylodactylids, Griffin (1968) on majid crabs, Grindley (1961) on Natal crabs, Berry (1969a,b 1971, 1979) on palinurans and nephropids, Hayashi (1975) on processids, de Freitas (1979) on penaeids, and Kensley (1968–1980) on a variety of groups.

Certain areas around the coast, for one reason or another, have received more attention than others, resulting in gaps in distributional knowledge. Decapods from Lüderitz, South West Africa, have been collected by the German South Pole Expedition, the University of Cape Town, and the South African Museum. Saldanha Bay, and more particularly Langebaan Lagoon, has been extremely well sampled because of the annual student camps and surveys of the Zoology Department of the University of Cape Town. Most estuaries have been sampled by the same institution, while False Bay, Cape Province, because of its easy accessibility and position, has been well sampled both intertidally and from greater depths by the U.S. Exploring Expedition, the Pieter Faure, the John D. Gilchrist, and the Thomas B. Davie, the latter two being research vessels of the University of Cape Town. A comprehensive checklist of the fauna of False Bay resulted from this work (Day, Field, and Penrith 1970). Delagoa Bay and Inhaca Island, Mozambique, received considerable attention following K. H. Barnard's visit in 1914. Up to the early 1970s the University of the Witwatersrand conducted annual visits to the island's research station and documented the fauna and flora of the region (MacNae and Kalk, 1958). The South African Museum collected from Inhaca Island to Vilanculos and Magaruque Island (22°01′S, 35°19′E) in the north during 1971 and 1973.

Until recently the continental shelf beyond the 200 m line had been poorly sampled, and only in scattered areas such as Lambert's Bay, Saldanha Bay, Table Bay, False Bay, and the Agulhas Bank in the Still Bay area. Up to 1975, the most comprehensive but still very inadequate report on shelf/slope decapods dealt with a very limited area off the Cape Peninsula, which was the result of the South African Museum-Division of Sea Fisheries deep trawling of the R.V. Africana II (Kensley, 1968). In 1975, the South African Museum initiated a five-year program of sampling the deep benthic and pelagic fauna off the east coast from the Mozambique border to Transkei, with the help of the C.S.I.R. R.V. Meiring Naude. The resulting 256 stations form the most comprehensive if still inadequate collection of decapods from deep water in southern Africa (Kensley, 1977a,b, 1980a).

There are areas which have enjoyed little or no sampling, and which accordingly weaken any zoogeographic conclusions. These include much of the continental shelf, especially the Agulhas Bank, and the entire West Coast; also the Transkei-Pondoland-Zululand shallow waters (with the exception of the Durban area). This latter omission is unfortunate, as the area includes the transitional zone from the Semitropical East Coast Province to the Warm Temperate South Coast Province.

Geographical Limits of the Present Study

The area covered by the present work has not been too rigidly defined so as to allow inclusion of as many records as possible. The northern limit on the west coast is taken as the mouth of the Kunene River, the brachyurans of the West African region from southern Angola northwards having been dealt with by Monod (1956) and Manning and Holthuis (1981). A list of intertidal decapods is included in a checklist of shore animals from Moçâmedes, southern Angola (Kensley and Penrith, 1973). On the east coast, Vilanculos in Mozambique is taken as the northern limit. (See Figure 1.) Barnard (1950) adopted the 15° latitude as his northern limit on both the east and west coasts in his monographic work on the decapods, as well as in his earlier work on fish (1925). This corresponds with Moçâmedes on the west and Mozambique Island on the east. With the exception of Kalk (1959), there are almost no published records of decapods north of Vilanculos. In his work on the Mollusca, Barnard (1974) stated that the 15° latitude seemed too wide an area, and accordingly placed his limits at the Tropic of Capricorn, i.e., Walvis Bay on the west and Inhambane on the east.

As to distance out to sea, almost no limit has been placed in this work. Apart from the 256 Meiring Naude stations mentioned earlier, little work has been done beyond the 200 m line. A very few Division of Sea Fisheries stations extend from about 5° to 45° east and to 45° south. These have been included in this survey, as well as the isolated stations on the seamounts Tripp and Vema, and Walter's Shoal. For comparative purposes, the decapod faunas of St. Helena Island, Ascension, and the Tristan da Cunha group, Marion and Prince Edward and Gough islands, have also been considered.

FIGURE 1.—Map of southern Africa showing major collecting localities.

Composition and Zoogeography of the Southern African Decapod Fauna

In an analysis of any large and heterogeneous group of organisms, a knowledge of the modes of life, life histories, and habitats is essential if nonsensical interpretations are to be avoided. Thus in the following discussion, the decapods are dealt with in several sections. The terrestrial and freshwater forms are only briefly mentioned. The marine forms are dealt with in three separate sections, viz., shallow benthic (intertidal to 200 m), deep benthic (beyond the 200 m line), and pelagic forms.

TERRESTRIAL AND FRESHWATER DECAPODA.—Two macruran, one anomuran, and three brachyuran families have representatives either on land or in freshwater in southern Africa. Freshwater shrimps are found amongst the Atyidae and Palaemonidae; the Coenobitidae contain the only terrestrial hermit crabs, while the Gecarcinidae and Grapsidae include the terrestrial crabs. The freshwater crabs all belong to the Potamonautidae.

Potamonautidae: With the exception of Gecarcinautes brincki Bott (1960), all the freshwater crabs of southern Africa belong to the genus Potamonautes, which is exclusively African. Eight species occur south of the Zambesi River. It is postulated (Bott, 1955) that the Potamonautidae originated from marine ancestors of the Tethys Sea in the Northern Hemisphere. Palaeontological evidence seems to indicate this origin somewhere near the end of the Cretaceous or the beginning of the Tertiary. One section of the ancestors left the sea and migrated onto land, while the ancestors of the Potamonautes group entered fresh water and migrated southward down the African continent via the major river systems and lakes of the Rift Valley complex. This southward migration, along with adaptive radiation, led to the formation of species or species-groups, each characteristic of a particular river or river-system. Where a river-system is relatively isolated, well-defined species such as P. bayonianus (Brito-Capello) of the Kunene River have arisen. Where several river-systems interlink or where a large area is drained by several small closely situated rivers, the definition of species becomes blurred. Thus the Cape Province is characterized by P. perlatus (H. Milne-Edwards), and Natal by P. sidneyi (Rathbun), but a wide range of overlap occurs from which transitional forms showing features of both species have been recorded. Similarly, where P. perlatus (H. Milne-Edwards) overlaps with P. warreni Calman in the Orange Free State and Transvaal, transitional forms are found. These forms lead one to suspect the subgeneric divisions proposed by Bott (1955).

Gecarcinautes brincki Bott, recorded from the mountain streams of the southwestern Cape Province, has its closest relatives in the rivers of Madagascar.

Gecarcinidae: There are not many records of land crabs from southern Africa, the gecarcinids being tropical in distribution and essentially nocturnal. Cardisoma carnifex (Herbst) has been recorded northwards from Durban, while C. armatum Herklots occurs on the west African coast from Moçâmedes northwards.

Grapsidae: Geograpsus stormi de Man, being a tropical Indo-Pacific species, has also been recorded northwards from Durban on the east coast.

Coenobitidae: Of the land hermit crabs, Coenobita rugosus H. Milne-Edwards is known from the east African coast north of Natal and the Indo-Pacific region, while C. cavipes Stimpson appears to be restricted to the Indian Ocean.

Atyidae: Of the more than 20 genera of the Atyidae, only Caridina occurs in southern Africa, where it is represented by four species, some of which are of doubtful validity. The uncertainty of the taxonomic position of the species is due to the variability of the forms and the paucity of collecting. Material, usually only a few specimens from isolated localities, is all that is available at present. Intensive collecting over a wide area is needed before reliable specific definitions may be derived.

Caridina typus H. Milne-Edwards, a species supposedly found on several Indian Ocean islands, the western Pacific, and Queensland, Australia, has been recorded from several rivers in Natal, including the Umhloti, Umgeni, Illovo, Umbilo, and Uvongo. Caridina nilotica (Roux) varies with regard to egg size, and this has given rise to the description of varieties such as C. nilotica var. paucipara Weber (1897) and C. nilotica var. natalensis de Man (1908a). Habitat also plays a role in the degree of variation, as lacustrine and fluviatile forms of this species have been distinguished. The species is widespread, occurring throughout East Africa, Egypt, the Indian Ocean area, as well as China and Australia. Caridina africana Kingsley, recorded from the Amamzimtoti River in Natal, and Zululand, may prove to be a form of C. nilotica. Caridina indistincta Calman was originally described from Australia, but specimens from the Congo and Zambesi rivers have been assigned to it.

It would seem that the southern African Caridina species, like the freshwater crabs, are either migrants or derived from migrants from the north.

Palaemonidae: The family Palaemonidae has freshwater, estuarine, and marine representatives in southern Africa. Again, due to lack of collecting, several of the freshwater forms are in uncertain taxonomic position.

Palaemon (Palaemon) capensis de Man, the Cape River prawn, is a true freshwater form, having been recorded from several rivers, including the Gamtoos, Duivenhoks, Buffeljachts, Palmiet, Zonderend, Baakens, and Breë. Its range would thus seem to be between the Palmiet River near Hermanus and the Baakens River near Port Elizabeth. The species has not been recorded from any of the west coast rivers.

The genus Macrobrachium contains the rest of the freshwater prawns in southern Africa and is represented by about seven species. Macrobrachium vollenhoveni (Herklots), recorded from the Kunene River, may be regarded as a true West African form, being known from northern Angola, Liberia, and the Cape Verde Islands. The Kunene form may well be on the way to developing a separate identity, showing more slender pereopods than the northern representatives.

Macrobrachium lepidactylus (Hilgendorf) was originally described from northern Mozambique, and has since been found in Tanzania and Madagascar. In southern Africa it has spread down the river systems of the east coast, and is known from southern Mozambique, Zululand, Natal, East London, and the eastern Transvaal. Macrobrachium equidens (Dana) inhabits the lower reaches and estuaries of rivers in Natal and southern Mozambique.

Macrobrachium rude (Heller), M. petersi (Hilgendorf), M. scabriculum (Heller), and M. idella (Hilgendorf) have all been recorded from Natal and southern Mozambique. All are typically tropical east African and Indian forms. As the southern African material is often immature, and as few specimens are collected from any single locality, some of these identifications are still open to doubt.

PELAGIC NATANTIA.—Before any discussion of the southern African Natantia can be attempted, some reservations regarding the data on meso- and bathypelagic species must be made.

The single overriding factor that prevents any firm conclusions from being drawn regarding vertical distribution is that opening/closing nets have not been used for macroplanktonic sampling. The earlier collections, including those of the Pieter Faure and the Pickle, were made with an assortment of dredges, trawls, and nets, while the more recent collections, such as those made by the Africana II off Cape Point, the midwater survey of Grindley and Penrith (1965) on the SS Natal, and the South African Museum's Meiring Naude survey, did not use closing nets (although the latter did use temperature/depth recording devices). The non-selective sampling is well illustrated by the deep-sea collection of the Africana II off Cape Point. Although a beam trawl was used to sample the benthic fauna, the samples included such genera as Sergestes, Gennadas, and pelagic Acanthephyra, as well as the mysid Gnathophausia, the jellyfish Periphylla, and a large number of pelagic fish. In such a case, it may be said that these organisms occur in the waters under discussion, but correlation with more precisely defined water-masses is not possible.

The effect of vertical migration in water-mass correlation will further weaken the available data. Some species may rise from one water-mass to another, and, depending on time of capture, may be associated with either water-mass. Foxton (1972) has shown that in the North Atlantic, mesopelagic species of the genus Acanthephyra tend to execute considerable vertical migrations, while the deeper-living bathypelagic species apparently do not migrate.

The term “pelagic,” when used in relation to a species, is here understood to imply the inhabiting of the main water body of the sea, living neither in the upper 200 meters (epipelagic) nor on the sea bed (benthic). “Mesopelagic” indicates living in the depth range 200–500 meters; “bathypelagic” indicates living in the depths beyond 500 meters (Briggs, 1974). In this study, some members of the caridean families Oplophoridae and Pasiphaeidae and the penaeidean families Aristeidae, Sergestidae, and Penaeidae are regarded as pelagic.

Kensley (1974c) attempted to demonstrate statistically the presence of species-groups significantly associated with specific water-masses, using data available up to 1974. An inherent weakness of this analysis was the diverse and non-selective collecting methods used, which necessitated a subjective decision to be made for each sample as to water-mass of origin. It was nevertheless thought useful to carry out the analysis for any information it might yield. The McConnaughey coefficient (McConnaughey, 1965), which reveals homogeneous groups within heterogeneous systems and employs as variables the occurrences of species A, occurrences of species B, and co-occurrences of species A and B, along with a generalised sorting strategy (Lance and Williams, 1966), was used to generate a dendrogram. Interpretation of the dendrogram could not be given any great significance; rather it was used as the starting point for discussion. The most interesting facts to emerge from this analysis were that the Aristaeomorpha foliacea was linked to South Indian Central Water and that a much larger group of 20 species, containing several species of Sergestes, Sergia, and Gennadas, and the common Acanthephyra quadrispinosa Kemp and Systellaspis debilis (A. Milne-Edwards), was loosely connected to South Atlantic Central Water. Beyond this very broad statement, the analysis gave some (admittedly dubious) weight to subjective conclusions arrived at mainly by direct observation. For example, the Aristeidae, described by Burkenroad (1936) as containing mainly oceanic forms, contains the genus Gennadas, which the analysis showed to be truly oceanic, being found only in the South Atlantic Central Water, as is the related Bentheogennema intermedia (Bate).

Chace (1940) noted that several of the Oplophoridae are truly bathypelagic (sensu latu) oceanic forms. The analysis bore this out, the genera Oplophorus, Acanthephyra, Systellaspis, and Hymenodora being found only in South Atlantic Central Water or in Antarctic Intermediate Water. Foxton (1972) showed that Acanthephyra pelagica (Risso) and A. purpurea A. Milne-Edwards were always associated with North Atlantic Central Water, whereas A. sexspinosa Kemp and A. acanthitelsonis Bate were always associated with South Atlantic Central Water. Also, A. prionota Foxton, A. curtirostris Wood-Mason, and A. stylorostrata (Bate) were species living in deeper water than the four aforementioned species. Foxton was able to state these facts with some certainty, as his specimens were caught with opening and closing nets. It is possible that a similar system prevails in the south Atlantic and the southwest Indian Ocean.

The following remarks may be made regarding the southern African oplophorids. Three species appear to be rare and are found in water deeper than 2000 m; these are A. corallina (A. Milne Edwards), A. brevirostris Smith, and A. gracilipes Chace taken off Cape Point in 2480–3000 meters. Acanthephyra armata A. Milne-Edwards (described from the Lesser Antilles) and A. indica Balss are known from the Indian Ocean, and have been recorded off the east coast to 850 meters. Six species appear to have a wide depth range, being found from 250–2000 meters, viz., A. curtirostris Wood-Mason, A. eximia Smith, A. pelagica (Risso), A. prionota Foxton, A. quadrispinosa Kemp, and A. stylorostrata (Bate). Acanthephyra quadrispinosa Kemp and A. pelagica (Risso), members of the purpurea species complex, are the two common species off South Africa. The former has been recorded from the surface to 3800 meters, while the latter has been recorded from 250–3800 meters, but both appear to be abundant in the 500–600 meter zone.

Amongst the Natantia from deeper waters, many genera are almost cosmopolitan in distribution, with the same species occurring in several water-masses having similar properties. The region off the Cape is sometimes regarded as a corridor for the Natantia. Burkenroad (1936) showed that many so-called Indo-Pacific species found off the Cape also occurred in the Atlantic off the Bahamas and the northeast coast of the United States. This connection via the Cape was used to explain the dissimilarity of the oceanic Natantia off the east and west coasts of the U.S.A.

Turning to the zoogeographic relationships of the Natantia under discussion, these may be summarized as follows: of the approximately 67 species regarded as truly pelagic, 27 (40%) have been recorded from both the Atlantic and Indo-Pacific; 17 (25%) are Atlantic forms; and 18 (27%) are Indo-Pacific forms. More interestingly, and in line with the idea of the southern African oceans being a corridor area, of the 17 purely Mediterranean/Atlantic species, 13 (76%) have been recorded east of Cape Agulhas, while of the 18 Indo-Pacific forms, 10 (55%) have been recorded west of Cape Point. The presence of these latter 10 species may be explained partially by the eddying of warm pockets of Agulhas water on the west coast (Welsh and Visser, 1970). Whether the foregoing figures indicate genuine penetration into neighbouring areas or merely a lack of knowledge concerning the distribution of pelagic species is difficult to assess.

BENTHIC DECAPODA.—Consideration of species having a depth distribution below 200 m, many of which are single records, has been omitted from the following discussion of zoogeographic relationships.

The overall composition of the southern African decapods is given in Table 1. The zoogeographic affinities of the benthic decapods from less than 200 m are given in Table 2, while the distribution of the major components of this group around the southern African coastline is illustrated in Figure 3.

The Indo-Pacific component constitutes the major section of this part of the decapod fauna. Predictably, this component shows a marked increase from west to east along the coastline, with 73% occurring at Durban and 93% at Maputo/Inhaca Island. There is a dramatic cutoff between Durban and East London, which would indicate the transition from a Subtropical East Coast Province to a Warm Temperate South Coast Province. This is obviously a reflection of the change in the temperature regime of the seas of the area. The Indo-Pacific, and more especially the Indo-West Pacific, is a predominantly tropical/subtropical marine region, with circulation in the northern part of the Indian Ocean by the North Equatorial Current and the Counter Current and the South-West and North-East Monsoon Drift, and in the southern part by the South Equatorial Current, communicating with the western Pacific through the various straits of the East Indies and northern Australia. (See Figure 2.) The westward-flowing North and South Equatorial currents could bring the larval planktonic or pelagic forms towards the East African coast, both north and south of Madagascar. The part of the North Equatorial Current that is diverted southward along the African coast passes down the Mozambique Channel as the Mozambique Current, to be joined by the westerly component of the South Equatorial Current, thereby forming the Agulhas Current. Depending on the width and direction of the continental shelf, the Agulhas Current may be either close inshore, as between Delagoa Bay and St. Lucia and between Durban and East London, or further offshore, as between St. Lucia and Durban and south of East London. Where this warm southerly flowing body of water is close inshore, the intertidal and shallow areas have a good chance of receiving larval forms of tropical and subtropical species and of allowing them to become established. This would account in part for the high numbers of species recorded from Durban and the Delagoa Bay areas. Where the Agulhas Current, by virtue of the width of the continental shelf, is forced offshore, a countercurrent of cold water flowing in a northeasterly direction develops, which may even reach the Limpopo River mouth. This countercurrent could to some extent explain the relative paucity of species in the area between Delagoa Bay and Inhambane (although poor collecting in the area cannot be discounted), and is almost certainly a limiting factor to the southward range extension of subtropical species. Some subtropical species have managed to colonize southern Natal and the Eastern Cape coastal areas in spite of the cold countercurrent (prawns of the genus Penaeus reach the Knysna estuary). This may be explained by the presence of a southward-flowing inshore current (Harris, 1961) along the Natal coast, which is especially noticeable in summer and thought to be caused by strong northeasterly sea breezes. On the other hand, the cold inshore countercurrent, especially noticeable between the Agulhas Bank and East London, may assist a few cold-water (i.e., west coast) species to become established east of False Bay.

TABLE 1.—Composition of the southern African decapod crustacean fauna

TABLE 2.—Zoogeographic components of the southern African benthic decapod crustacean fauna from less than 200 m

Component
Species
% of total

Indo-pacific

345
65.7

Atlantic/Mediterranean

  30
  5.7

Endemic

103
19.6

Other (Austral, widespread, uncertain)

  47
  8.9

Total             
525

FIGURE 2.—Major ocean currents around southern Africa. (Stippled area = Agulhas Bank.)

Yet another factor controlling the presence or absence of several decapod species is the presence or absence of specialized habitats. The three most notable of these are coral reefs, mangroves, and shallow-water seagrass beds. The former two are fully, and the latter to some extent, dependent on warm water for their existence. Thus, coral reefs are not found south of southern Mozambique, and those decapods always associated with corals, including several xanthid crabs and alpheid shrimps, do not occur south of northern Natal. Similarly, mangroves that occur in estuaries, embayments, or sheltered areas are noted for their associated crab fauna, especially species of Uca and Sesarma. As one moves down the coast from Mozambique to Natal and the Eastern Cape, under the combined lack of warm water and the influence of the cold countercurrent, the trees which form the basis of the mangal decrease both in number and diversity. MacNae (1968) showed that many of the specialized crabs that inhabit mangroves are dependent on a variety of substrates found chiefly in the mangal.

Most shallow-water seagrass beds, along with their associated fauna, are controlled by the presence of warm water. Mud flats are colonized by plants such as Cymodocea, Halodule, Halophila, Thalassia, and Zostera, in the shelter of which decapod genera such as Diogenes, Hippolyte, Tozeuma, Gnathophyllum, Periclimenes, and Periclimenaeus flourish. Only Zostera and Halophila reach southern Natal and the Cape Province, and the associated fauna thus decreases.

The Atlantic/Mediterranean component, forming only about 6% of the fauna, shows a peak around False Bay and Port Elizabeth, tapering off to the east, but with still an almost 10% presence at Maputo. The low point on the west coast at Lüderitz may be explained by sea temperatures. The area of maximum upwelling of cold Antarctic Intermediate water (9°–12° C) is at Lüderitz (Stander, 1964), while northwards in the area of the Kunene River mouth, and southwards around the Agulhas Bank area, water closer in temperature to that off Angola is found. The effect of the cold Benguela water may be adjudged by the fact that the only corals occurring off West Africa are north of the equator in the Gulf of Guinea, which is swept by the warm Guinea Current. Upwelling of cold nutrient-rich water on the west coast also contributes to the characteristic faunal and floral pattern, viz., few species but large numbers of individuals.

Whether the Atlantic species found east of False Bay are in genetic contact with the rest of the Atlantic populations, or whether these represent relict populations from warmer Pleistocene times needs to be investigated for the individual species. The slightly higher percentage (22.6%) of Atlantic forms in northern South West Africa reflects the presence of a few West African species. Bearing in mind the north-flowing direction of the Benguela Current System, it is surprising that so few Atlantic species manage to migrate and populate the southern west coast and the south-east coast. The actual method of migration from north to south is probably a step-by-step and relatively slow occupation of the continental shelf area. It is possible that invasion, especially of pelagic larval forms, is assisted by movement of Atlantic water around and into the southwest Indian Ocean. This has been demonstrated to occur in relatively deep water (le Pichón, 1960; Visser and van Niekerk, 1965; Shannon, 1966), and would account for the presence of true Atlantic pelagic forms in the southwest Indian Ocean. A similar movement of water, but in the opposite direction, would account for the presence of Indian Ocean species in the Saldanha Bay area. Shannon (1966) demonstrated that movement of Agulhas water around Cape Point does occur. Only species tolerant of the low temperatures of the west coast could survive such migration. Eddying of Agulhas water into the South Atlantic (Welsh and Visser, 1970) may also account for east coast species on the west coast.

A few species of the southern African decapod fauna appear to have a southern oceanic or austral distribution, viz., Plagusia chabrus (Linnaeus), Ovalipes punctatus (de Haan), Pilumnoides perlatus (Pöppig), and Lithodes murrayi Henderson.

Pilumnoides perlatus (Pöppig), first recorded from Chile, is also known from Panama, Queensland, Australia, Ireland, and Plymouth, England, and is known in southern Africa from northern South West Africa to False Bay. These localities would seem to suggest that the species has a cold-water preference. The unusual records from Britain and Panama may be due to transportation on ships' keels, the species having been recorded from such sites on several occasions. The southern distribution of this crab together with the other three species can be explained by the influence of the West Wind Drift. This current, flowing from west to east, communicates with the west coasts of South America, South Africa, and Australia by means of the Peru, Benguela, and Western Australian Currents, respectively, and also bathes the islands of Tristan da Cunha, Gough, Marion and Prince Edward, St. Paul and Amsterdam, and New Zealand. Pilumnoides perlatus has a larval life span of 43 to 56 days (Fagetti and Campodonica, 1973), sufficient for considerable transport by ocean currents.

Ovalipes punctatus (de Haan) is a swimming crab, while Plagusia, the rock crab, has been found floating on objects far out to sea. There is thus no reason for not invoking the West Wind Drift to explain the southern distribution of these species.

Lithodes murrayi Henderson, known from Macquarie Island, New Zealand, the Crozet Islands, and Prince Edward Islan, has been recorded from deep water of Natal and Lüderitz, which distribution is more difficult to explain.

ENDEMIC DECAPODA.—Any discussion on endemism obviously depends on the state of knowledge of the geographic and depth distribution of the species in question. Because of its accessibility, the intertidal fauna of any area almost always will be relatively well known, and a species can be designated “endemic” with some degree of confidence. When the infratidal region is considered, however, it soon becomes apparent that gaps of knowledge exist, as few infratidal areas of any extent have been thoroughly sampled. Species from the infratidal thus seldom can be designated “endemic” with any confidence. For the purposes of the present discussion, an endemic species is considered to be confined to the geographic area previously defined and limited to the depth range 0- ± 200 m, in agreement with Smith (1970). The species of the lower part of this range are called endemic with reservation, and only after consideration of all depth records.

Only seven genera are endemic, and of these, Projasus is known from off the east coast from depths of about 600 m. Macropetasma, Eudromidia, and Exodromidia have been recorded from the west coast, but are not confined to this area. The four crab genera all have been recorded on the east coast. Of the fifty-three endemic brachyuran species, twenty-two have been taken in the intertidal zone.

The family Dromiidae is worthy of note, fourteen of the twenty-eight species being endemic; another four species, although from depths of more than 200 m, have not been recorded elsewhere, but may not be endemic. Eleven species are found in the Saldanha Bay to Agulhas Bank area. It would seem that, although basically of Indo-Pacific affinity, these species tend to be stenothermic and prefer the relatively cooler water of the south coast, i.e., the area of mixing of cold west coast and warmer Agulhas water.

Of the 105 species of anomurans, twenty-seven are endemic, while another thirteen from more than 200 m are not recorded elsewhere. Fourteen species are intertidal dwellers. Seven species have been recorded west of Cape Point. A further four species recorded from deep water (+500 m) off the west coast are known only from the area between Saldanha Bay and Cape Point.

Of the thirty-two species of palinurans, one is endemic, and six are recorded from more than 200 m, but not elsewhere. Although Homarus capensis (Herbst) has been recorded four times between the Cape of Good Hope and Port Elizabeth, no depth information is available (see Wolff, 1978). Polycheles demani Stebbing and Willemoesia bonaspei Kensley, both from very deep water, have been recorded only from off the Cape Peninsula, the latter species only once.

Twenty-five (12%) of the 208 species of Natantia are endemics. Of these, four are intertidal dwellers, viz., Periclimenes delagoae Barnard, Periclimenaeus uropodialis Barnard, Alpheus edwardsii (Audouin), and Hippolyte kraussiana (Stimpson). Ten species have been recorded from the west coast, and of these, Leontocaris paulsoni Stebbing, Lebbeus saldanhae (Barnard), and Plesiopenaeus nitidus Barnard have not been recorded east of Cape Point. The remaining species from the west coast, Haliporoides triarthrus Stebbing, Metacrangon jacqueti bellmarleyi (Stebbing), Solenocera africana Stebbing, Macropetasma africana (Blass), and Ogyrides saldanhae Barnard have all been recorded eastward to Natal.

Further analysis of the endemic component casts additional light on the faunal affinities of the area. Of the 103 endemic species, thirty-three are known from the west coast; twenty of these are distributed eastward beyond Cape Agulhas for varying distances, some even as far as Mozambique. Most of these species are regarded as being part of the endemic group of the area between the Transkei and Cape Point that has penetrated to the west coast, especially to the sheltered Saldanha Bay area. Ogyrides saldanhae Barnard, Paguristes engyops Barnard, Dromidia hirsutissima Lamarck, and Eudromidia hendersoni (Stebbing) occur on the west coast and also between Cape Point and Cape Agulhas, i.e., in the overlap zone. These species, together with Nautilocorystes ocellata (Gray), known from Walvis Bay to Port Elizabeth, and Callianassa australis Kensley, may be said to be true west coast endemics.

From a perusal of the distribution, the endemic species (Figure 3) would seem to reach a maximum from False Bay to Port Elizabeth (i.e., the Agulhas Bank region), with another peak in the Durban area. The Agulhas Bank maximum may to some extent be explained by the overlap of a few species from the cold west coast, with a larger number from the warm-temperate area south of the Transkei. The endemic peak found at Durban may be due to concentrated collecting, but probably reflects a true peak if compared with other well-sampled areas such as Inhaca Island and East London. This peak may be emphasized by the presence of a few warm-temperate species from the south added to the subtropical endemics of Natal and Southern Mozambique.

The 24% endemism found at Inhaca Island and Maputo may be more apparent than real, as collecting north of Delagoa Bay and on the East African coast generally has been scant. These peaks of endemism are worthy of further comment. Day (1973), in a discussion of the affinities of the fauna of Morrumbene estuary in Mozambique, analyzed the views of various authors on the East African shallow marine fauna. He defined tropical species as those occurring north of 20°S, subtropical from 20°S to Transkei, and warm-temperate south of the Transkei to False Bay. These definitions are adopted here, and it follows from the geographical limits of this paper that all the east coast endemic species north of the Transkei and south of Vilanculos should be regarded as subtropical. Ekman (1967) regarded all Natal and Mozambique species as tropical, as did Kalk (1959) in her analysis of the fauna of Inhaca Island. This is perhaps an oversimplification for the decapods, but it is thought that the peak of endemics found at Durban does indicate a distinct faunal component. The situation in southern Mozambique is uncertain due to lack of information, but it is probable that members of the subtropical component are well represented, and that a distinct tropical component is also present as indicated by Day (1973).

The question of faunal provinces around southern Africa has given rise to a considerable literature (see Brown and Jarman, 1978), starting in the mid-19th century and continuing to the present. With each separate group of organisms examined, a slightly different picture emerges, which is to be expected when factors such as habits, reproductive modes, life histories, and physico-biological requirements are considered. The situation may be briefly described for the shallow benthic decapods.

FIGURE 3.—Distribution of decapod Crustacea from less than 200 m, indicated by percentages at 8 localities (from west to east): Kunene River mouth, Lüderitz, Saldanha Bay, False Bay, Port Elizabeth, East London, Durban, Maputo.

The Tropical West African Province (Figure 4) barely makes its presence felt in northern South West Africa, such forms as Ocypode cursor (Linnaeus), Hippolyte palliola Kensley, and Maja squinado (Herbst) being limited in the south by the cold Benguela System and its concomitant upwelling.

The Namaqua or Cold-Temperate West Coast Province probably extends to Cape Agulhas and is characterized by few Indo-Pacific forms and some typically Atlantic species. Most of the endemics of this area are to be found on both sides of Cape Point, with no marked division at the Cape Peninsula, the temperature regime below about 30 m being relatively uniform.

The Warm-Temperate South Coast Province stretching from the overlap area of False Bay/Agulhas to Transkei is characterized by high numbers of endemics centered around the Algoa Bay (Port Elizabeth) region, by relatively high numbers of Atlantic forms, and by far fewer Indo-Pacifics than are found from Durban northwards.

The Subtropical East Coast Province extends from Transkei to about Inhambane in Mozambique, with a major Indo-Pacific component, but with a strong endemic element centered around the Durban area.

Origin of the Southern African Decapod Fauna

Good fossil decapod material is nowhere abundant in southern Africa; in fact, only five site records exist in the literature.

From the Cretaceous of Uitenhage, Cape Province, Kitchin (1913) recorded the palinuran mecochirid Meyeria. The genus is known from the Lower to Upper Cretaceous of Europe, North America, Mexico, West Africa, Angola, and Antarctica. The family as a whole has not been recorded later than the Cretaceous and thus has little bearing on the history of the present-day forms.

FIGURE 4.—Faunal provinces around southern Africa: (1) Tropical West African, (2) Namaqua or Cold Temperate West Coast, (3) Warm Temperate South Coast, (4) Subtropical East Coast, (5) Tropical East Coast. (Areas of overlap are only approximate.)

From the lower Miocene of Pemba Island, Stubblefield (1927) recorded the cancrid crab Palaeocarpilius intermedius, and a portunid, Neptunus, while from the lower Miocene of Inhambane, Mozambique, Stubblefield (1939) recorded another Neptunus as well as the extinct leucosiid crab Typilobus cf. granulosus.

King (1953) mentioned the presence of a “neptunid” (= portunid) crab from the lower Miocene deposits of Uloa, Zululand.

Böhm (1926) recorded Callianassa erecta, Callianassa cf. fraasi, and Brachyuridarus sp. (possibly a xanthid) from the lower Miocene of Bogenfels, South West Africa.

From such slender records, no useful conclusions can be drawn. Any speculations regarding the present-day distribution and origin of the decapod fauna must come from a consideration of the geological history of the area.

The southern African subcontinent, together with South America, India, Madagascar, Australia, and Antarctica formed the southern landmass of Gondwanaland (Crowson, 1970). During the Jurassic (± 160 × 10⁶ years b.p.) this landmass began to split up, with India, Madagascar, and Australia breaking away from Africa and exposing the eastern and southeastern continental edge of southern Africa. It was only during the upper Valanginian of the Cretaceous that South America and Africa began to separate, the break being completed during the Lower Turonian of the Cretaceous, 70 million years ago. The southeastern edge of Africa was thus exposed to oceanic conditions as well as to the faunal pioneers of the early Indian Ocean long before the North and South Atlantic joined and washed the southwestern continental margin. From this time on, changes in sea level, whether due to polar ice activity of the mid-oceanic ridges, accompanied by marine transgressions and regressions, took place especially during the Tertiary and Quarternary eras. These sea-level changes were undoubtedly reflected in changes in the faunal composition. There is a good evidence that the overall water-mass and current picture was considerably different from that of today. Micropalaeontological evidence from the east coast suggests that the Agulhas Current was not the well-marked year-round phenomenon it now is, during the last Pleistocene interglacial (Hutson, 1980). Further, if, as claimed by Prell and Hutson (1979), the Indian Ocean surface waters 18,000 years ago were much cooler than at present, and the Agulhas Current much weaker, the tropical Indo-Pacific fauna of the east coast must represent a relatively recent incursion, which is probably still taking place. As for the west coast fauna, studies of the Plio-Pleistocene molluscan fauna of raised beaches (Carrington and Kensley, 1969; Kensley, 1972, 1974b, 1977d; Tankard, 1975) indicate that a far more tropical and typically warm-water fauna prevailed up to the last interglacial in an area which is at present cold-temperate. Relicts of this warm-water fauna may have survived in sheltered and therefore warmer pockets such as Langebaan Lagoon, Saldanha Bay, and False Bay.

Decapoda from Neighboring Islands, Seamounts, and Shoals

Ascension Island, situated at 7°55′S, 14°30′W on the mid-Atlantic ridge, has little affinity with the southern African decapod fauna. The single species in common is Grapsus grapsus (Linnaeus) which is widespread through the Atlantic, and which occurs in northern South West Africa.

St. Helena Island, further south and closer to the African continent than Ascension, has a well-developed decapod fauna showing some affinity with southern Africa. Of the 23 species recorded (Chace, 1966) six are known from southern Africa. Of these, Grapsus grapsus (Linnaeus) and Calappa gallus (Herbst) are widespread through the tropical Atlantic; Planes cyaneus Dana is well known from the Indo-Pacific; Plagusia depressa Lamarck is known from both sides of the Atlantic; Dardanus arrosor (Herbst) from the eastern Atlantic and the Indo-Pacific; and Metalpheus paragracilis (Coutière) from the Indo-Pacific.

Whereas the fish, molluscs, and echinoderms of St. Helena show affinities first with the West Indies, and then with the Mediterranean and eastern North Atlantic, the decapods seem more closely related to the West African and southern African fauna, seven species being common to both areas. Chace (1966) expressed the view that only Planes cyaneus Dana, an oceanic species often found clinging to floating objects, came from southern Africa, having been carried by the trade wind drift.

Seamount Vema, situated 720 km off the coast of South Africa at 31°38′S, 8°20′E lies in the central region of the Cape Basin. It has a plateau-like summit some 7 km in width, at a depth of 45–80 m. Of the decapods from Vema, Pseudodromia cacuminis Kensley and Macropodia cirripilus Kensley are regarded as endemic (Kensley, 1980b), the former being closely related to Pseudodromia spinosissima Kensley from the east coast of South Africa. The single palinuran, Jasus tristani Holthuis, is also known from Tristan da Cunha, while Pseudactea corallina (Alcock) has Indo-Pacific affinity. Three species are true mesopelagics, Notostomus auriculatus Barnard, Gennadas gilchristi Calman, and Funchalia villosa (Bouvier). Of these, the Notostomus and Funchalia are true Atlantic forms, while G. gilchristi has only been recorded from the Agulhas Basin and the south-west Indian Ocean. Pagurus cuanensis (Bell), Eualus ctenifera (Barnard), and Pontophilus sculpta (Bell) have been recorded from southern Africa, while Pagurus chevreuxi Bouvier, Alpheus macrocheles (Hailstone), and Synalpheus huluensis africanus Crosnier and Forest are known from the Mediterranean and/or West Africa. The isopod fauna of Seamount Vema shows a much stronger affinity with southern Africa (Kensley, 1980b).

Seamount Tripp is situated at 29°36′S, 14°18′E off the coast of Namaqualand, in about 3000 m of water, and rises to about 160–170 m from the surface. One sample from the Sea Fisheries Branch has yielded two decapods, Paromola cuvieri (Risso) and Eumunida picta Smith. For both species, this is the most southerly record. The former has been recorded from the Mediterranean, Senegal, Cape Verde Islands, Azores, Ireland, Scotland, the Orkneys, Norway, the Shetlands, Congo, and Angola, and is a true Atlantic form. Eumunida picta Smith is known from the Canary Islands, Massachusetts, Australia, and New Zealand. These records suggest a widespread distribution.

Tristan da Cunha at 37°00′S, 12°50′E is an outlier of the mid-Atlantic ridge and well within the influence of the West Wind Drift. It is thus not surprising that Plagusia chabrus (Linnaeus) and Ovalipes punctatus (de Haan), both characteristically austral species, have been recorded here. Jasus tristani Holthuis is confined to this island group and to Seamount Vema. The only other records are a species of Pachygrapsus and a Notostomus sp. taken from an albatross' gut.

Walter's Shoal situated on the South Madagascar Ridge at 33°13′S, 43°51′E lies in about 38–46 m of water, and has been sampled by the R. V. Anton Bruun. The following five species have been recorded: Homola barbata (Fabricius) and Macropodia formosa Rathbun, both of which are known from the Atlantic and the east coast of South Africa; Platypodia granulosa (Rüppell), a typically Indo-Pacific species; Eualus ctenifera (Barnard), recorded from Maputo to Cape Point and from Seamount Vema; and Alpheus waltervadi Kensley, known only from this area.

Comparison of Decapoda with Other Benthic Crustacea from Southern Africa

When other crustacean groups from southern Africa are considered, a picture not too different from the decapods emerges. Griffiths (1977) noted a 46% endemism for amphipod species, concentrated in the south-western Cape Province. Griffiths further suggested that in spite of the fairly high endemism, the region derived its basic stock from tropical and southern temperate areas. J. A. Day (1979), working with the Cumacea, came to a similar conclusion, and in particular noted the cutoff in numbers somewhere between Durban and East London, which she took to indicate the presence of a boundary between faunistic provinces. A similar cutoff has been noted above for the decapods. From unpublished records I have been able to determine that the isopods follow a pattern similar to that of the Cumacea and amphipods, but with even higher endemism centered in the southwestern Cape-Agulhas Bank area. The endemic peak noted for the decapods in the Durban area was not apparent for the abovementioned peracaridan groups.

Comparison of the Decapod Faunas of Australia, New Zealand, South America, and Southern Africa

Little useful information emerges from comparison of the decapod faunas of the Southern Hemisphere regions.

The decapod fauna of Australia (Griffin and Yaldwyn, 1968) is almost double the size of the southern African fauna, and may be broadly divided into a tropical component and southern temperate component. In general terms, the fauna of Australia is almost entirely derived from the Indo-Pacific. Although there are several genera in common with southern Africa, relatively few species are shared. Interestingly, of the 10 largest decapod families in southern Africa, seven of these occur in the 10 largest Australian families, with Xanthidae, Majidae, and Portunidae, in this order, being the three largest in both areas. The Majidae of southern Africa have both Atlantic and Pacific components, whereas the Australian majids (Griffin, 1966a) are almost entirely Indo-Pacific. Nine species are common to South Africa and Australia, with only Achaeopsis thomsoni (Norman) not having a tropical distribution. (See Table 3.)

The Australian Thalassinidea are richer and have more endemic species than in southern Africa, with no species in common. (See Table 4.)

TABLE 3.—Comparison of the majid crab fauna of southern Africa, Australia, and New Zealand (Australian figures from Griffin, 1966a; New Zealand figures from Griffin, 1966b)

TABLE 4.—Comparison of the thalassinidean fauna of Australia and southern Africa (Australian figures from Poore and Griffin, 1979)

The New Zealand brachyuran (Dell, 1968) as well as the anomuran and natantian (Yaldwyn, 1967) forms are oceanic in character with 53%–55% endemism, showing close affinity to the Australian fauna, and then to the Indo-Pacific in general, and with very little in common with southern Africa. Plagusia chabrus (Linnaeus), Lithodes murrayi Henderson, and Eumunida picta Smith seem to be the only benthic species shared. The New Zealand and southern African records for Palaemon (N.) tenuipes (Henderson) need to be reexamined.

Very little information is available from South America (see Coelho, Ramos-Porto, and Koening, 1978). Ovalipes punctatus (de Haan), Pilumnoides perlatus (Pöppig), and Plagusia chabrus (Linnaeus) from southern Africa and South America have been discussed above in relation to austral distribution. The South American records of Ogyrides occidentalis (Ortmann) from Brasil, and Ibacus incisus (Peron) from Chile are suspect, while the taxonomic position of the South American Cyclograpsus punctata H. Milne-Edwards needs reappraisal.

Checklist of Southern African Decapoda

SOURCES OF DATA.—Apart from the many published records on southern African decapods, for compilation of the present list I have drawn on the entire holdings and records of the South African Museum, many of which are unpublished, as well as those of the Department of Zoology of the University of Cape Town. Collections from the Sea Fisheries Branch of the Department of Industry donated to the South African Museum in 1979 have also been processed, and the records included.

NOTES ON THE CHECKLIST.—The author and date of publication for each species is provided. A recent (often the only) reference to the species' occurrence in southern African waters is provided. As Barnard (1950) mentions many of the species, and is often the most useful reference, for brevity this reference is indicated by the letter “B.”

The depth distribution information (in meters) pertains only to the southern African records.

The geographic range within the defined southern African region is given from west to east.

Where no reference to a southern African record is given, the record comes from the South African Museum's collections but has not previously been published.

No attempt has been made to separate the Indian Ocean or Indo-West Pacific regions. These are included under the broad heading of Indo-Pacific.

Although every attempt has been made to ensure that the most recently accepted nomenclature is used, inaccuracies may still be present, while taxonomic changes will certainly take place to alter the list.

Family PENAEIDAE

Family ARISTEIDAE

Subfamily ARISTEINAE

Subfamily BENTHESICYMINAE

Family SOLENOCERIDAE

Family SICYONIIDAE

Family SERGESTIDAE

Family STENOPODIDAE

Family OPLOPHORIDAE

Family ATYIDAE

Family NEMATOCARCINIDAE

Family STYLODACTYLIDAE

Family PASIPHAEIDAE

Family BRESILIIDAE

Family RHYNCHOCINETIDAE

Family PALAEMONIDAE

Subfamily PALAEMONINAE

Subfamily PONTONIINAE

Family GNATHOPHYLLIDAE

Family ALPHEIDAE

Family OGYRIDIDAE

Family HIPPOLYTIDAE

Family PROCESSIDAE

Family PANDALIDAE

Family CRANGONIDAE

Family GLYPHOCRANGONIDAE

Family NEPHROPIDAE

Family POLYCHELIDAE

Family PALINURIDAE

Family SCYLLARIDAE

Family AXIIDAE

Family CALLIANASSIDAE

Family UPOGEBIIDAE

Family PYLOCHELIDAE

Family DIOGENIDAE

Family COENOBITIDAE

Family PAGURIDAE

Family PARAPAGURIDAE

Family LITHODIDAE

Family GALATHEIDAE

Family CHIROSTYLIDAE

Family INCERTAE SEDIS

Family PORCELLANIDAE

Family RANINIDAE

Family ALBUNEIDAE

Family HIPPIDAE

Family HOMOLODROMIIDAE

Family DROMIIDAE

Family DYNOMENIDAE

Family TYMOLIDAE

Family HOMOLIDAE

Family LATREILLIDAE

Family DORIPPIDAE

Family CALAPPIDAE

Family LEUCOSIIDAE

Family MAJIDAE

Family HYMENOSOMATIDAE

Family PARTHENOPIDAE

Family CORYSTIDAE

Family ATELECYCLIDAE

Family GERYONIDAE

Family PORTUNIDAE

Family XANTHIDAE

Family GONEPLACIDAE

Family HEXAPODIDAE

Family GRAPSIDAE

Family GECARCINIDAE

Family PINNOTHERIDAE

Family POTAMONAUTIDAE

Family OCYPODIDAE

Family RETROPLUMIDAE

Family PALICIDAE

Family HAPALOCARCINIDAE

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1971a.  The Family Sergestidae in the Waters around Southern Africa (Crustacea, Decapoda, Natantia). Annals of the South African Museum, 57:215–264.

1971b.  The Genus Gennadas in the Waters around Southern Africa. Annals of the South African Museum, 57: 271–294.

1972.   Pliocene Marine Invertebrates from Langebaanweg, Cape Province. Annals of the South African Museum, 60:173–190.

1973.   A New Species of Hermit Crab from Natal, South Africa (Decapoda, Anomura, Paguridae). Durban Museum Novitates, 9:285–290.

1974a.  The Genus Callianassa (Crustacea, Decapoda, Thalassinidea) from the West Coast of South Africa, with a Key to the South African Species. Annals of the South African Museum, 62:265–278.

1974b.  The Status of the Plio-Pleistocene Panopea in Southern Africa (Mollusca, Bivalvia, Hiatellidae). Annals of the South African Museum, 1974:199–215.

1974c.  A Review of the Southern African Decapod Crustacean Fauna. Doctoral dissertation, Univeristy of Cape Town.

1975.   Records of Mud-Prawns (Genus Callianassa) from South Africa and Mauritius (Crustacea, Decapoda, Thalassinidea). Annals of the South African Museum, 69:47–58.

1977a.  The South African Museum's Meiring Naude Cruises, 5: Crustacea, Decapoda, Reptantia and Natantia. Annals of the South African Museum, 74:13–44.

1977b.  The South African Museum's Meiring Naude Cruises, 2: Crustacea, Decapoda, Anomura and Brachyura. Annals of the South African Museum, 72: 161–188.

1977c.  Lambrachaeus ramifer Alcock, a Rare Spider Crab from the East Coast of Southern Africa (Decapoda, Brachyura, Majidae). Zoologica Africana, 12: 323–327.

1977d.  A Second Assemblage of Pliocene Invertebrate Fossils from Langebaanweg, Cape. Annals of the South African Museum, 72:189–210.

1978.   Decapod Crustaceans Collected in Southern African Waters by the Th. Mortensen Java-South Africa Expedition (Crustacea, Decapoda). Steenstrupia, 4:249–261.

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1980b.  Decapod and Isopod Crustaceans from the West Coast of Southern Africa, Including Seamounts Vema and Tripp. Annals of the South African Museum, 83:13–32.

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