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Kak SAY Nae Re Cot ONE Dearest Renan ee pee wy : SIV OT Vey TATU eRe tam DENY Ree oA Rts iedsSyi wea ark ait i Fp Athy Means 2D aR Ahern ke Ae III Ged PIE Te My Deri ey deed Py haMehe rows VR RAM TACOMA AL OM as iniker teria ead WV eye Ge RY bw re nists Tape tea o® recone vile “wih Be A Suton ti) tte wt » Lali reate den, “ten rect Hag Met MLAs eS SUM NS nea a ERO eng eke Gesell ah) Sitti Beha gurearh, Dies Pawn: ” wU AS y OR tes A Wo Vette Pha Ma Aeon Ral tslp Be we Mee “AOA AP VISES ATR EDC OF EEWE 2 AN Whi BN, MUTI TOY EY Be oF ait aig ot ey iaitie te ey NEAL Ae sane Amits iss ely (beim, PNM ey BO i ovens Wy hohe eather YY paver Plame’, é, Permineide Ns say PALO ESO Ogre eer MER we Pid k eGR MALO 5 hee oui dif eter a of D THAD Teale tay te? COTININSES FOS Nice Vea bell RDF setae ir ta yet ere he tl OW ES NN Tata F . seas + Trae Wn abayeanes ahahs IAN Ee bas te Bey TerbeRalaTe SAG Me Svein? Geet ice Ms the Se ees my NEM es Syke, ET ye Chet es teehee ath Sash ei oe edie i Wee Me a th tags ety meh ee RN carat Seria ently MR EY gat MRT Me Siete eee ae ee ae Geb det intehs Neihaciea Niicits res wate AER DMN oes Ws ate embed Gavyhrk clawed Se ae eae gt Ae EO STO Cond 6 Bed Vetere RS ens ne m0 Seatetrt Serer ror O¥ Sey OS A Se RE ae reel et BADER Dud er eee isn ove sie SGIRZ ES toa Wt ene Fe cme it ee er ee SSerninapoyny oR Po bnasine = Fr ae Ms eV Nay Heath ahh EMS AS Nea SUP nde DR Me hae te Yaga Wey MOOT WME CEA ODE MATA ee eat dy Mone eer tgt gh Mc site AANA NTR oN agate Behe BeBe TREN By Rest TSO, 1 vor kyla Ray ee AS pea yall Teh Ms eee Mie. re on ee Oe a Ne Dee eee es fc i Ce Lovee atheee she UF EAE aye Theta teat POL ras Stet by dain Patel Ses Se tte Sy ey Nyt Te eg oN eel La pelea Lee rR nae Sie ec Son PAGE Tt eh Te be DeeMVW faytks eaFae ie Hey ASHES MUD EAL! BE Ty Se Meta Ta eres By ote sett Fe ANE e Paaet ia Doybstite 3 = iy Artes st Seer rae i or? nivergligitee apm 2 Men BA oie why tea hema Tag Pte ae ee ee ere tae ed ees Fe er ee oot be roaye moras Setlist OP ALO Geb ome eraee hye Oe Rae red. sents td De Me Mae pete ERSTE CMON Ln Manca ten ace Ss Beha tay 8, wh ce eo SP Nae Sahin? 2 nell ete Fed rata +a ek ab ylod ge at ore Mortar ge 1 REP I on We pcre ete HARVARD UNIVERSITY Library of the Museum of Comparative Zoology Ate ©: Gas ee, | ; tnd a = i -Q 4 * 1 ) \ i i 1 4 et MEY s 4 ¥ ANNALS OF THE ANNALE VAN DIE SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM VOLUME 74 BAND 74 \a ba « ev ¢ = « » a ‘ mA ‘ 7 1 ae 3 F é ; 5 4 Th —f t i o Y : , ; ’ a,* . c “4 . bh ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM VOLUME 74 BAND THE TRUSTEES OF THE DIE TRUSTEES VAN DIE SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM CAPE TOWN KAAPSTAD 1977-1978 # SET, PRINTED AND BOUND IN THE REPUBLIC OF SOUTH AFRICA BY THE RUSTICA PRESS (PTY.) LTD., WYNBERG, CAPE 524 LIST OF CONTENTS Avery, D. M. Past and present distribution of some rodent and insectivore species in the southern Cape Province, South Africa: new information. (Published December 1977.) EAsTWwoop, E. B. Notes on the scorpion fauna of the Cape. Part 3. Some observations on the distri- bution and biology of scorpions on Table Mountain. (Published January 1978.) EASTWOop, E. B. Notes on the scorpion fauna of the Cape. Part 4. The burrowing activities of some scorpionids and buthids (Arachnida, Scorpionida). (Published January 1978.) GLAESSNER, M. F. Re-examination of Archaeichnium, a fossil from the Nama Group. (Published April 1978.) .. ie me GRIFFITHS, C. The South African Museum’s a tea Naude cruises. Part 6. Giese area November 1977.) é ; GRINDLEY, J. R. A new species of Tortanus (Crustacea, Copepoda) from South Africa. (Published January 1978.) : a se We : KENNEDY, W. J. & KLINGER, H. C. Cretaceous faunas from Zululand and Natal, South Africa. A Jauberticeras from the Mzinene Formation (Albian). (Published November 1977.) KENNEDY, W. J. & KLINGER, H. C. Cretaceous faunas from Zululand and Natal, South Africa. A Flickia from the Cenomanian of northern Zululand. (Published January 1978.) KENNEDY, W. J. & KLINGER, H. C. Cretaceous faunas from Zululand and Natal, South Africa. The ammonite family Lytoceratidae Neumayr, 1875. (Published February 1978.) _KENSLEY, B. The South African Museum’s Meiring Naude cruises. Part 5. Crustacea, Decapoda, Reptantia and Natantia. (Published December 1977.) .. : KENSLEY, B. The South African Museum’s Meiring Naude cruises. Part 7. Marine Isopoda. (Published January 1978.) .. KLINGER, H. C. see KENNEDY, W. J. McKENZIE, K. G. Illustrated generic key to South African continental Ostracoda. (Published Novem- ber 1977.) ; MILLARD, N. A. H. The Se distribution of southern African hydroids. (Published January 1978.).. : a Ke me By aN es S% id Page 201 Vipes) 249 335 105 US) ZAM PEST) 13 2S 45 159 NEW GENERIC NAMES PROPOSED IN THIS VOLUME Page Izinkala Griffiths, 1977 ae ie ra A: he ra ar od An 115 Kapcypridopsis McKenzie, 1977 Bes Ae a: a ae ae 4 ae 46 Sarscypridopsis McKenzie, 1977 ae ae at fe Bat Ls ake He 48 Spinarcturus Kensley, 1978 .. as Me Bie a as ae Spe Se) Valettiella Griffiths, 1977 5 Sc =i on oe Be si si’ a 2G) a i ; rag > | . 1 CuUsy lath . ; ey Ct eA et Ae “an oe 1 M00 @eeca Py Bivorweaas ate ey inane iia + -- ‘VGA Salter Sia iat Y a ; — SOU VOLUME 74 PART 1 NOVEMBER 1977 | ISSN 0303-2515 MUS. COMP. ZOOL. LIBRARY ‘SAN 3 1 1978 HARVARD UNIVERSITY ANNALS OF THE SOUTH AFRICAN CAPE TOWN - INSTRUCTIONS TO AUTHORS 1. MATERIAL should be original and not published elsewhere, in whole or in part. 2. LAYOUT should be as follows: (a) Centred masthead to consist of Title: informative but concise, without abbreviations and not including the names of new genera or species Author’s(s’) name(s) Address(es) of author(s) (institution where work was carried out) Number of illustrations (figures, enumerated maps and tables, in this order) (b) Abstract of not more than 200 words, intelligible to the reader without reference to the text (c) Table of contents giving hierarchy of headings and subheadings (d) Introduction (e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents (f) Summary, if paper is lengthy (g) Acknowledgements (h) References (i) Abbreviations, where these are numerous 3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced with 2,5 cm margins all round. First lines of paragraphs should be indented. 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REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: ‘Smith (1969) describes... .’ ‘Smith (1969: 36, fig. 16) describes . “As described (Smith 1969a, 19695; ee pales ‘As described (Haughton & Broom ee ‘As described (Haughton et al. 1927) . Note: no comma separating name and: year pagination indicated by colon, not p. names of joint authors connected by ampersand et al. in text for more than two joint authors, but names of all authors given in list of references. (b) Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, b, etc. to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a). For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list o scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number (only if independently paged) in parentheses, pagination (first and last pages of article). Examples (note capitalization and punctuation) BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140. FISCHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. gen. 74: 627-634. Konan, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320. Konn, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51 THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. (continued inside back cover) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 74 Band November 1977 November Part 1 Deel CRETACEOUS FAUNAS FROM ZULULAND AND NATAL, SOUTH AFRICA A JAUBERTICERAS FROM THE MZINENE FORMATION (ALBIAN) By WILLIAM JAMES KENNEDY & HERBERT CHRISTIAN KLINGER Cape Town Kaapstad The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular intervals as material becomes available Obtainable from the South African Museum, P.O. Box 61, Cape Town Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM word uitgegee in dele op ongereelde tye na beskikbaarheid van stof Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad OUT OF PRINT/UIT DRUK 1, 201-3, 5-8), 3(1-2, 4—5, 8, t.-p.i.), 51-3, 5, 7-9), 6(1, t.-p.i.), 701-4), 8, 911-2, 7), 10(1-3), 11(1-2, 5, 7, t.-p.i.), 15(4—-5), 24(2), 27, 31(1-3), 33 Price of this part/Prys van hierdie deel R1,90 Trustees of the South African Museum © Trustees van die Suid-Afrikaanse Museum 1977 ISBN 0 908407 25 4 Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk., Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap CRETACEOUS FAUNAS FROM ZULULAND AND NATAL, SOUTH AFRICA A JAUBERTICERAS FROM THE MZINENE FORMATION (ALBIAN) By WILLIAM JAMES KENNEDY Geological Collections, University Museum, Oxford & HERBERT CHRISTIAN KLINGER South African Museum, Cape Town (With 6 figures) [MS. accepted 28 April 1977] ABSTRACT The ammonite subfamily Gabbioceratinae Breistroffer, 1953, is represented in the Cretaceous of southern Africa by a unique specimen of Jauberticeras collignoni Murphy from the Albian Mzinene Formation exposed along the Mzinene River, Zululand. The specimen shows details of suture, lirae and constrictions not previously known in the species, until now based on the unique Madagascan holotype alone, allowing a fuller diagnosis, description and comparison with other species. The holotype and the types of the closely related species J. jaubertianum (d’Orbigny) and J. villoutreysi Murphy are reillustrated for comparative purposes. CONTENTS PAGE Introduction : : ; i i l Systematic palaeontology . : ; 2 Acknowledgements . ‘ , 4 1] References : , ‘ , ; 11 INTRODUCTION In two previous papers (Kennedy & Klinger 1977, and in press) the authors have documented the representatives of the great majority of the ammonite superfamily Tetragonitaceae Hyatt, 1900 known from the Cretaceous sediments of Zululand and Natal. In the course of examining the collections of the late Dr E. C. N. van Hoepen, recently transferred to the South African Museum from the National Museum Bloemfontein, one of the authors (H. C. K.) recog- nized a further representative of this superfamily, in the form of a Jauberticeras Jacob, 1907, from the Upper Albian Mzinene Formation of Zululand. Jauberticeras is one of the two genera referred to the subfamily Gabbio- ceratinae Breistroffer, 1953 (the other is the genus Gabbioceras Hyatt, 1900); the specimen represents not only the first record of the genus from southern Africa, but also the first record of the subfamily from the region. l Ann. S. Afr. Mus. 74 (1), 1977: 1-12, 6 figs. D) ANNALS OF THE SOUTH AFRICAN MUSEUM SYSTEMATIC PALAEONTOLOGY Class CEPHALOPODA Cuvier, 1797 Order AMMONOIDEA Zittel, 1884 Suborder LYTOCERATINA Hyatt, 1889 Superfamily TETRAGONITACEAE Hyatt, 1900 Family Tetragonitidae Hyatt, 1900 Subfamily Gabbioceratinae Breistroffer, 1953 The subfamily Gabbioceratinae has been the subject of two exhaustive accounts in recent years (Wiedmann 1962; Murphy 1967), and there is a diver- gence of opinion over which species should be referred to Gabbioceras versus Jauberticeras, as well as over the systematic position of a number of species regarded by Wiedmann as Gabbioceras and by Murphy as Eofetragonites. More serious is the disputation of the position of Parajaubertella Matsumoto, 1943, regarded by Wiedmann as a synonym of Gabbioceras and by Murphy as an independent homoeomorphous offshoot of the Eogaudryceras stock. There is, however, agreement that the subfamily has its origin in the group of Eogaudry- ceras (Eogaudryceras) numidum (Coquand). From this the line of descent runs to Gabbioceras, with G. lamberti (Breistroffer) as the earliest known member of that genus giving rise to Jauberticeras via J. jaubertianum during early late, or late early Aptian time. Jauberticeras is confined to the late Aptian and Albian; Gabbioceras extends to the Cenomanian. Genus Jauberticeras Jacob, 1907 Type species Ammonites jaubertianus d Orbigny (1850: 200, figs 9-10) by original designa- tion (see Howarth 1974 for clarification of the date and place of publication of Jauberticeras and of Ammonites jaubertianus as type species). Diagnosis Relatively small cadicones with a funnel-shaped umbilicus, no flank and a marked angulation demarcating umbilicus from the broad, flattened venter. Shell surface lirate with constrictions (sometimes only visible on the mould). Suture with three umbilical lobes. Discussion The authors take this opportunity to illustrate (Fig. |!A—G) a plaster cast of the lectotype of J. jaubertianum, the original of which is from the Aptian of Hyeges, Basses Alpes, France. When compared with Gabbioceras (a typical example of which is shown in Fig. 2A—G), the two genera are best distinguished on whorl section (following Murphy 1967: 604): in Gabbioceras the venter is strongly curved (Fig. 2A, C, D, G): in Jauberticeras it is flat (Fig. 1A—B, D-F). CRETACEOUS FAUNAS FROM SOUTH AFRICA 3 The angulation is commonly sharper in Jauberticeras when compared with early Gabbioceras (Murphy 1967, fig. 1). Occurrence Late Aptian to mid Albian of southern France, Spain, the Caucasus, north Africa, Madagascar and Zululand. F G ° Fig. 1. Jauberticeras jaubertianum (d’Orbigny). The lectotype from the Aptian of Hyéges, Basses Alpes, France (Peticlerc Collection, Sorbonne, Paris). A-C x 2; D-G ~x 1. 4 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 2. Gabbioceras lamberti (Breistroffer). BMNH C10780, from the Aptian of Castellane, Basses Alpes, France. A-C x 2; D-G x 1]. CRETACEOUS FAUNAS FROM SOUTH AFRICA 5 Jauberticeras collignoni Murphy, 1967 Figs 3-5 Jauberticeras latecarinatum Collignon non Anthula, 1963: 28, pl. 251 (fig. 1080). Jauberticeras collignoni Murphy, 1967: 606, pl. 64 (fig. 16). Holotype M. Collignon Collection no. 435, the original of Collignon (1963: 28, pl. 251 (fig. 1080)) by monotypy. Reproduced here as Fig. 3A-H. From the Albian sensu lato of Betioky, Madagascar. Material . Specimen D293 in the E. C. N. van Hoepen Collection. Emended diagnosis A Jauberticeras with a blunt lateral angle, moderately expanding whorls, weak lirae and irregularly spaced prorsiradiate convex constrictions and a prominent second saddle on the internal suture. Description Specimen D293 is a beautifully preserved mould of a phragmocone 21,5 mm in diameter, infilled with sparry calcite, and retaining a short portion of the body chamber filled with black quartzose glauconitic sandstone. Traces of the original shell material with delicate external ornament are preserved on the umbilical wall, but on the outer surface the shell is partially or wholly absent. Dimensions of the specimen, and others with which it is compared, are shown below. (D = diameter; Wb = whorl breadth; Wh = whorl height; U = umbilical diameter from seam to seam; K = umbilical diameter from keel to keel. Figures in parentheses are dimensions expressed as percentages of the total diameter.) D Wb Wh Wb:Wh U K J. collignoni 22,4 (100) 16,3 (74) 6,9 (31) 2,36 ~—-9,8 (44)_—‘:18,7 (84) D293 8200), 13.3,a)) 5.5 CO. 24N 74441) 14.978) MC435 OFS) (OO) lS (68) 523 (27) ae 4 8,1 (42) 14,9 (77) J. yilloutreysi | BMNH C68145 27,8 (100) 18,5 (67) 8,8(32) 2,13 12,0(43) 22,5 (81) J. jaubertianum lectotype SOR COO) Fe IOSD Os G2) e485 058833) = (=) The coiling is very evolute, so that the angulation of the successive whorls is exposed in the umbilicus, giving it terraced, rather than evenly sloping sides. The umbilical wall is gently convex, whilst the angulation between it and the venter is abruptly rounded and blunt. The venter is broadly and evenly rounded, the distance between angulations comprising 84 per cent of the maximum diameter. The whorls expand quite slowly, so that there is a distinct tapering 6 ANNALS OF THE SOUTH AFRICAN MUSEUM of the whorls towards the apex in both dorsal and ventral views. The mould bears low, faint, irregular folds and interspaces, which are markedly prorsiradiate on the flanks and sweep across the venter with a broad and shallow convex curvature. Two of these folds are strengthened into broad collars, one at the beginning of the body chamber, the other at a diameter of just over 20 mm. These are succeeded by distinct if shallow constrictions. On the venter, patches of partially exfoliated shell appear smooth, save for the low folds already mentioned; a few tiny areas preserve fine, dense lirae, invisible to the naked eye. In contrast, areas of umbilical wall where shell is well preserved show distinct if fine prorsiradiate lirae, concave close to the umbilical seam, but weakly flexed so as to be gently convex across the angulation. So far as can be discerned the whole of the shell was finely lirate, lirae paralleling the folds and constrictions visible on the mould. The external suture is well exposed (Fig. 4) and 1s deeply and intricately subdivided, as is characteristic of other species of the genus. The median saddle of the external lobe is prolonged into a series of narrow, tapering processes which strongly recall the siphonal band of certain tetragonitids (Kennedy & Klinger 1977). Fig. 3. Jauberticeras collignoni Murphy. Holotype, M. Collignon Collection no. 435, from the Albian sensu lato of Betioky, Madagascar. A-D x 2; E-H ~x 1. CRETACEOUS FAUNAS FROM SOUTH AFRICA 7 Fig. 4. Sutures of Jauberticeras collignoni Murphy. Based on specimen D293 (Van Hoepen Collection) from the Mzinene Formation of the Mzinene River, Zululand, and of late middle or early late Albian age. A « 12: B < 6 to show details of siphonal structures. 3) ANNALS OF THE SOUTH AFRICAN MUSEUM E F G H Fig. 5. Jauberticeras collignoni Murphy. D293 (Van Hoepen Collection) from the Mzinene Formation of the Mzinene River, Zululand, and of late middle or early late Albian age. A-Dix 22. E=H << Ii, CRETACEOUS FAUNAS FROM SOUTH AFRICA 9 Discussion Murphy based this species on the specimen figured by Collignon (1963, pl. 251 (fig. 1080)) which he refigured in dorsal view only (Murphy 1967, pl. 64 (fig. 16), text-fig. 1 (9)). The original description is short, and in it Murphy characterized the species by its low expansion rate, smooth shell, and strong second saddle in the internal suture, lack of lirae and constrictions and similar whorl section to that of the type species. As can be seen from the holotype, here reillustrated (Fig. 3), the specimen does not quite fit Murphy’s description in possessing constrictions. There are at least three distinct constrictions on the outer whorl, markedly prorsiradiate on the flank, and feebly convex across the venter, each being preceded by a slight swelling. The whorl section, with a breadth to height ratio of 2,54 at the maximum preserved diameter and 2,08 360° before this, comprises only 68 per cent of the total diameter whereas in J. jaubertianum, to which it was said to be similar, the figure is 71 to 94 per cent. The whorl angulation is also significantly blunter. The latter, however, may be partially due to preservation or even of intra-specific value only, being dependent on ontogeny (see Murphy 1967, pl. 64 (fig. 23)). The second internal saddle is, however, prominently developed, whilst the shell surface is smooth, even under a lens. Close inspection shows that this may be no more than an artefact of preservation; the originally aragonitic shell has been replaced by iron-stained sparry calcite, and the whole of the surface of the umbilicus of the specimen is corroded and pitted. The venter is similarly pitted, and in places the angulation is covered and accentuated by traces of a succeeding whorl, now lost. This preservation and the remaining adherent matrix of fine sandstone suggest the specimen to be from a precisely similar setting to many of the Albian and Cenomanian ammonites from Zululand which the authors have studied, especially those found weathering out along the terraces of the Mzinene and on the Skoenberg, where identically preserved specimens of diverse ammonites have lost delicate growth lines and lirae by post-mortem processes. Lack of liration is not, therefore, a diagnostic feature of the species in the authors’ view. With these qualifications in mind, the Zululand specimen, although larger than the Madagascan type, can be referred to Jauberticeras collignoni on the basis of similarities of whorl section, bluntness of angulation, relative propor- tions (which are similar but not identical), expansion rate, form and direction of constrictions, and associated collar-like swellings. Because the present specimen does not show the septal face (retaining as it does part of the body chamber) the nature of the internal suture is not determinable. The principal difference between J. collignoni and the type species, J. jauber- tianum (of which J. latecarinatum (Anthula) (1899: 101, pl. 7 (fig. 2a—c)) is probably a synonym according to Murphy (1967: 605-6)), lies in the more prominent development of the second saddle in the internal suture. Jauberticeras villoutreysi Murphy (1967: 606, pl. 64 (figs 25—29)) (the holo- type BMNH C68148 is reproduced here as Fig. 6A—G) has much coarser lirae on the umbilical wall, which give a distinctive appearance to the whorl angulation 10 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 6. Jauberticeras villoutreysi Murphy. Holotype, BMNH C68148, from the Albian of Clar, Alpes-Maritimes, France (Astier Collection). A-C x 2; D-G x 1. CRETACEOUS FAUNAS FROM SOUTH AFRICA II (Fig. 6A). In addition, the expansion rate is such that the last half whorl is almost parallel-sided in profile (Fig. 6B, F), the venter flatter, and constrictions straight across the venter with a broad, band-like collar associated. In addition the saddles in the internal suture are of different sizes (Fig. 6C). Jauberticeras lanternoi Wiedmann (1962: 32, pl. 2 (fig. 2)) has quite different proportions and whorl section, and may possibly be a juvenile Gabbioceras (M. A. Murphy pers. comm. 7.3.1977). J. subbeticum Wiedmann (1962: 31, pl. 2 (fig. 3a—c)} has much coarser umbilical liration and a different form of whorl section and constrictions, and again is of dubious generic affinity. According to Murphy (pers. comm. 7.3.1977) it is probably a Scaphites. | Occurrence The holotype of Jauberticeras collignoni is from the Albian sensu lato of Collignon’s locality 435, ‘Pied Est du Vohimaranitra (Betioky), Madagascar’. The Zululand specimen is from the Albian part of the Mzinene Formation along the Mzinene River near Hluhluwe. Precise locality data are lacking, but preserva- tion suggests it to be from locality 51 of Kennedy & Klinger (1974: 288, fig. 4) where concretions are exposed in the stream bed and bank extending around the broad meander of the river east-north-east of the Sisal Factory at 27°53'43’S, 32°19'22”E. The specimen is thus of late middle or early late Albian age (Albian IV-V). ACKNOWLEDGEMENTS Our best thanks are due to General M. Collignon of Moirans (Jsere), who kindly allowed us to study the holotype of Jauberticeras collignoni, and has helped us throughout our work on the Zululand Cretaceous. We are also grateful to Dr M. K. Howarth and Mr D. Phillips (British Museum, Natural History) for allowing us to study the specimens in their care, and Dr J. Sornay of the Muséum Nationale d’Histoire Naturelle, Paris, for help in our study of Specimens in the D’Orbigny collection. REFERENCES ANTHULA, D. J. 1899. Ueber die Kreide Fossilien des Kaukasus. Beitr. Paldont. Geol. Ost. Ung. 12: 54-159. COLLIGNON, M. 1963. Atlas des Fossiles caractéristiques de Madagascar (Ammonites), Fascicule X (Albien). Tananarive: Service Géologique. Howarth, M. K. 1974. The Lower Cretaceous ammonite genera proposed by C. Jacob in 1907. Palaeontology 17: 727-728. KENNEDY, W. J. & KLINGER, H. C. 1974. Cretaceous faunas from Zululand and Natal, South Africa. Introduction, Stratigraphy. Bull. Br. Mus. nat. Hist. (Geol.) 25: 263-315. KENNEDY, W. J. & KLINGER, H. C. 1977. Cretaceous faunas from Zululand and Natal, South Africa. The ammonite family Tetragonitidae Hyatt, 1900. Ann. S. Afr. Mus. 73: 149-197. KENNEDY, W. J. & KLINGER, H. C. In press. Cretaceous faunas from Zululand and Natal, South Africa. The ammonite family Gaudryceratidae Spath, 1927. Bull. Br. Mus. nat. Hist. (Geol.). 1 ANNALS OF THE SOUTH AFRICAN MUSEUM Murpny, M. A. 1967. The Ammonoid subfamily Gabbioceratinae Breistroffer. J. Paleont. 41: 595-607. OrBIGNY, A. D’. 1850. Note sur quelques nouvelles espéces remarquables des étages Neocomien et Aptien de France. J. Conchyliol. 1: 196-201. WIEDMANN, J. 1962. Die Gabbioceratinae Breistroffer. Newes Jb. Geol. Paldont. Abh. 115: \-43. 6. SYSTEMATIC papers must conform to the International code of zoological nomenclature (particularly Articles 22 and 51). Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov., etc. An author’s name when cited must follow the name of the taxon without intervening punctuation and not be abbreviated; if the year is added, a comma must separate author’s name and year. The author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is transferred from its original genus. The name of a subsequent user of a scientific name must be separated from the scientific name by a colon. Synonymy arrangement should be according to chronology of names, i.e. all published scientific names by which the species previously has been designated are listed in chronological order, with all references to that name following in chronological order, e.g.: Family Nuculanidae Nuculana (Lembulus) bicuspidata (Gould, 1845 Figs 14-15A Nucula (Leda) bicuspidata Gould, 1845: 37. Leda plicifera A. Adams, 1856: 50. Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b). Nucula largillierti Philippi, 1861: 87. Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. Note punctuation in the above example: comma separates author’s name and year semicolon separates more than one reference by the same author full stop separates references by different authors figures of plates are enclosed in parentheses to distinguish them from text-figures dash, not comma, separates consecutive numbers Synonymy arrangement according to chronology of bibliographic references, whereby the year is placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable. In describing new species, one specimen must be designated as the holotype; other speci- mens mentioned in the original description are to be designated paratypes; additional material not regarded as paratypes should be listed separately. The complete data (registration number, depository, description of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.: Holotype SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973. Note standard form of writing South African Museum registration numbers and date. 7. SPECIAL HOUSE RULES Capital initial letters (a) The Figures, Maps and Tables of the paper when referred to n the text e.g. “.. . the Figure depicting C. namacolus...’; ‘. . . in C. namacolus (Fig. 10)...’ (b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded by initials or full names e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene (c) Scientific names, but not their vernacular derivatives e.g. Therocephalia, but therocephalian Punctuation should be loose, omitting all not strictly necessary Reference to the author should be expressed in the third person Roman numerals should be converted to arabic, except when forming part of the title of a book or article, such as “Revision of the Crustacea. Part VIII. The Amphipoda.’ Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial capital letter, provided the same generic name is used consecutively. Name of new genus or species is not to be included in the title: it should be included in the abstract, counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts. WILLIAM JAMES KENNEDY & HERBERT CHRISTIAN KLINGER CRETACEOUS FAUNAS FROM ZULULAND AND NATAL, SOUTH AFRICA A JAUBERTICERAS FROM THE MZINENE FORMATION (ALBIAN) \ as WKS 2 DECEMBER 1977 ISSN 0303-2515 ; MLSs, NOV 1 9 ygQQn Ft A. ine 47 fovin, at me) ANNALS OF THE SOUTH AFRICAN UM ‘APE TOWN INSTRUCTIONS TO AUTHORS 1. MATERIAL should be original and not published elsewhere, in whole or in part. 2. LAYOUT should be as follows: (a) Centred masthead to consist of Title: informative but concise, without abbreviations and not including the names of new genera or species Author’s(s’) name(s) Address(es) of author(s) (institution where work was carried out) Number of illustrations (figures, enumerated maps and tables, in this order) (b) Abstract of not more than 200 words, intelligible to the reader without reference to the text (c) Table of contents giving hierarchy of headings and subheadings (d) Introduction (e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents (f) Summary, if paper is lengthy (g) Acknowledgements (h) References (i) Abbreviations, where these are numerous 3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced with 2,5 cm margins all round. First lines of paragraphs should be indented. Tables and a list of legends for illustrations should be typed separately, their positions indicated in the text. All pages should be numbered consecutively. Major headings of the paper are centred capitals; first subheadings are shouldered small capitals; second subheadings are shouldered italics; third subheadings are indented, shouldered italics. Further subdivisions should be avoided, as also enumeration (never roman numerals) of headings and abbreviations. Footnotes should be avoided unless they are short and essential. Only generic and specific names should be underlined to indicate italics; all other marking up should be left to editor and publisher. 4. ILLUSTRATIONS should be reducible to a size not exceeding 12 « 18 cm (19 cm including legend); the reduction or enlargement required should be indicated; originals larger than 35 x 47 cm should not be submitted; photographs should be rectangular in shape and final size. 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REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: ‘Smith (1969) describes...’ ‘Smith (1969: 36, fig. 16) describes...’ “As described (Smith 1969a, 1969b; Jones 1971)’ ‘As described (Haughton & Broom 1927)...’ ‘As described (Haughton et al. 1927)...’ Note: no comma separating name and year Dagination indicated by colon, not p. names of joint authors connected by ampersand et al. in text for more than two joint authors, but names of all authors given in list of references. (b) Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, b, etc. to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a). For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list o, scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number (only if independently paged) in parentheses, pagination (first and last pages of article). Examples (note capitalization and punctuation) BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140. FiscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. gen. 74: 627-634. Konn, A. J. 1960a. Ecological notes on Carus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Ann, Mag. nat. Hist. (13) 2: 309-320. Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4 ): 51. THIELE, J. 1910. Mollusca: B. Boleeceehees? Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270 (continued inside back cover) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 74 Band | December 1977 Desember Part 2 Deel THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES PART 5 CRUSTACEA, DECAPODA, REPTANTIA AND NATANTIA By BRIAN KENSLEY Cape Town Kaapstad The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular intervals as material becomes available Obtainable from the South African Museum, P.O. Box 61, Cape Town Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM word uitgegee in dele op ongereelde tye na beskikbaarheid van stof Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad OUT OF PRINT/UIT DRUK 1, 21-3, 5-8), 3(1-2, 4-5, 8, t.—p.i.), 5(1-3, 5, 7-9), 6(1, t.—p.i.), 701-4), 8, 9(1-2, 7), 10(1-3), 11(1-2, 5, 7, t.-p.i.), 15(4-5), 24(2), 27, 311-3), 33 Price of this part/Prys van hierdie deel - R3,00 Trustees of the South African Museum © Trustees van die Suid-Afrikaanse Museum 1977 ISBN 0 908407 27 0 Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., . Die Rustica-pers, Edms., Bpk., Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES PART 5 CRUSTACEA, DECAPODA, REPTANTIA AND NATANTIA By BRIAN KENSLEY South African Museum, Cape Town (With 16 figures) LMS. accepted 10 May 1977] ABSTRACT Sixty-one species of Reptantian and Natantian decapod crustaceans from deep water off Natal are listed. Of the thirteen new records of Natantia dealt with from the area, three are described as new species. These are Benthesicymus expansus, Hymenopenaeus kannemeyeri and Pasiphaea meiringnaudei. CONTENTS PAGE Introduction : ‘ i hk Species List : : : ene AS Systematic Discussion . : 5 4N) Acknowledgements ck : SiS References . : : : « |) 43 INTRODUCTION The present paper forms part of a series based on material collected by the South African Museum during two cruises off the Natal coast, on the R/V Meiring Naude (see Fig. 1). For the scope of, and the background to, this programme, as well as for all station data, the reader is referred to Louw (1977). All the species of Reptantia and Natantia collected are listed systematically, but only new species, new records for South Africa, and a few additional species requiring further comment are dealt with in the systematic text. The following abbreviations are used throughout this paper: SAM—South African Museum catalogue number; SM—Meiring Naude station numbers; CB—carapace breadth; CL—carapace length; RL—rostral length; TL—total length; ovig.— ovigerous; juv.—juvenile. SPECIES LIST Where specimens are not identified to species level, these were either immature or damaged. ‘ * New record for South African waters. SM Station Ovig. no. 33 29 29 Juv. PALINURA Family Eryonidae Stereomastis sculpta (Smith) . A i : 58 — ~- 1 — (Barnard 1950: 572) 83 1 — — — (Eryoneicus caecus) ; 5 F ee LOT — — _ 1 13) Ann. S. Afr. Mus. 74 (2), 1977: 13-44, 16 figs. 14 ANNALS OF THE SOUTH AFRICAN MUSEUM SOG Ta PAP INGA ho72 50' A DURBAN Fig. 1. Map showing localities of collecting stations. 50° 28° 29 SI9453 Sd os % a) 5 oY) @ ve j 33° 10 20° THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES SM Station no. 3d eee Family Palinuridae Palinurus delagoae Barnard . : : : 22 I = (Berry & Plante 1973: 374) Projasus parkeri (Stebbing) . , ' : 38 — ox (George & Grindley 1964: 87) ASTACURA Family Astacidae Nephropsis atlantica Norman. : 38 3 D (Barnard 1950: 530) 66 3 — NATANTIA PENAEIDEA Family Penaeidae Subfamily Sicyoninae Sicyonia longicauda Rathbun . ‘ : 86 — , 1 (Barnard 1950: 635) Subfamily Aristeinae Aristaeomorpha foliacea (Risso). , ; fi 1 y) (Barnard 1950: 625) 22 2 4 28 1 - 38 3 3 44 — 2 66 — 1 83 1 _ 92 — 2 99 1 _ 100 3 3 * Aristeus semidentatus (Bate) . : ; ee Os 1 _ (see systematic section, p. 20) *Aristeus virilis (Bate) . ; : : : 58 2 — (see systematic section, p. 21) 91 1 — 107 ~- 1 *Benthesicymus expansus sp.nov. . : ee hO7 1 — (see systematic section, p. 22) *Benthesicymus investigatoris Alcock & Anderson 38 — | (see systematic section, p. 26) 58 — 1 66 -- 1 107 _ y) Gennadas bouvieri Kemp : : ; ; 5 = 1 (Kensley 19716: 273) 56 1 — 63 — 4 88 — i 105 — 1 111 -- 1 Gennadas capensis Calman. : ; : 99 — 1 (Kensley 19715: 277) 105 1 1 Gennadas clavicarpus De Man : : 5) 1 1 (Kensley 19715: 278) 63 — 1 88 — 1 96 1 1 105 — 1 111 3 — Gennadas gilchristi Calman . : 5 : 25 — 3 (Kensley 19715: 280) 33 2 4 49 -- 2) 55 3 4 — 16 ANNALS OF THE SOUTH AFRICAN MUSEUM SM Station Ovig. no. = +O | | Gennadas parvus Bate . : ; : F 25 (Kensley 19715: 287) ) 33 Gennadas scutatus Bouvier . : x 4 56 (Kensley 19715: 288) 71 Gennadas tinayrei Bouvier. : ; : 56 (Kensley 19715: 290) 63 | | Plesiopenaeus edwardsianus (Johnson) . ; 7 (Crosnier & Forest 1973: 292) 28 lswleeal | | | | | Plesiopenaeus ni‘idus Barnard ; ‘ , 7 (Barnard 1950: 622) 22 — me OY BRENANIUNERE XA | | ww Subfamily Solenocerinae Haliporoides triarthrus (Stebbing) : : 2) (Barnard 1950: 619) 92 * Hymenopenaeus kannemeyeri sp.nov. . ; 38 (see systematic section, p. 27) Hymenopenaeus sp. : dU 1 _ — = == an | | | THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES SM Station no. Subfamily Penaeinae *Funchalia villosa (Bouvier) . , (see systematic section, p. 29) Parapenaeopsis acclivirostris Alcock (Barnard 1950: 604) Penaeopsis rectacuta (Bate) (Hall 1962: 18) Penaeus indicus Edwards (Barnard 1950: 588) Family Sergestidae Sergestes (Sergestes) armatus Kroyer (Kensley 1971la: 232) Sergestes (Sergestes) atlanticus Edwards . (Kensley 1971la: 234) Sergestes (Sergestes) curvatus Crosnier & Forest (Crosnier & Forest 1973: 315) Sergestes (Sergestes) orientalis Hansen (Kensley 1971a: 238) Sergestes (Sergestes) sargassi Ortmann (Kensley 1971la: 241) Sergestes (Sergestes) pectinatus Sund (Kensley 1971a: 240) lwme leo! 3d Psi | w —_— | Sy eal op load ol Ovig. e2 juv. 18 ANNALS OF THE SOUTH AFRICAN MUSEUM SM Station ovig. no. +O +O +O +O a c < Sergestes (Sergia) laminatus Burkenroad . : 56 (Kensley 1971la: 251) 88 \O ~ Ox | RerRUmOBRWN OG NwUNueas | | Sergestes (Sergia) potens Burkenroad . : U (Kensley 1971la: 253) Sergestes (Sergia) prehensilis Bate . : 5 (Kensley 1971a: 253) Ne 00 — e || (oe) (oe) | Ne WN) DB KH OW — RK Pe Sergestes (Sergia) regalis Gordon . : F DS (Kensley 1971a: 256) 55 ore) SS loos fool Sergestes (Sergia) scintillans Burkenroad . : WS) (Kensley 1971a: 257) 88 Sergestes (Sergia) splendens Sund . : : 88 (Kensley 1971a: 260) .. ‘ : : 96 * Sergestes (Sergia) talismani Barnard . ; 55) (see systematic section, p. 31) 63 — — N b— | | — N | NATANTIA CARIDEA Family Oplophoridae * Acanthephyra armata Edwards i ; 38 4 (see systematic section, p. 31) 58 3 Acanthephyra eximia Smith . : : 2 3 — (Crosnier & Forest 1973: 34) 38 — —=re UN = COR WD | | * Acanthephyra indica Balss_ . 4 ; é 56 (see systematic section, p. 31) 88 THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES Acanthephyra quadrispinosa Kemp . (Barnard 1950: 668) Oplophorus gracilirostris Edwards . (Chace 1936: 30) Oplophorus spinicauda Edwards (Kensley 1969: 169) Oplophorus spinosus (Brullé) . (Crosnier & Forest 1973: 25) Systellaspis debilis (Edwards) (Crosnier & Forest 1973: 87) Family Nematocarcinidae Nematocarcinus sp. Family Pasiphaeidae * Eupasiphae gilesii Wood-Mason (see systematic section, p. 32) *Pasiphaea sivado (Risso) , (see systematic section, p. 32) *Pasiphaea meiringnaudei sp. nov. . (see systematic section, p. 34) Family Hippolytidae Hippolysmata tugelae Stebbing (Barnard 1950: 712) SM Station no. | A) WD YIN 9 UD CIS QW VIS fw | bt | oy) Uy Vea IS er | hes hoes] | | — CON = = = — | =e oon | oa | N 20 ANNALS OF THE SOUTH AFRICAN MUSEUM SM Station Ovig. no. Oy +0 +0 +0 +O et | < Family Pandalidae Heterocarpus dorsalis Bate. . ; 3 7 (see systematic section, p. 38) 28 Heterocarpus laevigatus Bate . ; : : 38 (see systematic section, p. 38) 66 co WwW Heterocarpus tricarinatus Alcock & Anderson . 22 (see systematic section, p. 38) Parapandalus richardi (Coutiére) . : s 56 Plesionika martia (Edwards) . : , ' 7 (Crosnier & Forest 1973: 212) 22 ereel) cytes 4 Samm csp) es S | | Family Glyphocrangonidae Glyphocrangon dentatus Barnard . ' ; 22 (see systematic section, p. 40) 28 ieee | | @& | ld *Glyphocrangon regalis Bate . : 3 : 7 (see systematic section, p. 40) 28 | sree | | | | oneeenl | Ne) Ne) | Family Crangonidae Pontophilus sculptus (Bell) . ; . ° 86 _ (Barnard 1950: 810) — | | SYSTEMATIC DISCUSSION (new species, new records, and selected other species) PENAEIDEA Family Penaeidae Aristeus semidentatus (Bate) Figs 2A, 3A Hemipenaeus semidentatus Bate, 1888: 305, pl. 49 (fig. 1). Aristaeus semidentatus: Wood-Mason & Alcock, 1891: 280. Alcock, 1901: 31. Alcock & McArdle, 1901: pl. 49 (fig. 3). Aristeus semidentatus: De Man, 1911: 29. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 21 Previous records Laccadive Islands; Cape Comorin; Bay of Bengal; off Banda; Kei Islands; Kermadec Islands. Material SAM-A15139 SM 107 12 gg CL + RL 26,8 mm x TL 77,0 mm to CL + RL 31,5 mm x 92,0 mm Remarks The genus Aristeus is characterized by the lack of an hepatic spine, an indistinct cervical groove, the third pereiopods possessing an epipodite but no podobranch, the fourth pereiopods lacking an epipodite, the exopodite of the second maxilliped longer and stronger than the exopodite of the third maxilliped. There is some variation in the rostral dentition of the present material, several specimens showing a small dorsal and ventral tooth close to the apex (Fig. 2A). (Burucovsky & Romensky 1972, also describe rostral variability in Aristeus varidens.) Aristeus virilis (Bate) Figs 2B, 3B Hemipenaeus virilis Bate, 1888: 303, pl. 44 (fig. 4). Hemipenaeus tomentosus Bate, 1888: 307, pl. 49 (figs 2-3). Aristaeus virilis: Wood-Mason & Alcock, 1891: 279. Alcock, 1901: 30: Aristeus virilis:; De Man, 1911: 27. Fig. 2. A. Aristeus semidentatus. B. Aristeus virilis. DY ANNALS OF THE SOUTH AFRICAN MUSEUM Previous records Andaman Sea; China Sea; Philippines; Bali Sea; New Hebrides. Material SAM-A15159 SM 58 2 3g CL 41,2 mm x TL 110,5 mm; 37,0 mm x + 95 mm (telson damaged) SAM-A1I5160 SM 91 1 ¢43,0 mm x 109,0 mm SAM-A1I5161 SM 107 1 ° (rostrum damaged) Remarks The present material shows the characteristic thickening of the distal part of the scaphocerite, but the males lack the pubescence of the integument. A twist in the base of the inner antennular flagellum is also present. The two species of Aristeus dealt with here may be distinguished by the differences in the petasmata (Fig. 3) and rostra (Fig. 2), and the pubescent carapace of some specimens of A. virilis. Benthesicymus expansus sp. nov. Figs 4—5 Description Male Carapace (Fig. 4A) integument soft, membranous. Rostrum an entire crest, apex missing in holotype, ventral border bearing slight ridge carrying setae. No A Fig. 3. A. Aristeus semidentatus petasma. B. Aristeus virilis petasma. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 23 antennal spine; orbitoantennal groove extended posteroventrally to branchial region; rounded ridge situated above orbitoantennal groove; branchiostegal spine small, supported by short carina, no hepatic spine; cervical groove strong, almost reaching orbitoantennal groove; suprabranchial ridge fairly strong, rounded. Thoracic sternite between fourth pereiopods bearing short narrowly rounded, forwardly directed lobe. Posterior half of third pleon segment, and pleon segments four to six with mediodorsal carina, none bearing posterior spine. Telson carrying four pairs of lateral spines on distal third, apex acute (Fig. 4B). Antennal scaphocerite (Fig. 5F) blade two and one-third times longer than wide, outer margin reinforced by tapering ridge, ending in small spine some distance from apex of scale; basal segment carrying small dorsal hook-like process. Antennular peduncle segments setose. Ocular tubercle present on inner margin of proximal half of eye-stalk. Mandibular palp two-segmented, basal segment one and a half times longer and considerably broader than distal segment; mandible consisting of strong, straight cutting edge and three small teeth on inner face (Fig. 5A). First maxilliped (Fig. 5B) exopod narrowing abruptly to segmented distal portion. Second maxilliped (Fig. 5E) with merus very expanded, distally rounded, inner margin setose. Third maxilliped longer than first perelopod (Fig. 4A), merus and ischium expanded, fringed with setae on ventral margin; propodus and carpus relatively slender, of equal length, dactylus broadest at midpoint, tapering distally. First pereiopod chela relatively slender, finger and thumb slightly longer than palm; carpus slender; merus and ischium expanded. Second pereiopod (Fig. 4A) longer than first, very slender, finger and thumb of chela equal in length to palm; carpus and merus of equal length. Third to fifth pereiopods missing in holotype. Petasma slender (Fig. 4C), divided distally into median narrow lobe con- nected to truncate, rounded portion, and outer smoothly convex lobe bearing spinules. Inner scale of appendix masculina narrow, elongate, apically rounded, bearing short setae; outer scale broad, longer than inner. Material Holotype SAM-A15140 SM 107 1 ¢ CL 50,5 mm TL 150 mm Remarks The lack of mediodorsal pleonal spines, the ocular tubercle being proximal in position, the lack of an hepatic spine, and the presence of a mediodorsal keel on pleonal segments 3-6, place this specimen close to B. iridescens Bate (see Burkenroad 1936: 45; Roberts & Pequegnat 1970: 42). The latter authors note the difficulty of sorting out the ‘Brasiliensis’ group of species of Benthesicymus, due mainly to a lack of information on intraspecific variation. In spite of this 24 ANNALS OF THE SOUTH AFRICAN MUSEUM Z Yi YY YY A Vy Fig. 4. Benthesicymus expansus sp. nov. A. Holotype, carapace in lateral view. B. Telson. C. Anterior and posterior views of petasma. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 25 limitation, the present new species is described, albeit from a single specimen. The species has two features which easily separate it from B. iridescens. These are the non-dentate rostrum, and the expanded meri and ischia of the third maxilliped and first pereiopods. (The specific name is derived from this latter feature.) In addition, differences from B. iridescens are seen in the petasma, the appendix masculina, the telson, the lack of an antennal spine, and in the carapace sculpture. ( May HA’ AK WS S f iy / Y tt G y 4 Z Z ; & 4 yy Y Y, Y Y Y Y j Y q Y Y Y Z ANS MMA Fig. 5. Benthesicymus expansus sp. nov. A. Mandible. B. First maxilla. C. Second maxilla. D. First maxilliped. E. Second maxilliped. F. Antennal scaphocerite. 26 ANNALS OF THE SOUTH AFRICAN MUSEUM Benthesicymus investigatoris Alcock & Anderson Fig. 6 Benthesicymus investigatoris Alcock & Anderson, 1899: 282. Alcock, 1899: pl. 41 (fig. 2); 1901: 44. Burkenroad, 1936: 25. Previous records Saya de Malha (mid Indian Ocean); Andaman Sea; Gulf of Manar; Halmahera Sea; Ceram Sea; Hawaii. Material 5 292 CL + RL 24,0 mm x TL 76 to 31,0 mm x 89,5 mm Fig. 6. Benthesicymus investigatoris. A. Carapace in lateral view. B. Thelycum. THE SOUTH AFRICAN MUSEUM’S ME/RING NAUDE CRUISES pe Hymenopenaeus kannemeyeri sp. nov. Fig. 7 Description Carapace (Fig. 7A) firm, glabrous. Rostrum about one- Ania length of carapace directed obliquely upward at angle of about 15°, with eight dorsal teeth (three postorbital) in uninterrupted series, apical tooth very small, ventral margin bearing fringe of setae. Carapace with strong antennal and postantennal spine, each supported by stout carina; hepatic spine lacking keel; branchiostegal spine marginal, supported by keel extending posteriorly to hind margin of cara- pace; cervical sulcus well developed, with posterior margin of groove at midpoint marked by a short keel, joining the hepatic groove ventrally. Hepatic groove marked posteroventrally by short keel. Branchiocardiac sulcus well developed with supra-branchial portion having a straight, rounded ridge ventrally. First pleon segment dorsally rounded, segment two with posterior half keeled, third to sixth segments fully keeled, segments four to six with short posterior spine. Telson (Fig. 7D) with two fixed spines subterminally, with strong dorsolateral keel extending from spines to proximal margin; outer uropodal ramus just reaching telsonic apex. Eye reaching slightly beyond midpoint of rostrum, tubercle at midpoint of medial eye-stalk margin (Fig. 7E). Prosartema of antennule short and rounded, carrying long setae; basal peduncular segment with strong distal spine; stylocerite a strong spine; terminal peduncular segment with lateral setose ridge and thick setose patch dorsally; flagellum circular in cross-section. Scaphocerite extending well beyond antennular peduncle. Mandibular palp conspicuous, setose; distal segment slender, tapering, shorter than proximal segment, latter broadly oval. Third maxilliped extending slightly beyond scaphocerite. First pereiopod with strong distoventral spine on ischium and basis. Rudimentary exopods on first to fifth pereiopods and third maxilliped; epipods on second and third maxillipeds and first to fourth pereiopods. Inner margin of fifth perelopod basis in male with anteriorly directed triangular lobe above genital aperture. Thelycum of female (Fig. 7C) relatively simple, sternites of fourth and fifth pereiopods with rounded, convex central areas. Petasma divided distally into two lobes; inner lobe longer than outer, truncate, with distal hollowed portion produced laterally into triangular point; outer lobe also truncate, with posterior strengthening ridge. Appendix masculina of second pleopod (Fig. 7F) about half length of appendix interna; both bearing marginal spinules; leaf-like expansion present on outer side of base of endopod. Material Holotype SAM-A15141 SM 38 3 CL + RL 43,5 mm TL 119,5 mm Allotype SAM-A15141 SM 38 9 CL + RL 39,8 mm TL 110,8 mm. 28 ANNALS OF THE SOUTH AFRICAN MUSEUM Remarks The present specimens with the rostral and post-rostral teeth not separated, with a strong branchiostegal spine, and lacking a pterygostomian spine, fall into Burkenroad’s (1936) Group II. This group includes H. robustus, H. modestus, H. lucassi and H. muelleri. H. kannemeyeri differs from all these in the structure of the petasma and thelycum. The latter most closely resembles that of H. fattahi Ramadan, 1938 (as figured by Crosnier & Forest 1973, fig. 85g—h) but the petasma of H. fattahi differs markedly from the present species. Fig. 7. Hymenopenaeus kannemeyeri sp. nov. A. Carapace in lateral view. B. Anterior and posterior view of petasma. C. Thelycum. D. Telson. E. Eye. F. Pleopod 2, 3, appendix interna and appendix masculina. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 29 The species is named for S. X. Kannemeyer, of the Department of Marine Biology, South African Museum, in appreciation for his invaluable help in the Meiring Naude cruises. Genus Funchalia Johnson As Funchalia villosa and F. woodwardi have been confused in the past, the petasma and thelycum of each is figured, and all the South African Museum’s holdings of both species are listed. Funchalia (Funchalia) villosa (Bouvier) Fig. 8C-D Funchalia (Funchalia) villosa: Burkenroad, 1936: 129. Crosnier & Forest, 1973: 296. Previous records Eastern and western North Atlantic; Caribbean; southern central South Atlantic. Material and South African records SAM-A15368 SM 18 12 CL 22,8 mm SAM-A15369 SM 33 12 CL 21,0 mm SAM-A15370 SM 88 13192CL 13,0 mm, 14,2 mm SAM-A15371 SM 111 13 CL 18,4 mm SAM-A13195_ off Natal, 30°30’S 31°45’E 200 m 1 3 SAM-A13197_ off Natal, 26°30’S 42°40’E 500 m 1 d SAM-A13198 off Agulhas Bank, 37°10’S 21°50’E 500 m1 4d SAM-A13236 off Natal, 31°44’S 44°35’E 500 m 3 gd Funchalia (Funchalia) woodwardi Johnson Fig. 8A-B Funchalia (Funchalia) woodwardi: Barnard, 1950: 609, fig. 112. Gordon & Ingle, 1956: 478 Paulinose, 1974: 433. Previous records Eastern North Atlantic; Mediterranean; south-eastern South Atlantic. South African records SAM-A8481_ off Table Bay, from stomach of stock fish 9 ¢3 SAM-A10914 off Cape Peninsula, from tunny stomach several dg & 29 SAM-A10995_ off Cape Peninsula, from tunny stomach | SAM-A11025 off Dassen Island, from tunny stomach 25 g¢ 19 9° SAM-A11027 off Cape Peninsula, several g¢ & 99 SAM-A12014 off Dassen Island, 500 m 2 ¢¢ 2 99 SAM-A12482 Vema Seamount, from tunny stomach 60 immature 3g & 2° SAM-A13196 off Cape Peninsula, 250 m 3 99 SAM-A13237 off Cape Peninsula, 350 m7 gg 7 22 SAM-A13238 off Cape Peninsula, 500 m 2 g¢ 3 99 30 ANNALS OF THE SOUTH AFRICAN MUSEUM B Fig. 8. A. Funchalia woodwardi petasma. B. F. woodwardi thelycum. C. Funchalia villosa petasma. D. F. villosa thelycum. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES Sil Family Sergestidae Sergestes (Sergia) talismani Barnard Sergestes talismani Barnard, 1947: 384. Sergestes (Sergia) talismani:; Crosnier & Forest, 1973: 325, figs 11la—c, 112a—b. Previous records Cape Verde Islands; Portuguese Guinea; Gabon; Congo. Material See species list. Remarks This would seem to be the first record of the species from the Indian Ocean. CARIDEA Family Oplophoridae Acanthephyra armata Edwards Fig. 9B Acanthephyra armata: Bate, 1888: 744, pl. 125 (fig. 2). De Man, 1920: 61, pl. 6 (fig. 13). Chace, 1936: 27. Previous records West Indies; Bali Sea; Straits of Macassar; off Kei Islands; off Banda Island. Material SAM-A15154 SM 38 434499 SAM-A15150 SM 58 3 3¢3 99 gd CL 16,9 mm-29,0 mm RL 27,0 mm—37,5 mm TL 92,0 mm—137 mm oe) 24,5 mm-—33,0 mm 34.7 mm-—35,2 mm 121 mm—-146 mm Acanthephyra indica Balss Fig. 9A Acanthephyra indica Balss, 1925: 264, fig. 34. Chace, 1936: 26. Acanthephyra sp. De Man, 1920: 68, pl. 6 (fig. 16). Previous records Between Sumatra and Ceylon; Banda Sea; off Ambon. Material See species list. 3 gg 123 ovig. 22 gG¢CL10,0 mm-10,.2mm 2 CL 10,9 mm-13,1 mm 32 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 9. A. Acanthephyra indica. B. Acanthephyra armata. Family Pasiphaeidae Eupasiphae gilesii (Wood-Mason & Alcock) Fig. 10B Parapasiphae (Eupasiphae) gilesii Wood-Mason & Alcock, 1893: 166. Alcock, 1892: pl. 3 (fig. 8); 1901: 66. Eupasiphae gilesii: Holthuis, 1955: 36. Tirmizi, 1969: 214, figs 1-4. Foxton, 1970: 958. Eupasiphae gilesi: Fisher & Goldie, 1961: 78. Crosnier & Forest, 1973: 150, fig. 44. Previous records Bermuda; Cape Verde Islands; Canary Islands; Madeira; Arabian Sea; Gulf of Oman; Andaman Sea. Material SAM-A15151 SM 105 1g CL + RL 32,4 mm TL 97 mm Pasiphaea sivado (Risso) Fig. 10A Pasiphae sivado: Wood-Mason & Alcock, 1893: 161. Alcock, 1892: pl. 3 (fig. 6); 1901: 59. Sivertsen & Holthuis, 1956: 29. Crosnier & Forest, 1973: 133. 33 THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES Fig. 10. A. Pasiphae sivado. B. Eupasiphae gilesi, 34 ANNALS OF THE SOUTH AFRICAN MUSEUM Previous records Eastern North Atlantic; Mediterranean; Red Sea; Bay of Bengal; Andaman Sea. Material SAM-15153 SM 63 2 99 SAM-A15152 SM 85 12° 2° CL 10,0 mm-14,5 mm Pasiphaea meiringnaudei sp. nov. Figs 11-14 Description Carapace (Fig. 11) mid-dorsally carinate for almost its entire length. Rostrum a postfrontal triangular spine directed obliquely upward, only rarely reaching beyond anterior carapace margin. Anterior margin of rostrum straight, almost vertical, apparently more variable in females than in males (Fig. 13). Orbital angle a rounded lobe; branchiostegal spine directed outwards; supra- branchial ridge rounded, not reaching posterior carapace margin. Second to sixth abdominal segments dorsally keeled, not ending in a posterior spine; sixth segment with curved lateral ridge. Telson shorter than Fig. 11. Pasiphae meiringnaudei sp. nov. Holotype in lateral view, THE SOUTH AFRICAN MUSEUM’S MEIJRING NAUDE CRUISES 35 uropodal endopod, dorsally grooved, distally forked, with seven spines on inner margin of sinus (Fig. 121). Antennular peduncle extending for two-thirds of scaphocerite (Fig. 11); stylocerite (Fig. 12G) broadening distally, twisted, apically acute, dorsolateral flagellum with about eighteen enlarged basal segments in male and female. | Antennal base bearing ventrodistal spine. Fig. 12. Pasiphae meiringnaudei sp. nov. A, Anterior carapace and rostrum. B. Mandible. C. First maxilla. D. First maxilliped. E. Second maxilliped. F. Second maxilla. G. Antennular base. H. Antennal base. I. Telsonic apex. J. Pleopod 2, ¢, appendix interna and appendix masculina. K. Pleopod 1, 3. L. Pleopod 1, 3, endopod. 36 ANNALS OF THE SOUTH AFRICAN MUSEUM Third maxilliped just reaching to end of scaphocerite. First perelopod overreaching scaphocerite by half length of palm plus fingers (Fig. 11); palm longer than fingers; merus with one to four ventral spines; basis with ventral distal triangular spine. Second pereiopod overreaching scaphocerite by half length of palm plus fingers; palm equal in length to fingers; carpus with broad ventral tooth; merus with fifteen to twenty-one ventral spines; ischium with one spine, basis with four to seven spines (Fig. 14B). Third pereiopod overreaching anterior carapace margin by about half length of propodus plus dactylus; very slender; dactylus tapering; propodus one- third length of dactylus; carpus very elongate, almost four times length of merus. Fourth pereiopod reaching to midpoint of carpus of third pereiopod. Fifth pereiopod longer than fourth pereiopod, reaching to end of carpus of third pereiopod. Endopod of first pleopod ¢ short, with triangular process on inner margin bearing very small denticles (Fig. 12L). Second pleopod 3 appendix masculina twice length of appendix interna, latter curved, with seven marginal setae (Fig. 12J). Uropodal endopod two-thirds length of exopod, latter with small terminal spine on outer margin. Material | Holotype SAM-A15316 SM 107 ovig. 9 CL 57,0 mm TL 166 mm SM 38 134 CL 39,9 mm TL 126 mm SM 58 4 493 CL 35,5 mm-36,3 mm TL 109 mm—-110 mm SM 107 5 ovig. 992 50,0 mm—58,0 mm 153 mm—-168 mm SM 91 192 42,0 mm 125 mm SM 28 12 34,5 mm 108 mm SM 40 2 99 31,8 mm—42,4 mm 95-130 mm Fig. 13. Pasiphae meiringnaudei sp. nov. rostral variation. Top row, ovigerous 22. Bottom row, gd. Bottom right, rostrum modified by ellobiopsid parasites. EE a THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 3) Remarks J. C. Yaldwyn (National Museum of New Zealand) has kindly supplied most of the following information on the closely related species of Pasiphaea, from an as yet unpublished review of the genus by him and F. A. Chace Jr (Smithsonian Institution). The combination of three characters, viz. the distally forked telson, carinate carapace and abdomen, and basis of the second pereiopod ventrally spined, places the present species in a group with four previously described species. These are P. multidentata Esmark, P.: tarda Kroyer, P. rath- bunae Stebbing, and P. barnardi Yaldwyn. The rostral shape of P. meiringnaudei differs from these four, but comes closest to P. barnardi. The finger-to-palm ratio of the first and second pereiopods, however, differs from Yaldwyn’s species where the fingers of the first pereiopod are subequal to the palm and the fingers of the second pereiopod are distinctly longer than the palm. P. meiringnaudei overlaps with P. tarda (as P. princeps in Kemp 1910, pl. 4 (figs 1—-7)) in the spine counts of the first and second pereiopods, but differs markedly in the rostral shape and in the relative lengths of the thoracic appendages. The species is named for the R/V Meiring Naude. Fig. 14. Pasiphae meiringnaudei sp. nov. A. First pereiopod, with chela further enlarged. B. Second pereiopod, with chela further enlarged. 38 ANNALS OF THE SOUTH AFRICAN MUSEUM Family Pandalidae Heterocarpus dorsalis Bate Fig. ISA Heterocarpus dorsalis: De Man, 1920: 156, 171, pl. 15 (fig. 43). Barnard, 1950: 684, fig. 127a Previous records Off Durban; off East Africa; East Indies; Japan. Material See species list. 7 33 CL 20 mm—41 mm RL 35 mm-48 mm TL 98 mm-179 mm IQD 27 mm—42 mm 35 mm-44 mm~—~—s«126 mm-—171 mm Remarks The ovigerous female of total length 171 mm is at least 20 mm longer than the largest ovigerous female recorded by De Man (1920). Heterocarpus laevigatus Bate Fig. 15B Heterocarpus laevigatus Bate, 1888: 636, pl. 112 (fig. 3). Stebbing, 1914: 40. De Man, 1920: 154, 159, pl. 13 (fig. 37). Barnard, 1950: 684, fig. 127b. Crosnier & Forest, 1973: 195, fig. 6lc. ; Previous records Cape Verde Islands; Madeira; off East London; Arabian Sea; East Indies; Hawaii. Material See species list. 233 CL 65 mm-74 mm RL 28 mm-38 mm TL 121 mm—160 mm 39° 64 mm—-105 mm 36 mm—5!1 mm 121 mm—200 mm Remarks The East London record was of an immature specimen measuring approxi- mately 63 mm TL. Heterocarpus tricarinatus Alcock & Anderson Fig. 15C Heterocarpus tricarinatus: De Man, 1920: 155, 161, pl. 13 (fig. 38), pl. 14 (fig. 38). Calman, 1939: 204. Barnard, 1950: 682, fig. 127c—d. Previous records Off East London; Arabian Sea; East Indies. Material SAM-A15264 SM 22 | 3 CL 35mm RL 30 mm TL 135 mm ovig. 9 37 mm 32 mm 146 mm THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 39 a Wal 0. 10 2:0 #30 £0 55 ey a On. 10 2 30. «660 50 B pg, we “ilitiyyyy, tikiy tig a Vi On 10 20. 30 210 50 Aj Cc Fig. 15. A. Heterocarpus dorsalis. B. Heterocarpus laevigatus. C. Heterocarpus tricarinatus. 40 ANNALS OF THE SOUTH AFRICAN MUSEUM Remarks Using both Barnard’s (1950) and De Man’s (1920) keys to the genus Heterocarpus, H. tricarinatus is arrived at. The genus is obviously variable with regard to the rostral dentition, proportions of the pereiopodal dactyli, and the lengths of the carapace keels. This led Calman (1939) to doubt the validity of this species, suggesting that it might be synonymous with H. gibbosus Bate. The variability of this latter species is well illustrated by De Man (1920). The present material could well be placed in Bate’s species, but until this species complex is resolved, the lower carapace keel reaching back about two-thirds of the cara- pace length necessitates placing these two specimens in H. tricarinatus. Family Glyphocrangonidae Glyphocrangon dentatus Barnard Glyphocrangon gilesii var. dentata Barnard, 1926: 128. Glyphocrangon mabihissae Calman, 1939: 217, fig. 8. Glyphocrangon dentarus Barnard, 1947 (typ. err.): 387. Glyphocrangon dentatus Barnard, 1950: 722, fig. 134h. Previous records Off Mozambique, 540 m; off Zanzibar, 640-658 m. Material 8 dd 1 2 8 ovig. 29, see species list. So CL 16,5 mm-19,8 mm RL 17,8 mm-22 mm TL 73 mm-85 mm ovig. 2° 19,1 mm—23 mm 20,8 mm-27,5 mm 83 mm—-102 mm Remarks Only slight differences are apparent between Calman’s specimens from Zanzibar and the present material. These differences are to some extent probably due to individual variation, as well as to variations between populations. The spine on the midpoint of the posterior antennal carina is present in some of the present specimens, absent in others, while the row of small tubercles just lateral to the submedian carina seen in the Zanzibar specimens is represented by one or two almost indistinct tubercles. Although the rostral length always exceeds the carapace length, the ratio is variable. Glyphocrangon regalis Bate Fig. 16 Glyphocrangon regalis Bate, 1888: 517, pl. 93 (figs 3-4). De Man, 1920: 220. Description Rostrum curved ventrally, with apex upturned, slightly less than two-thirds carapace length, with pair of lateral rostral teeth above eyes, second pair of spines in postorbital position. Antennal spine strong, directed forward, separated THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 4] by short. curved carapace margin from longer branchiostegal spine directed anteroventrally. Anterior submedian carinae consisting of seven elongate tubercles. Posterior submedian carinae consisting of four elongate tubercles. Row of small, rounded tubercles median to both anterior and posterior sub- median carinae. Small median spine at base of rostrum flanked by two postero- lateral spines. Anterior antennal carina forming wing-like flattened structure ending in outward-pointed strong spine. Posterior antennal carina situated some distance above anterior antennal carina, ending anteriorly in outwardly directed spine, not as strong as that of anterior carina; at about midpoint, a slight indica- tion of a notch, never strong enough to be called a spine or tubercle. Anterior intermediate carina not well defined, consisting of three or four large irregularly placed tubercles. Posterior intermediate carina consisting of four or five elongate tubercles. Area between anterior submedian and anterior intermediate carinae with two rows of small tubercles; three rows of tubercles between posterior submedian and posterior intermediate carinae. Area between anterior antennal carina and lateral groove carrying a scattering of small rounded tubercles. Areas between posterior antennal and posterior lateral carinae also carrying irregular Fig. 16. Glyphocrangon regalis. 42 ANNALS OF THE SOUTH AFRICAN MUSEUM scattering of rounded tubercles. Anterior and posterior lateral carina entire, not tuberculate. Sublateral carina only distinct in posterior region, consisting of a foveolate ridge. Region between sublateral carina and margin bearing a foveolate band. Abdomen (excluding telson) equal in length of carapace plus rostrum. Median carina of abdominal segments strong, only on first segment forming a forwardly directed spine. Pleura of first segment considerably shorter than second, with forwardly directed spine. Pleura of second segment ventrally bispinose, spines of equal strength. Pleura of third segment ventrally bispinose, anterior spine longer than posterior. Pleura of fourth segment ventrally bispinose, spines subequal. Pleura of fifth segment bispinose, spines directed posteroventrally. Sixth abdominal segment ending posteroventrally in strong spine. Telson with median spine at base, elongate triangular, with strong dorsolateral keel, apex acute, flexed slightly dorsally. Eyes well developed, with large black corneas, small spinose tubercle on inner distal margin of eye-stalk. Scaphocerite ovate, fringed with setae, lacking any spine on inner margin. Second pereiopods unequal, left shorter and slightly squatter than right, carpus of right (30-31 segments) considerably longer than left (19 segments), chela markedly smaller. Third pereiopod dactylus lanceolate, oval in cross section, one-third length of propodus. Fourth pereiopod dactylus slightly more than one-third length of propodus, dorsally grooved. Fifth pereiopod dactylus similar to fourth. Previous records Banda Island; off Makassar; Kermadec Island; Fiji; Paternoster Island. Material 5 33 7 98 10 ovig. 29, see species list. $3o CL 20,5 mm—-29,9 mm RL 16,5 mm-19,9 mm TL 82,5 mm—108 mm ovig. 2° 29,3 mm—34,9 mm 17,8 mm—21,0 mm 108 mm—-128 mm Remarks The species recorded by Wood-Mason (1891) as G. investigatoris and G. investigatoris var. andamanensis, was stated by De Man (1920) to be possibly synonymous with G. regalis. Wood-Mason’s species, however, shows some differences, e.g. in the degree of ‘flare’ of the spine of the postantennal carina, the possession of midline tubercles on the rostrum which together suggest that this is not the same species. Examination of the ‘Challenger’ types of G. regalis reveal almost no differences from the present material. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 43 ACKNOWLEDGEMENTS In addition to the people and institutions thanked in the introductory paper of this series, I should like to thank Mr S. X. Kannemeyer of the South African Museum for the photographs, Dr R. B. Manning of the Smithsonian Institution for the loan of Pasiphaea material, Dr J. Yaldwyn of the National Museum of New Zealand and Dr F. A. Chace Jr of the Smithsonian Institution for informa- tion on Pasiphaea, and Prof. J. H. Day of the University of Cape Town and Dr J. Yaldwyn for reading the manuscript and for their many useful criticisms. REFERENCES Atcock, A. 1892. Crustacea Part I. ///ustrations of the Zoology of the Investigator Plates 1-S. Calcutta: Superintendent of Government Printing. Atcock, A. 1899. Crustacea Part VII. ///lustrations of the Zoology of the Investigator Plates 36-45. Calcutta: Superintendent of Government Printing. Awcock, A. 1901. A descriptive catalogue of the Indian deep-sea Crustacea Decapoda Macrura and Anomala, in the Indian Museum. Calcutta: Trustees of the Indian Museum. ALcock, A. & ANDERSON, A. R. S. 1899. Natural history notes from H.M. Royal Indian Marine Survey Ship ‘Investigator’, Commander T. H. Heming, R.N., commanding. Series III, no. 2. An account of the deep-sea Crustacea dredged during the surveying season 1897-98. Ann. Mag. nat. Hist. (7) 3: 278-282. AuLcock, A. & McARDLE, A. F. 1901. Crustacea Part IX. //lustrations of the Zoology of the Investigator Plates 49-55. Calcutta: Superintendent of Government Printing. BALss, H. 1925. Macrura der Deutschen Tiefsee-Expedition. 2. Natantia, Teil A. Wiss. Ergebn. dt. Tiefsee-Exped. ‘Valdivia’ 20: 221-315. BARNARD, K. H. 1926. Report on a collection of Crustacea from Portuguese East Africa. Trans. R. Soc. S. Afr. 13: 119-130. BARNARD, K. H. 1947. Descriptions of new species of South African decapod Crustacea, with notes on synonymy and new records. Ann. Mag. nat. Hist. (11) 13: 361-392. BARNARD, K. H. 1950. Descriptive catalogue of South African decapod Crustacea (crabs and shrimps). Ann. S. Afr. Mus. 38: 1-837. Berry, P. F. & PLANTE, R. 1973. Revision of the spiny lobster genus Palinurus in the south- west Indian Ocean. Trans. R. Soc. S. Afr. 40: 373-380. BATE, C. S. 1888. Report on the Crustacea Macrura dredged by HMS Challenger during the years 1873-1876. Rep. Voy. Challenger, 1873-1876 24: 1-942. BURKENROAD, M. D. 1936. The Aristaeinae, Solenocerinae and pelagic Penaeinae of the Bingham Oceanographic Collection. Bull. Bingham oceanogr. Coll. 5 (2): 1-151. BURUCOVSKY, R. N. & ROMENSKY, L. L. 1972. On rostrum changeability of Aristeus varidens Holthuis (Decapoda, Penaeidae). At/. Nauchno-Issled. Inst. Rybn. Khoz. Okeanogr. 42: 156-161. CALMAN, W. T. 1939. Crustacea: Caridea. Scient. Reps. John Murray Exped. 6: 183-223. CHACE, F. A. 1936. Revision of the bathypelagic prawns of the family Acanthephyridae, with notes on a new family, Gomphonotidae. J. Wash. Acad. Sci. 26: 24-31. CROSNIER, A. & Forest, J. 1973. Faune Tropicale. XIX. Les Crevettes profondes de I’ Atlantique Oriental Tropical. Paris: O.R.S.T.O.M. FisHER, L. R. & Goupig, E. H. 1961. New records of two deep-sea decapods. Crustaceana 2: 78-79. Foxton, P. 1970. The vertical distribution of pelagic decapods (Crustacea: Natantia) collected on the SOND cruise 1965. I. The Caridea. J. mar. biol. Assoc. U.K. 50: 939-960. GEORGE, R. W. & GRINDLEY, J. R. 1964. Projasus—a new generic name for Parker’s crayfish, Jasus parkeri Stebbing (Palinuridae: ‘Silentes’). J. Proc. R. Soc. West. Aust. 47: 87-90. Gorpon, I. & INGLE, R. W. 1956. On a pelagic penaeid prawn, Funchalia woodwardi Johnson, new to the British fauna. J. mar. biol. Assoc. U.K. 35: 475-481. HALL, D. N. F. 1962. Observations on the taxonomy and biology of some Indo-West Pacific Penaeidae. Col. Off. Fish. Publs 17: 1-230. 44 ANNALS OF THE SOUTH AFRICAN MUSEUM Ho.tuuts, L. B. 1955. The recent genera of the Caridean and Stenopodidean shrimps (Class Crustacea, Order Decapoda, Supersection Natantia) with keys for their determination. Zool. Verhandl. 26: 1-157. Kemp, S. 1910. The Decapoda Natantia of the coasts of Ireland. Scient. Invest. Fish. Brch Ire. 1908 (1): 1-190. KENSLEY, B. F. 1969. Decapod Crustacea from the south-west Indian Ocean. Ann. S. Afr. Mus. 52: 149-181. KENSLEY, B. F. 1971a. The Family Sergestidae in the waters around southern Africa (Crustacea, Decapoda, Natantia). Ann. S. Afr. Mus. 57: 215-264. KENSLEY, B. F. 1971b. The genus Gennadas in the waters around southern Africa. Ann. S. Afr. Mus. 57: 271-294. Louw, E. M. 1977. The South African Museum’s Meiring Naude cruises. Part 1. Station data 1975, 1976. Ann. S. Afr. Mus. 72: 147-159. Man, J. G. DE. 1911. The Decapoda of the Siboga Expedition. Part I. Family Penaeidae. Siboga Exped. monogr. 39A: 1-131. Man, J. G. DE. 1920. The Decapoda of the Siboga Expedition. Part IV. Families Pasiphaeidae, Stylodactylidae, Hoplophoridae, Nematocarcinidae, Thalassocaridae, Pandalidae, Psalido- podidae, Gnathophyllidae, Processidae, Glyphocrangonidae, and Crangonidae. Siboga Exped. monogr. 39A3: 1-318. PAULINOSE, V. T. 1974. Decapod Crustacea from the International Indian Ocean Expedition: The species of Funchalia (Penaeidae) and their post-larvae. J. nat. Hist. 8: 433-443. RAMADAN, M. M. 1938. Crustacea, Penaeidae. Scient. Reps John Murray Exped. 5: 35-76. RosBertTs, T. W. & PEQUEGNAT, W. E. 1970. Deep-water decapod shrimps of the Family Penaeidae. Jn: Pequegnat, W. E. & Chace, F. A. Texas A & M University Oceanographical Studies 1. Contributions on the Biology of the Gulf of Mexico. Houston, Texas: Gulf Publishing Co. SIVERTSEN, E. & HOLTHUuIS, L. B. 1956. Crustacea Decapoda (the Penaeidae and Stenopodidae excepted). Rep. scient. Results Michael Sars N. Atlant. Deep Sea Exped. 5 (12): 1-54. STEBBING, T. R. R. 1914. South African Crustacea. Ann. S. Afr. Mus. 15: 1-55. TirMiIZzI, N. 1969. Eupasiphae gilesii (Wood-Mason, 1892) from the northern Arabian Sea (Decapoda, Caridea). Crustaceana 16: 213-218. Woop-Mason, J. & ALCOcK, A. 1891. On the results of deep-sea dredging during the season 1890-91. Ann. Mag. nat. Hist. (6) 8: 268-286. Woop-Mason, J. & ALcock, A. 1893. On the results of deep-sea dredging during the season 1890-91. Ann. Mag. nat. Hist. (6) 11: 161-171. ~~ 6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature (particularly Articles 22 and 51). Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov., etc. An author’s name when cited must follow the name of the taxon without intervening punctuation and not be abbreviated; if the year is added, a comma must separate author’s name and year. The author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is transferred from its original genus. The name of a subsequent user of a scientific name must be separated from the scientific name by a colon. Synonymy arrangement should be according to chronology of names, 1.e. all published scientific names by which the species previously has been designated are listed in chronological order, with all references to that name following in chronological order, e.g.: Family Nuculanidae Nuculana (Lembulus) bicuspidata (Gould, 1845) Figs 14-15A Nucula (Leda) bicuspidata Gould, 1845: 37. Leda plicifera A. Adams, 1856: 50. Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b). Nucula largillierti Philippi, 1861: 87. Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. Note punctuation in the above example: comma separates author’s name and year semicolon separates more than one reference by the same author full stop separates references by different authors figures of plates are enclosed in parentheses to distinguish them from text-figures dash, not comma, separates consecutive numbers Synonymy arrangement according to chronology of bibliographic references, whereby the year is placed in front of each entry, and the synonym repeated in full for each entry, is ‘not acceptable. In describing new species, one specimen must be designated as the holotype; other speci- mens mentioned in the original description are to be designated paratypes; additional material not regarded as paratypes should be listed separately. The complete data (registration number, depository, description of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.: Holotype SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973. Note standard form of writing South African Museum registration numbers and date. 7. SPECIAL HOUSE RULES Capital initial letters (a) The Figures, Maps and Tables of the paper when referred to in the text e.g. °... the Figure depicting C. namacolus ...’; ‘. .. in C. namacolus (Fig. 10)...’ (b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded by initials or full names e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene (c) Scientific names, but not their vernacular derivatives e.g. Therocephalia, but therocephalian Punctuation should be loose, omitting all not strictly necessary Reference to the author should be expressed in the third person Roman numerals should be converted to arabic, except when forming part of the title of a book or article, such as “Revision of the Crustacea. Part VIII. The Amphipoda.’ Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial capital letter, provided the same generic name is used consecutively. Name of new genus or species is not to be included in the title: it should be included in the abstract, counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts. BRIAN KENSLEY THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES PART 5 CRUSTACEA, DECAPODA, REPTANTIA AND NATANTIA ’ ale, = VOLUME 74 PART 3 NOVEMBER 1977 ISSN 0303-2515 MUS. COMP. Z LIBRARY JAN 3 1 1978 HARVARD UNIVERSITY. OF THE SOUTH AFRICAN ~ MUSEUM CAPE TOWN INSTRUCTIONS TO AUTHORS 1. MATERIAL should be original and not published elsewhere, in whole or in part. 2. LAYOUT should be as follows: (a) Centred masthead to consist of Title: informative but concise, without abbreviations and not including the names of new genera or species Author’s(s’) name(s) Address(es) of author(s) (institution where work was carried out) Number of illustrations (figures, enumerated maps and tables, in this order) (b) Abstract of not more than 200 words, intelligible to the reader without reference to the text (c) Table of contents giving hierarchy of headings and subheadings (d) Introduction (e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents (f) Summary, if paper is lengthy (g) Acknowledgements (h) References (i) Abbreviations, where these are numerous 3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced with 2,5 cm margins all round. First lines of paragraphs should be indented. 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REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: ‘Smith (1969) describes .. .’ ‘Smith (1969: 36, fig. 16) describes . ‘As described (Smith 1969a, 19696; aubs nes ‘As described (Haughton & Broom Fees ‘As described (Haughton et al. 1927) . Note: no comma separating name and: year pagination indicated by colon, not p. names of joint authors connected by ampersand et al. in text for more than two joint authors, but names of all authors given in list of references. (b) Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, b, etc. to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a). For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list o scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number (only if independently paged) in parentheses, pagination (first and last pages of article). Examples (note capitalization and punctuation) BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140. FISCHER, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. zen. 74: 627-63 4. Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320. Konn, A. J. 1960b. Spawning behaviour, eee! masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): THIELE, J. 1910. Mollusca: B. Polypiecohorss Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. (continued inside back cover) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 74 ~ Band November 1977 November Part 3 Deel ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA By K. G. McKENZIE Cape Town Kaapstad The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular intervals as material becomes available Obtainable from the South African Museum, P.O. Box 61, Cape Town Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM word uitgegee in dele op ongereelde tye na beskikbaarheid van stof Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad OUT OF PRINT/ UIT DRUK 1, 201-3, 5-8), 3(1-2, 4—S, 8, t.-p.i.), 51-3, 5, 7-9), 6(1, t.—p.1.), 71-4), 8, 9(1-2, 7), 10(1-3), 11(1-2, 5, 7, t.-p.i.), 15(4—-5), 24(2), 27, 31(1-3), 33 Price of this part/Prys van hierdie deel R4,10 Trustees of the South African Museum © _ Trustees van die Suid-Afrikaanse Museum 1977 ISBN 0 908407 24 6 Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., | Die Rustica-pers, Edms., Bpk., Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA By K. G. MCKENZIE Riverina College of Advanced Education, Wagga Wagga, Australia (With 24 figures) [MS. accepted 7 June 1977) ABSTRACT An illustrated generic key to South African continental ostracodes is provided together with the diagnoses of two new cypridopsinid genera, Sarscypridopsis and Kapcypridopsis. CONTENTS PAGE Introduction . : ; » 45 Systematics . ; , ft 46 Key : : : : . 49 Acknowledgements . ; : 53 References. A We HS Appendix : 2 ; Bhs) INTRODUCTION In a previous paper (McKenzie 1971a) the known continental and mainly freshwater Ostracoda of South Africa were listed together with a complete bibliography of such ostracode studies in the Republic. This paper complements that previous one by providing an illustrated key to the known ostracode genera of South Africa. The key is meant to be used with reference to illustrations in earlier papers by Sars (1896, 1924a, 19246) and Rome (1965) as well as to the illustrations in this text. Ostracoda are microscopic crustaceans with a bivalve shell which usually completely covers the body when it is closed. This shell is commonly called the carapace. The ostracode body has 5-7 pairs of limbs—7 pairs in freshwater taxa—and, usually, a furca at the posterior, the rami of which attach to paired chitin supports. From the anterior, the limbs are termed as follows: antennule; antenna; mandible; maxillule; and the first through to the third thoracic limbs (PI to PIII). The furca normally carries two claws and two bristles on each ramus but can be strongly modified, e.g. into a whip-like seta in Cypridopsinae. Sometimes, as in Darwinula, it is absent. The furca in the genus Gesa is described in the key as having two claws only. This follows the original description (Hartmann 1957). However, this 45 Ann. S. Afr. Mus, 74 (3), 1977: 45-103, 24 figs. 46 ANNALS OF THE SOUTH AFRICAN MUSEUM original description was of a juvenile form not of an adult. It is probable that the furca in an adult Gesa will be found to carry the normal complement of claws and bristles on each ramus. Some characters which are illustrated in the Figures are not normally recorded for ostracodes. These include the «, 6 and y bristles on the mandible palp which have been so named by Danielopol (personal communication 1970) and which appear to have some utility in taxonomy (e.g. compare Figures 6 and 14). Similarly, rake-like organs are very rarely illustrated but may also prove to be useful taxonomic characters when known for more genera (McKenzie 19715). There is also a special terminology for characters of the ostracode carapace. These include: adductor muscle scar patterns (Fig. | et seq.); marginal or radial pore canals; normal pore canals which can be either simple and open or sieve- like (Fig. 1); and various types of hingement. The hingement terminology used here is that used in the American Treatise on Ostracoda (Moore ed. 1961) and is illustrated in Figure 1. Marginal pore canals are used to distinguish between genera in the tribe Stenocypridini. In particular, the genus Chrissia Hartmann, 1957 is distinguished by its very short marginal pore canals, whereas the species which were ascribed to Parastenocypris (McKenzie 1971a) have well-developed branching marginal pore canals (Figs 15-17). In this paper the new taxa comprise two new cypridopsinid genera which were foreshadowed earlier (McKenzie 1971a), namely: the genus Sarscypridopsis ; and a new genus from Table Mountain, Kapcypridopsis. These new genera are described in the systematics section below. SYSTEMATICS Family Cyprididae Subfamily Cypridopsinae Tribe Cypridopsini Genus Kapcypridopsis gen. nov. Figs 6-7 Type species Kapcypridopsis barnardi sp. nov. Derivation of name Kap = Cape (Afrikaans), barnardi for the late Dr K. H. Barnard, a tireless worker on South African crustaceans and molluscs and former director of the South African Museum. Diagnosis In lateral view, carapace small; subreniform; smooth; greenish or yellowish; valves subequal; dorsal margin convex; ventral margin inflexed medially: ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 47 anterior margin slightly more broadly rounded than the posterior; valve height slightly more than half valve length. In dorsal view, elliptical. Internally, inner lamellae broad anteriorly and posteriorly; inner margin regularly curved; vestibule large; line of concrescence following the outer margin; marginal pore canals short, fairly numerous; normal pore canals open and simple; muscle scar pattern including 4 adductor and 2 mandibular scars (sometimes individual adductor scars are divided and the adductor group appears to consist of 5 or 6 scars); hingement of simple ridge and groove type. Antennule 7-segmented; length ratios of the 5 proximal segments are 20:13:10: 10:15 (or 15:10: 8:8: 11). Antenna 4-segmented, length ratios of the 3 proximal segments are 7:4: 1; ‘natatory’ setae 6 in number, short, reaching about the middle of the second endopod segment. Mandible coxa normal; respiratory epipodial plate with 8 Strahlen; palp 4-segmented; «, f and y bristles as illustrated. Maxillule respiratory epipod with 4 downwards- directed setae plus 17 Strahlen; lobes and palp elongate; palp 2-segmented, 3rd lobe with two smooth Zahnborsten. PI epipod with 4 Strahlen; palp strongly dimorphic as is usual in cypridaceans with 3 terminal bristles in females but modified into a clasping organ in males. PII normal, with a powerful terminal claw. P HI with a short terminal segment which bears a fairly short reflexed seta. Furca strongly modified, whip-like. Rake-like organs each with about 10 teeth. Eye with 2 ommatophores. In males, other dimorphic characters include a Zenkers organ with 13 whorls; and hemipenis as illustrated (Fig. 7: 7). Dimensions Length, about 0,6 mm; height, about 0,35 mm. Material Small population of six individuals including mature males and ovigerous females. Some forms were greenish, others yellowish, when collected. Syntypes: South African Museum catalogue number SAM-A15451. Locality Top of Table Mountain, in small pool about 400m from the cable car terminus; Cape Town, Cape Province, the Republic of South Africa. Collector and data collected K. G. McKenzie; September 1970. Discussion The regularly subreniform shape of this taxon is sufficient to distinguish it from Plesiocypridopsis Rome, Cypridopsis Brady, and Sarscypridopsis gen. nov. since Plesiocypridopsis and Sarscypridopsis are subtriangular in shape while Cypridopsis is subovate and this new genus is subreniform. Also, unlike these genera, Kapcypridopsis has broad anterior and posterior inner lamellae. 48 ANNALS OF THE SOUTH AFRICAN MUSEUM Genus: Sarscypridopsis gen. nov. Fig. 8 Type species Sarscypridopsis gregaria (Sars) 1896. Derivation of name For G. O. Sars, pioneer worker on South African Ostracoda. Diagnosis In lateral view, carapace small-medium; subtriangular; surface pitted and frequently hirsute; often greenish to brown; valves subequal; dorsal margin strongly convex; ventral margin inflexed medially; anterior margin rounded, tending to narrow anteroventrally; posterior margin more broadly rounded than the anterior margin; valve height usually well over half valve length. In dorsal view elongate subovate moderately broad (breadth about half the length); greatest breadth behind the middle. Internally, inner lamellae moderately broad anteriorly and narrow posteriorly; inner margin regularly curved, vestibule large, line of concrescence following the outer margin; marginal pore canals short, numerous; normal pore canals simple, open; central muscle scar pattern of 4 adductors and 2 mandibulars; hingement of simple ridge and groove type. Antennule 7-segmented; antenna 4-segmented; natatory setae extending well beyond the tips of the terminal claws; mandible normal, «, 8 and y bristles as illustrated (Fig. 8: 2); maxillule palp 2-segmented, elongate-cylindrical; 3rd lobe with 2 smooth Zahnborsten; PI epipod with 2 Strahlen; endopod modified in males into a clasping organ which is dissimilar between the right and left limbs (Fig. 8: 10, 13); PII normal with powerful terminal claw; PIII with a short reflexed seta; furca whip-like. In males, other dimorphic characters include a Zenkers Organ with 10-15 whorls; and hemipenis as illustrated. Discussion Of the three species in the original designation of Cypridopsis by Brady and Norman in 1889, one, Cypridopsis vidua (O. F. Miller, 1776), is the type species; C. villosa (Jurine, 1806) was made the type species of Cypridopsella Kaufmann, 1900, which is now synonymised with Potamocypris Brady, 1868, and C. aculeata Costa, 1847, was included by Sars (1924a: 160) with the South African species of Cypridopsis in his understanding of the genus at that time and indeed was made its type. However, C. vidua has priority as the type species of Cypridopsis s.s. which means that a different generic name is required for the African and possibly one European species (note that the type material of Costa’s species has been lost and his original figures and descriptions were poor). The tribe Cypridopsini is differentiated from other cypridopsine ostracodes by the fact that it possesses an elongate maxillule palp and lobes, the palp being ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 49 further characterized by an elongate second segment. This feature is in contra- distinction to the broad palp, especially at the second segment, of Potamocypris and its related cypridopsine genera. Considered in this sense, Cypridopsini include the genera Cypridopsis, Plesiocypridopsis, Neocypridopsis, Kapcypridopsis, the present new genus Sarscypridopsis, and two Korean subspecies (McKenzie 1972) which appear to be generically distinct from these others although similar in some features to Kapcypridopsis. This genus differs from Cypridopsis in that the shape is subtriangular and moderately broad rather than subovate and broad; the anterior inner lamellae are narrower, the ventral margin of the right valve is not denticulate; and there are only up to two Strahlen on the epipod of the PI. It differs from Plesio- cypridopsis in lacking the elongation of male PI palps and the characteristic upturned anterior hemipenes of that genus. Kapcypridopsis is subreniform in shape with broader anterior and posterior inner lamellae and short antennal ‘natatory’ setae, differing in all these characters from Sarscypridopsis. The Korean taxa differ somewhat in shape especially posteriorly and have broad anterior and posterior inner lamellae and short antennal ‘natatory’ setae. The genus Neocypridopsis differs from Sarscypridopsis in general shape and also in the setation of the terminal segment of the PIII, the shapes of the clasping palps of the male P I and in possessing an upturned not a downturned anterior hemipenis. Of the cypridopsinid species listed by Sars (1924a) aculeata Costa, 1847, spinifera Sars, 1924a, and echinata Sars, 1924a, form a subgroup characterized by an armature of spines as well as hairs on the shell surface. These species are at present classed with Sarscypridopsis but may merit transfer to a new category when aculeata Costa, 1847, is better understood. Apart from this latter species, which has been recorded from Iceland, Europe, central Asia, the Americas and Africa, Sarscypridopsis has a southern distribution. KEY 1. All three pairs of thoracic legs more or less pediform and non- reflexed a ve sae a ue ae ae = Been ie First pair of thoracic legs non- or subpediform, strongly dimorphic; second pair pediform; third pair strongly reflexed i Cy priGidaen. /- 2. Shell small; smooth; elongate-ovate in lateral view; adductor-muscle scar pattern feather-shaped; first pair of thoracic legs (PI) sub- pediform; males not reported since 1870 Darwinulidae, Darwinula (Fig. 1) Shell small to medium; surface usually ornamented; subrectangular, subreniform, subtrapezoidal or subrhomboidal in lateral view; adductor-muscle scar pattern of four discrete scars in a subvertical row; all three pairs of thoracic limbs pediform; typically bisexual Cytheridae 3. 50 Il, ANNALS OF THE SOUTH AFRICAN MUSEUM Shells small or medium; ornamented; subrectangular to subreniform; radial pore canals few, straight; normal pore canals simple; hinges modified lophodont ot i. Limnocytherinae 4. Shells small or medium; prnamentede sulargetenoule, subtrapezoidal or subrhomboidal; adel pore canals few or numerous, usually straight; normal pore canals of sieve type; hinge types merodont, entomodont or gonglyodont He Cytherideinae, Loxoconchinae 5. Shell small; subrectangular to subreniform; non-dimorphic Limnocythere Shell medium; subrectangular; strongly dimorphic (females very broad, but males slender, posteriorly) .. .. Gomphocythere (Fig. 2) Shell small to medium; subrectangular or subtrapezoidal; radial pore canals few or numerous; hinge type merodont or entomodont Cytherideinae 6. Shell small; subrhomboidal; radial pore canals few, straight; hinge type gonglyodont ..__.... a ie Loxoconcha (Fig. 1) Shell small-medium; subtrapezoidal; adic pore canals few; hinge type merodont ee ah .. Perissocytheridea Shell medium; dubrestaneulan: bacliel pore canals numerous; hinge type ene medont ne a ie a Cypriden (Fig. 1) Shell medium; subrectangular; “ gipiaes pitted, dorsomedial sulci present; palps of first pair of thoracic legs two-segmented Ilyocypridinae, I/yocypris (Fig. 3) Shell differing from this; palps of first pair of thoracic legs non- segmented .. a 8. Third thoracic limb beating at ee one 5 lone fefiexed feaninall ae: Zenkers Organ (males only) with 5-6 whorls Paracypridinae, Candoninae and Cyclocypridinae 9. Third thoracic limb with short reflexed terminal seta; Zenkers Organ (males only) with more than 6 whorls .. ae a, Shell medium, subreniform or elongate sutenernaulle in fareral view; elliptical in dorsal view, antennal natatory setae poorly developed or absent e ae Paracypridinae, Candoninae 10. Shell small- aaceliunns “gtibewathats in lateral view; antennal natatory setae well developed (extending to the terminal claws or beyond) Cyclocypridinae 11. Shell medium, elongate-subtriangular in lateral view; furca with a posterior bristle, furcal claws without spurs Paracypridinae, Paracypris Shell medium, subreniform in lateral view; furca lacking a posterior bristle, furcal claws with spurs... es Candoninae, Candonopsis Shell small-medium; right valve denticulate ventrally especially on anteroventral margin; compressed in dorsal view; furca normal Cyclocypridinae, Physocypria (Figs 3-4) Shell small; non-denticulate ventrally; broadened in dorsal view, furca normal io a cn - .. Cyclocypridinae, Cyclocypris 12? 20. Ale ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 51 Shells small to very large; furca with at least two terminal claws .. 19. Shells small to medium; furca whip-like. . _ Cypridopsinae 13. Shell with distinctive pustulose surface ornamentation and charac- teristic lunette-bearing inner lamellae; maxillule 3rd lobe with at least four Zahnborsten .. ie Oncocypris (Fig. 5) Shells different; maxillule ard fone with ine Zahnborsten .. Ae 14: Shell subtriangular, produced at each end; left valve overlapping right valve anteriorly and posteriorly, but overlapped by the right valve dorsally; maxillule palp distal segment spatulate Cyprilla (Fig. 5) Shell of a different type; maxillule palp elongate oo Ue me ilps): . Shell small-medium; subtriangular to subovate in lateral view; ovate in dorsal view; left valve overlapping right valve anteriorly; anterior inner lamellae broad, posterior inner lamellae narrow ue: ae NG: Shells of a different type .. we we Ld . Shell typically whitish to arcenich: Rewally Ge 408 or - nies irregular dark strigae on each valve in life; right valve denticulate anteriorly; inner lamellae as illustrated Ns wl Cypridopsis (Fig. 4) Shell typically whitish; ornamented with nite arranged concentrically or else tuberculate; without strigae; inner lamellae as illustrated Zonocypris (Fig. 9) . Shell small-medium to medium; subtriangular in lateral view; relatively compressed in dorsal view; right valve overlapping left valve anteriorly; anterior and posterior inner lamellae rather narrow; antennal natatory setae well developed .. Wf Me AS: Shell small to small-medium; valves subequal; aiibrenifornn in etal view; without strigae on the valves; anterior and posterior inner lamellae both relatively broad; antennal natatory setae short Kapcypridopsis (Figs 6-7) . In males, palp of first thoracic legs elongate, hemipenis anterior lobe curved upwards .. set a Plesiocypridopsis (Fig. 9) In males, palp of first ores. ines of normal length, hemipenis anterior lobe pointing downwards Me Sarscypridopsis (Fig. 8) Shell small-medium to medium; smooth (Cyprettini, Bradycyprini) or pitted (Cyprettini) or striate (Bradycyprini); subglobose in lateral and dorsal view; radial septa well developed at least in right valve; furca ‘normal’ but slender (Bradycyprini) or very slender (Cyprettini) | Cyprettinae 20. Shell medium to very large, typically smooth; compressed to globose in dorsal view; without radial septa (exception Stenocypris, cf. 29); furca ‘normal’, not conspicuously slender i ie ie Hi SohDD Radial septa developed in both valves .. Cyprettini—Cypretta (Fig. 10) Radial septa developed in right valve only ae Bradycyprini 21. Shell striate .. ie ae ee ss Paracypretta (Figs 10-12) Shell smooth iM ie as, a .. Bradycypris (Figs 11-12) 3 DD, D3: 24. 2c 26. Die IES. aD), 30) Sill Se 33) ANNALS OF THE SOUTH AFRICAN MUSEUM Chitin supports with loculi where they branch proximally, furca flexuous he cz Cypricercini 23. Chitin supports “fle oeull here hey branch jraees furca typically straight or evenly curved, rarely flexuous... se 29 Shell oval, with distinct posterior process in some species; chitin supports with anterior branch well developed; spermatic vesicles ($ only) coiled anteriorly .. xy ae S Cypricercus (Fig. 12) Shell oval-oblong, without posterior processes; chitin supports without an anterior branch ie < oh wm, 2Ae Furca normally developed (2 claws and 2 BnBtes) ti Tanycypris Furca with two claws only © ee) .. Gesa (cf. Introduction) Shell without a posterior inner lamella (the inner margin coincides posteriorly with the posterior shell margin cf. Rome 1965) .. wo 203 Shells with a posterior inner lamella es i ake 208 Distal segment of the. PII with a short donsal isi (Rome 1965) Isocypridini, [socypris (Fig. 13) Distal segment of the PII with a dorsal bristle which is almost as long as the terminal claw (Rome 1965) Amphibolocypridini, Amphibolocypris Chitin supports widened distally into triangular chitinous processes Herpetocypris, Stenocypridini 28. Chitin supports without distal triangular chitinous processes ie” Sale Furcal rami similar, furca with a posterior bristle Herpetocypris (Figs 13-14) Furcal rami dissimilar; furca without a posterior bristle Stenocypridini 29. Shells with radial septa... es on Ze eh Stenocypris Shells without radial septa RS I ae ee 7 Lie Marginal pore canals very short .. oe an .. Chrissia (Fig. 17) Marginal pore canals well developed and branched Parastenocypris (Figs 14-17) Either right or left valve of shell denticulate ventrally except Homocypris which, however, is of medium size only (length 1,1—1,4 mm) and has anormal furca.. as .. Cyprinotini 32. Neither right nor left valve a shel dendioulic ventrally (except Apatelecypris cf. 42); carapace large, at least 1,75 mm, except Mesocypris (0,91 mm) which has an unique furca, cf. 34 te .. 34. Shell elongate-subreniform in lateral view; anterior inner lamellae distinctly broader than posterior inner lamellae Homocypris (Figs 17-18) Shell elongate-ovate in lateral view; both anterior and posterior inner lamellae relatively, and subequally narrow ae a We OSBE Right valve denticulate antero- and/or poten ea uciln Heterocypris (Fig. 18) 34. 3) 30: Bi: 38. jy). 40. 41. 42. ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA Do) Left valve denticulate antero- and/or posteroventrally Hemicypris Furca normally developed (2 claws, 2 bristles) and not modified as below se » We Furca with the oeted ar iyaeile ransomed ate a sialbibs Clas fie spine; furcal rami weakly dissimilar sf E, Mesocypris Carapace large, with prominent secondary eieedl zone in right valve Afrocypris (Figs 18-20) Carapace large, without prominent secondary fused zone in right valve 36. Carapace large; maxillule palp elongate a ce er Oil Carapace large; maxillule palp relatively broad (i.e. approximate square or spatulate) = , oi Carapace large, without ventral ‘ale: oemulifanete seamen: o P Ul divided oe ae .. Eucypridini, Eucypris (Fig. 19) Carapace large, quant or quot ventral ala; penultimate ‘segment’ of PII not divided .. se oe 5s bts ee CyoriGdinigess Carapace large, with ventral ala .. oA .. Pseudocypris (Fig. 20) Carapace large, without ventral ala iu 3 Cypris Carapace large, tuberculate in one species; anfennal natatory setae well developed, reaching about as far as the distal part of the terminal segment iy us ve Sclerocypris (Fig. 21) Carapace large, eiiooth 6 distinetly hirsute; antennal natatory setae strongly reduced .. a . 40. Carapace large, in general shape dexeineline Pidioewprise PI male ($ only) strikingly dissimilar on left and right limbs; hemipenes (¢ only) unlike those of any other ostracode; shape of female genital lobe also distinctive cS, iar Liocypris (Fig. 22) Carapace large; general shape nnlike that of Homocypris; P 1 palps (g only) not strikingly dissimilar on left and right limbs, hemipenes (¢ only) different to Liocypris .. ae ue Megalocypridini 41. Carapace large; with a small but distinct posteroventral caudal spine on each valve ss ne Hypselecypris Carapace large; without ouch a spine on (Sihen selke ts oe . 42. Carapace large; in dorsal view indented at both ends; not denuetiate posteroventrally .. er a Megalocypris (Figs 23-24) Carapace large; in dorsal view narrowing anteriorly, but rounded, not indented, posteriorly; denticulate posteroventrally Apatelecypris (Fig. 24) ACKNOWLEDGEMENTS I am grateful to the National Institute for Water Research, Pretoria and to the British Museum (Natural History) London for the opportunity to work in South Africa and to Dr R. G. Noble who organized my visit. My original drawings were traced for publication by Mr D. Goode of the Transvaal Museum. My thanks are due to Professor P. H. A. Sneath and Dr M. J. Sackin of the 54 ANNALS OF THE SOUTH AFRICAN MUSEUM University of Leicester, who prepared the computer key of the appendix, using the PANKEY program. I am particularly grateful to Dr T. H. Barry, Director of the South African Museum, for the extended use of its laboratory and library facilities during my working visit to Cape Town, also to Dr B. F. Kensley at the South African Museum for his considerable assistance in reorganizing the illustrations. Mrs L. Vennell, Riverina College of Advanced Education, typed the manuscript. REFERENCES HARTMANN, G. 1957. Ostracoden aus dem Namaland und Transvaal. Veroff. naturw. Osnabruck 28: 50-60. McKeEnzIE, K. G. 1971a. Species list of South African freshwater Ostracoda with an appendix listing museum collections and some further determinations. Ann. S. Afr. Mus. 57: 157-213. McKenzie, K. G. 1971b. Ostracoda from Lake Peunde, near Mt. Wilhelm, New Guinea. Zool. Anz. 186: 391-403. McKeEnzigE, K. G. 1972. Results of the speleological survey in South Korea 1966. XXII. Subterranean Ostracoda from South Korea. Bull. Natl. Sci. Mus., Tokyo 15: 155-166. Moore, R. C. ed. 1961. Treatise on Invertebrate Paleontology. Part Q. Arthropoda 3 Crustacea Ostracoda. Lawrence: Geological Society of America and University of Kansas Press. Rome, D. R. 1965. Crustacea: Ostracoda. Jn Hanstrom, B., Brinck, P. & Rudebeck, G., eds. South African animal life 11: 9-58. Stockholm: Swedish Natural Science Research Council. SARS, G. O. 1896. On some South African Entomostraca raised from dried mud. Skr. Vidensk Selsk. Christiania 1895 (8): 1-56. Sars, G. O. 1924a. The freshwater Entomostraca of the Cape Province (Union of South Africa). Part Il: Ostracoda. Ann. S. Afr. Mus. 20: 105-193. Sars, G. O. 1924b. Contributions to a knowledge of the fauna of South West Africa. Ann. S. Afr. Mus. 20: 195-211. APPENDIX South African Freshwater Ostracoda Generic Key (prepared by the Program PANKEY). il. Shell: surface not smooth De 2: Furca reduced to whip-like seta 8: 3, Dorsal view of shell broad 4. 4, Maxillule 3rd lobe with 4 or more Zahnborsten, maxilla epipod with less than 3 Strahlen, shell denticulation in one or both valves absent, shell: radial septa or lunettes in inner lamellae present in both valves, shell surface pustulose not tuberculate-costate. Oncocypris 4. Maxillule 3rd lobe with 2 Zahnborsten, maxilla epipod with more than 3 up to 6 Strahlen, shell denticulation in one or both valves present, shell: radial septa or Junettes in inner lamellae, shell surface tuberculate- costate not pustulose Zonocypris 3 Dorsal view of shell ‘normal’ 5) Da Parthenogenesis: males virtually unknown, shell denticulation in one or both valves present. Cypridopsis De Parthenogenesis: both sexes known 6. 6. Maxilla male epipod not elongated. : Cyprilla 6. Maxilla male epipod elongated Plesiocypridopsis ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 53 De, Furca ‘normal’ ie Us 7th limb reflexed, terminal seta long oye 8. Maxilla endopod 2-segmented, chitin supports: distally with simple point, Zenker’s Organ with more than 15 whorls, shell: dorsal sulci present Ilyocypris 8. Maxilla endopod unisegmented ; 9. Shell denticulation in one or both valves present Physocypria oF Shell denticulation in one or both valves absent Cyclocypris Is 7th limb reflexed, terminal seta short 10. Furcal rami very slender, Zenker’s Organ with 7-14 whorls, shell: radial septa or lunettes in inner lamellae‘ present in both valves, shell length 0,45—0,90 mm Cypretta 10. Zenker’s Organ with more than 15 whorls Ht Wi. Furcal rami slender, shell: radial septa or lunettes in inner lamellae present in one valve, shell surface striate, shell length 0,90-1,75 mm Paracypretta iN Furcal rami ‘normal’ IZ, 12. Dorsal view of shell broad Cypris I. Dorsal view of shell ‘normal’ Sclerocypris Shell: surface smooth (includes weak pitting) 13. I. 7th limb penultimate segment undivided 14, 14. Parthenogenesis: both sexes known, maxilla epipod with less than 3 Strahlen Candonopsis 14. Parthenogenesis: males virtually unknown IS Sy Mandible palp comb present, maxilla epipiod with more than 7 Strahlen, maxilla endopod 3-segmented, 7th limb not reflexed, furca absent, shell: inner lamellae absent. Darwinula 15. Mandible palp comb absent, maxilla epipod with more than 3 up to 6 Strahlen, maxilla endopod unisegmented (but ‘ghost’ segmentation indicated), 7th limb reflexed, furca ‘normal’, shell: inner lamellae present. Paracypris 13. 7th limb penultimate segment divided ! 16. 16. Natatory setae of antennae weakly developed Ws We Maxilla epipod with less than 3 Strahlen, furca: posterior margins dissimilar, shell densely hirsute Mesocypris 7: Maxilla epipod with more than 3 up to 6 Strahlen 18. 18. Furca reduced to whip-like seta, Zenker’s Organ with 7-14 whorls, Shell length 0,45—0,90 mm Kapcypridopsis 18. Furca ‘normal’ 19. 19. Chitin supports: distally with simple point Liocypris 19. Chitin supports: distally elaborated 20. 20. Shell denticulation in one or both valves absent, shell length 3,60-7,30 mm Megalocypris 20. Shell denticulation in one or both valves present, shell length 1,75-3,60 mm Apatelecypris 16. Natatory setae of antennae well developed Die le Chitin supports: distally elaborated DI UD, Furca: posterior bristle absent M23 D3) Shell length 1,75-3,60 mm 24. 24. Shell: radial septa or lunettes in inner lamellae present in both valves Stenocypris 24. Shell: radial septa or lunettes in inner lamellae absent ‘Parastenocypris’ W3). Shell length 0,90-1,75 mm WD Di: 6th limb: distal bristle almost as long as the distal claw, furca: posterior margins similar, chitin supports: triangular process absent, dorsal view of shell very compressed, shell: inner lamellae absent posteriorly Amphibolocypris 56 ANNALS OF THE SOUTH AFRICAN MUSEUM Dy 6th limb: distal bristle short, furca: posterior margins dissimilar, chitin supports: triangular process present, dorsal view of shell normal, shell: inner lamellae present Chrissia DOP Furca: posterior bristle present 26. 26. Chitin supports: triangular process present, shell length 1,75—-3,60 mm Herpetocypris 26. Chitin supports: triangular process absent 24 Dap Dorsal view of shell very compressed, shell: inner lamellae absent posteriorly, shell length 0,90-1,75 mm __Isocypris ile Dorsal view of shell ‘normal’ 28. 28. Shell posterior processes absent Afrocypris 28. Shell posterior processes present Hypselecypris Ml. Chitin supports: distally with simple point ; BS) Dorsal view of shell broad 30. 30. Furcal rami slender, shell: radial septa or lunettes in inner lamellae present in one valve, shell length 0,90-1,75 mm Bradycypris 30. Furcal rami ‘normal’ 31. 31. Dorsal view of shell: alae absent Eucypris 31. Dorsal view of shell: alae present Pseudocypris IES). Dorsal view of shell ‘normal’ Syd oe Chitin supports: loculi present where they branch proximally 338 33. Chitin supports: loculi present and prominent anterior branch, shell posterior processes present Cypricercus 333), Chitin supports: loculi present but without prominent anterior branch, shell posterior processes absent Tanycypris 3, Chitin supports: loculi absent where they branch proximally 34. 34. Shell denticulation in one or both valves absent, shell inner lamellae: anterior broad posterior narrow Homocypris 34. Shell denticulation in one or both valves present 335). 35.0 Shell denticulation in right valve Heterocypris 35}, Shell denticulation in left valve Hemicypris REFERENCE PANKHURST, R. J., 1970. A computer program for generating diagnostic keys. Computer J. 12: 145-151. Fig. 1 1. Darwinula sp.; ovigerous 2; adductor muscle scars, left valve. x 175. 2. Darwinula sp.; ovigerous °; outline right valve. x 125. 3. Generalized rimmed normal pore canal. x 1400. 4. Generalized rimless normal pore canal. x 2500. 5. Generalized sieve type normal pore canal. x 1000. 6. Generalized lophodont hingement right valve (after Moore ed. 1961). 7. Generalized merodont hingement right valve (after Moore ed. 1961). 8. Generalized entomodont hingement right valve (after Moore ed. 1961). 9. Generalized gonglyodont hingement right valve (after Moore ed. 1961). 10. Cyprideis sp.; hingement right valve. x 100. 11. Cyprideis sp.; adductor muscle scars and fulcral scar (dashes). x 225. 12. Loxoconcha sp.; adductor muscle scars and fulcral scar (dashes). x 300. 13. Loxoconcha sp.; anterior hingement right valve. x 250. ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA >)// ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 59 Fig. 2 Gomphocythere expansa Sars, 1924; SAM-A11303; ovigerous 9; PI and PII. x 400. Posterior of body with caudal lobes. x 400. P Ill. x 400. Antennule. x 400. Antenna. x 400. ARWN- ANNALS OF THE SOUTH AFRICAN MUSEUM 60 ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 61 Fig. 3 fe oe ee ee Ilyocypris australiensis Sars, 1889; SAM-A11; adult 9; detail anterior margin right valve showing radial pore canals and muscle scars. x 160. Antennule, six terminal segments. x 400. P III, terminal segments. x 400. PI. x 400. Mandible palp. x 400. Furca. x 400. Physocypria capensis (Sars, 1896); SAM-A11136; adult 3; internal view right valve. x 160. ANNALS OF THE SOUTH AFRICAN MUSEUM 62 ws 4 ZN | d ry \ Des e// 2 rT Be ||) t/a) || eee zane = <8 os Sekt | RL OW : fe ys if e & ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 63 Fig. 4 Physocypria capensis (Sars, 1896); SAM-A11136; adult ¢; Zenkers Organ. x 400. Mandible palp, detail 8 and y bristles. x 400. Antennule, six terminal segments. x 400. P III, terminal segment. x 400. Cypridopsis viduella Sars, 1896; SAM-A11171; adult 2; PI. x 400. Rake-like organ. x 400. P III, terminal segment. x 400. Antennule, six terminal segments. x 400. Mandible palp, detail «, 8 and y bristles. x 400. Internal view right valve. x 160. Sie EON EA ee eS Seo — G4) ) (WO Sea), GOOPANINIATE S OF THE SOUTH AFRICAN MUSEUM ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 65 Fig. 5 1. Oncocypris cf. voeltzkowi Miller, 1898; SAM—A11374; ovigerous 2; maxillule palp and lobes, third lobe with some Zahnborsten. x 400. 2. PI. x 400. 3. Anterior margin of left valve. x 160. 4. Mandible palp, detail of «, B and y bristles. x 400. 5. Antennule, six terminal segments. x 400. 6. Cyprilla arcuata Sars, 1924; SAM-A11158; ovigerous 2; antennule, six terminal segments. x 400. 7. Internal view left valve. x 160. 8. Maxillule palp. x 400. 9. PII. x 400. 0. Antenna, first endopod segment. ~ 400. 1. PII. x 400. ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 67 Fig. 6 1. Kapcypridopsis barnardi McKenzie, 1977; SAM-A15451; adult 2 (yellowish form); internal view left valve. x 160. 2. Antennule segments. x 400. 3. Antenna, with ‘natatory’ setae. x 400. 4. Rake-like organ. x 400. 5. Mandible palp, detail of «, 8 and y bristles. x 400. 6. Maxillule palp and lobes. x 400. 7. PIII, terminal segments. x 400. sae IT. >< -400. Ss Burca. < 400. 0. —" PI. x 400. 68 ANNALS OF THE SOUTH AFRICAN MUSEUM ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 69 Fig. 7 Ile NDAR WH Kapcypridopsis barnardi McKenzie, 1977; SAM-—A15451; adult ¢ (greenish form); internal view right valve. x 160. P I right palp. x 400. PII. x 400. Zenkers Organ. x 400. Eye. x 400. Antennule, six terminal segments. x 400. Hemipenis. x 400. 70 ANNALS OF THE SOUTH AFRICAN MUSEUM . Lee @ aneeeBane =anOe = 1 = ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 1 Fig. 8 1. Sarscypridopsis striolata (Sars, 1924); SAM-—AI1201; ovigerous 9; internal view right valve. x 160. 2. Sarscypridopsis ochracea (Sars, 1924); SAM-A11220; adult 3; mandible palp detail -a, B and y bristles. x 400. 3. Antennule, six terminal segments. x 400. 4. Antenna, first endopod segment with sensory bristle. x 400. 5. Posteroventral right valve. =< 160. 6. PII. x 400. 7. Hemipenis. =< 400. 8. Maxillule third lobe with Zahnborsten. x 400. 9. Muscle scars. x 160. 10. PI, left palp. x 400. - 11. PIII, terminal segment. = 400. 12. Maxillule palp. x 400. 13. PI, right palp. x 400. ANNALS OF THE SOUTH AFRICAN MUSEUM 2 a Th ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 73 Eig, 9 I. eg 3g 4. GN Plesiocypridopsis cf. aldabrae (Miller, 1898); SAM-A5940; adult ¢; P I right palp. x 400. P I left palp. x 400. Antenna, sensory bristle first endopod segment. x 400. Zonocypris cordata Sars, 1924; SAM-—A11144; ovigerous 2; anterior margin left valve. x 160. Plesiocypridopsis cf. aldabrae (Miller, 1898); SAM-A5940; adult 2; ventral view of labia with rake-like organs. x 400. Mandible palp, detail of «, 68 and y bristles. 400. Antennule, six terminal segments. x 400. P III, terminal segment. x 400. Adult 3; head and oral region. x 160. Anteroventral right valve. ~ 160. Hemipenis. x 400. ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA Is) Fig. 10 . Cypretta cf. turgida (Sars, 1895); SAM—A11159; ovigerous 9; anterior margin right valve with radial septa. = 160. . Antenna endopod segments with sensory bristle. 400. Antennule six terminal segments. x 400. Mandible palp, detail of «, 8 and y bristles. x 400. Maxillule palp and third lobe with Zahnborsten. x 400. P I, terminal segments. x 400. Distal furca. x 400. . Paracypretta ampullacea Sars, 1924; SAM-A11278; ovigerous 2; sensory bristle first endopod segment of antenna. x 400. P III, distal terminal segment. x 400. Mandible palp, detail of «, 8 and y bristles. x 400. . Maxillule palp and third lobe with Zahnborsten. x 400. ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 77 Fig. 11 I — SO WIA Paracypretta ampullacea Sars, 1924; SAM—A11278; ovigerous @; internal view left valve. x 40. 2. Chitin support. = 160. 3. 4. Bradycypris intumescens (Brady, 1907); SAM-A11299; ovigerous 2; internal view right Detail anterior margin radial pore canals left valve. x 160. valve, showing radial septa. x 40. Rake-like organ. x 400. Antennule, six terminal segments. x 160. Antenna, sensory bristle first endopod segment. x 400. Maxillule palp and lobes, third lobe with Zahnborsten. x 160. Mandible palp, detail of «, 8 and y bristles. x 400. Mandible coxale. x 160. 78 ANNALS OF THE SOUTH AFRICAN MUSEUM ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 719 Fig. 12 1. Paracypretta ampullacea Sars, 1924; SAM-A11278; ovigerous 9; furca. x 160. 2 Bye, < 400: 3. Antennule, terminal six segments. x 160. 4. Antennule, detail second segment. x 400. 5. Bradycypris intumescens (Brady, 1907); SAM-A11299; ovigerous 2; furca. x 160. 6. Chitin support. x 160. 7. Cypricercus cuneatus Sars, 1896; SAM-~A11140; adult 3; antennule, detail first and second segments with gland and chemo-receptor. x 400. Chitin support. x 160. Antenna, sensory bristle of first endopod segment. x 400. . Cap of Zenkers Organ. x 400. . Mandible palp, detail of «, 8 and y bristles. x 400. . PI, right palp. x 400. . Furca. x 160. . PI, left palp. x 400. 80 ANNALS OF THE SOUTH AFRICAN MUSEUM ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 8] Fig. 13 1. Isocypris priomena Miller, 1908; SAM-A11177; ovigerous 2; antennule second segment. x 400. Internal view left valve. x 40. P II, three terminal segments. x 160. Antenna endopod segments. x 160. Anterior margin, detail radial pore canals. x 160. Mandible palp, detail of «, 8 and y bristles. x 160. P III terminal segment. x 400. Maxillule palp. x 160. Maxillule third lobe with Zahnborsten. x 160. Herpetocypris chevreuxi (Sars, 1896); SAM-A1148; ovigerous 2; antennule second and third segments. x 160. 11. Antennule second segment chemo-receptor. x 400. 12. Rake-like organ. x 160. 13. PIII, terminal segment. x 160. 14. Internal view left valve. x 40. SESE ON a S — ANNALS OF THE SOUTH AFRICAN MUSEUM ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 83 Fig. 14 1. Herpetocypris chevreuxi (Sars, 1896); SAM-A1148; ovigerous 2; maxillule third lobe. x 400. 2. Chitin support. x 160. 3. PI, terminal segments and part of claw. x 160. 4. Mandible palp, detail of «, B and y bristles. x 400. 5. Parastenocypris declivis (Sars, 1924); SAM-A3797; ovigerous 2; internal view, left valve. x 40. 6. Distal furca. x 160. 7. Mandible palp, detail of «, 8 and y bristles. x 160. 8. Antennule second segment with chemo-receptor. x 400. 84 ANNALS OF THE SOUTH AFRICAN MUSEUM ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 85 Fig. 15 igh se oe ee Parastenocypris declivis (Sars, 1924); SAM-A3797; adult 2; chitin support. x 160. Detail anterior margin left valve. x 160. Detail posteroventral margin left valve. x 160. Maxillule third lobe with Zahnborsten. x 400. Parastenocypris hodgsoni (Sars, 1924); SAM-—A11126; ovigerous 9; internal view left valve. x 16. Furca, detail of posterior margin. x 160. Detail anteroventral margin left valve. x 160. Furca. x 40. Furca, other ramus. x 160. Antennule, detail of second segment chemo-receptor. 400. . Parastenocypris pardalis (Sars, 1924); SAM-—A11167; internal view right valve. x 16. Internal view left valve. x 16. . Antenna, sensory bristle of basal endopod segment. x 400. 86 ANNALS OF THE SOUTH AFRICAN MUSEUM ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 87 Fig. 16 1. Parastenocypris pardalis (Sars, 1924); SAM-—-A11167; adult 3; detail anterior margin left valve. x 160. 2. PI, left palp. x 160. 3. PI, right palp. x 160. 4. Antennule second and third segments. x 160. 5. Parastenocypris olivacea (Sars, 1924); SAM-A3797; adult 3; internal view left valve. x 16. 6. Muscle scars. x 160. 7. Detail anterior margin left valve. =< 160. 8. Antenna terminal segment, detail of comb bristle. x 400. 9. Mandible palp, detail of « and f bristles. =< 400. 10. Mandible palp, detail of y bristle. x 400. 11. Terminal rake-like organ. = 160. 12. Antenna basal endopod segment, showing exopod and sensory bristle. < 400. 13. Hemipenis. x 160. 88 ANNALS OF THE SOUTH AFRICAN MUSEUM ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 89 Rig. 17 We Parastenocypris olivacea (Sars, 1924); SAM-—3793; adult 3; distal furca. <= 160. 2. Frontal head region and labrum. x 160. 8. ?Chrissia fascigera (Sars, 1924); SAM-A11361; adult 9; internal view posteroventral left valve. 160. P II, two terminal segments and claw. » 160. Outline hemipenes. 40. Chitin support. < 160. Distal furca. = 160. Antenna, sensory bristle. = 160. P III, distal segment. = 160. . Antennule, six distalmost segments. = 160. . Maxillule third lobe showing Zahnborsten. 160. Rake-like organ. x 160. Mandible palp, detail x, 8 and y bristles. < 160. ANNALS OF THE SOUTH AFRICAN MUSEUM 96 © i li 4 a, ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 97 Fig. 21 1. Sclerocypris clavularis Sars, 1924; SAM-A11118; ovigerous °; antennule, six distalmost segments. x 160. Rake-like organ. x 160. P I, four terminal segments. x 160. Chitin support. < 160. Maxillule palp and third lobe showing Zahnborsten. x 160. Carapace, muscle scars. = 40. Mandible palp, detail «, 8 and y bristles. x 160. Anterior margin left valve. =< 160. Anterior margin right valve. = 160. Internal view right valve. x 16. Distal furca. x 160. FHSS SNIDARYWNY pay 98 ANNALS OF THE SOUTH AFRICAN MUSEUM —_* ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA Fig. 22 1. —a ee aS 99 Liocypris grandis Sars, 1924; SAM-6281; adult $; carapace, detail anterior margin reticulate valve microstructure (in part). x 40. Internal view right valve. x 16. Furca, distal detail. « 160. Chitin support. x 40. Antennule, detail of second segment showing gland and bristle. < 160. Maxillule third lobe with Zahnborsten. x 160. Maxillule palp. x 160. PI palp. x 40. Rake-like organ. 160. Mandible palp, indicating complexity of setation. < 160. Mandible epipod. x 160. [ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 101 Fig. 23 1. Megalocypris hispida Sars, 1924; SAM-A11845; adult 2; mandible palp, detail of «, p and y bristles. x 160. 2. SAM-A11844; adult $; antennule, six distalmost segments. x 40. 3. SAM-A11845; adult 2; mandible coxale, detail of toothbrush bristles between first and second teeth. x 400. 4. Megalocypris sp. (smooth form with hemipenis similar to M. Hiberelitatt Sars, 1924); SAM-A11849; adult 3; antenna, endopod segments. x 40. and 6. Rake-like organs. x 160. Mandible palp, detail of « and f bristles. Note that this male carries only one y bristle. x 160. 8. Megalocypris princeps Sars, 1898; SAM-1479; adult ¢; rake-like organ. x 160. 9. and 10. Megalocypris tuberculata Sars, 1924; SAM-A11273; adult 3; rake-like organs. x 160. 11. PI palp. x 40. 12. Furca. x 40. 13. Chitin support. x 40. 14. PII, three terminal segments. x 160. 15. Maxillule palp. x 160. 16. Maxillule third lobe with Zahnborsten. x 160. 17. Antenna, sensory bristle of first endopod segment. x 160. 18. Carapace, muscle scars. x 40. 19. Outline of hemipenis. x 40. apes 102 ANNALS OF THE SOUTH AFRICAN MUSEUM ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 103 Fig. 24 Megalocypris tuberculata Sars, 1924; SAM—A1i1273; adult $; mandible palp, detail of «, B and y bristles. Note that the specimen may be a freak since it carries two y bristles instead of the normal complement of one only. x 160. Antennule, detail of second segment showing gland, chemo-receptor and bristle. x 160. Apatelecypris brevis Sars, 1924; SAM-—A11369; adult ¢; carapace, internal view right valve, muscle scars not shown. x 16. Detail anteroventral radial pore canals. = 160. Detail anterior margin right valve. x 40. Antenna, detail ‘natatory’ setae. =< 160. P I, detail of left palp. x= 160. P I, detail of right palp. x 160. Antennule, detail of second segment showing chemo-receptor and bristle. x 160. Mandible palp, detail of y bristle. = 160. Mandible palp, detail of « and f bristles. = 160. . Tips of rake-like organs. x 160. Furca. x= 40. Maxillule, palp and third lobe showing Zahnborsten. = 160. . Chitin support. x 40. . PII, three terminal segments. x 160. Dat Poke a , gat: Hay Se: ad r nua OF ty a » an ’ 6. SYSTEMATIC papers must conform with the International code of zoological nomenclature (particularly Articles 22 and 51). Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed by the appropriate Latin (not English) abbreviation, €.g. gen. NOV., Sp. Nov., comb. nov., syn. nov., etc. ‘An author’s name when cited must follow the name of the taxon without intervening punctuation and not be abbreviated; if the year is added, a comma must separate author’s name and year. The author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is transferred from its original genus. The name of a subsequent user of a scientific name must be separated from the scientific name by a colon. Synonymy arrangement should be according to chronology of names, i.e. all published scientific names by which the species previously has been designated are listed i in chronological order, with all references to that name following in chronological order, e.g.: Family Nuculanidae Nuculana (Lembulus) bicuspidata (Gould, 1845) Figs 14-15A Nucula (Leda) bicuspidata Gould, 1845: 37. Leda plicifera A. Adams, 1856: 50. Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b). Nucula largillierti Philippi, 1861: 87. Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. Note punctuation in the above example: comma separates author’s name and year semicolon separates more than one reference by the same author full stop separates references by different authors figures of plates are enclosed in parentheses to distinguish them from text-figures dash, not comma, separates consecutive numbers Synonymy arrangement according to chronology of bibliographic references, whereby the year is placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable. In describing new species, one specimen must be designated as the holotype; other speci- mens mentioned in the original description are to be designated paratypes; additional material not regarded as paratypes should be listed separately. The complete data (registration number, depository, description of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.: Holotype SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach, Port Elizabeth (33°51’S 25°39’E), collected by A. ‘Smith, 15 January 1973. Note standard form of writing South African Museum registration numbers and date. 7. SPECIAL HOUSE RULES Capital initial letters (a) The Figures, Maps and Tables of the paper when referred to in the text e.g. *... the Figure depicting C. namacolus...’; ‘*. . . in C. namacolus (Fig. 10)...” (b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded by initials or full names e.g. Du Toit but A.L. du Toit; Von Huene but F. von Huene (c) Scientific names, but not their vernacular derivatives e.g. Therocephalia, but therocephalian Punctuation should be loose, omitting all not strictly necessary Reference to the author should be expressed in the third person Roman numerals should be converted to arabic, except when forming part of the title of a book or article, such as ‘Revision of the Crustacea. Part VIII. The Amphipoda.’ Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial capital letter, provided the same generic name is used consecutively. Name of new genus or species is not to be included in the title: it should be included in the abstract, counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts. K. G. McKENZIE ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA e SOU Wy a* ee - VOLUME 74 PART 4 NOVEMBER 1977 | ISSN 0303-2515 -_ MUS. COMP. ZOOL: | » LIBRARY JAN 3 1 1978 HARVARD UNIVERSITY _ANNALS CAPE TOWN | — INSTRUCTIONS TO AUTHORS 1. MATERIAL should be original and not published elsewhere, in whole or in part. 2. LAYOUT should be as follows: (a) Centred masthead to consist of Title: informative but concise, without abbreviations and not including the names of new genera or species Author’s(s’) name(s) Address(es) of author(s) (institution where work was carried out) Number of illustrations (figures, enumerated maps and tables, in this order) (b) Abstract of not more than 200 words, intelligible to the reader without reference to the text (c) Table of contents giving hierarchy of headings and subheadings (d) Introduction (e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents (f) Summary, if paper is lengthy (g) Acknowledgements (h) References (i) Abbreviations, where these are numerous 3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced with 2,5 cm margins all round. First lines of paragraphs should be indented. 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REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: ‘Smith (1969) describes...’ ‘Smith (1969: 36, fig. 16) describes... .’ “As described (Smith 1969a, 1969b; Jones 1971)’ ‘As described (Haughton & Broom 1927)...’ ‘As described (Haughton et al. 1927)...’ Note: no comma separating name and year pagination indicated by colon, not p. names of joint authors connected by ampersand et al. in text for more than two joint authors, but names of all authors given in list of references. (b) Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, b, etc. to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a). For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list o scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number (only if independently paged) in parentheses, pagination (first and last pages of article). Examples (note capitalization and punctuation) BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140. FISCHER, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. gén. 74: 627-634. Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320. Konn, A. J. 1960b. Spawning behaviour, eae masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4 ): 1. THIELE, J. 1910. Mollusca: B. Dalene oubea Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. (continued inside back cover) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 74 Band November 1977 November Part 4 Deel THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES PART 6 AMPHIPODA By CHARLES GRIFFITHS Cape Town Kaapstad The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular intervals as material becomes available Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000 Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM word uitgegee in dele op ongereelde tye na beskikbaarheid van stof Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 8000 OUT OF PRINT/ UIT DRUK 1, 201-3, 5-8), 3(1-2, 4-5, 8, t.-p.i.), 51-3, 5, 7-9), 6(1, t.—p.i.), 711-4), 8, 9(1-2, 7), 10(1-3), 11(1-2, 5, 7, t-—p.i.), 15(4-5), 24(2), 27, 31(1-3), 33 Price of this part/Prys van hierdie deel R2,20 Trustees of the South African Museum © Trustees van die Suid-Afrikaanse Museum 1977 ISBN 0 908407 22 X Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk., Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES PART 6 AMPHIPODA By CHARLES GRIFFITHS Zoology Department, University of Cape Town (With 10 figures) [MS. accepted 4 August 1977] ABSTRACT Amphipoda recovered during the 1975, 1976 Meiring Naude cruises are listed and species of note discussed. The collection includes two new genera —Jzinkala gen. nov. and Valettiella gen. nov. (Lysianassidae)—represented by J. fihla sp. nov. and V. castellana sp. nov. Three further species are recorded from southern Africa for the first time and Bathyamaryllis conocephala (K. H. Barnard) is redescribed. CONTENTS PAGE Introduction yi. ae ee LO> IPiStrofaspeciesi 49 ate ee a LOO Systematic discussion. . . 108 Acknowledgements . . . 123 References Aa ahaa hi eB) 128) INTRODUCTION The material described herein is derived from collections made by the staff of the South African Museum during exploratory cruises by the R.V. Meiring Naude in 1975 and 1976. These cruises are of particular interest since they have investigated deeper waters off Natal which have previously received scant attention, despite their potential zoogeographical interest. A full listing of station data from both cruises has been presented elsewhere (Louw 1977), but few of these samples in fact used gear suitable for the capture of benthic amphipods. Although the number of amphipods recovered was limited, the proportion of new or unusual species in the collection bears witness to the potential for further exploration of this area. All material described below has been deposited in the South African Museum, Cape Town. 105 Ann. S. Afr. Mus. 74 (4), 1977: 105-123, 10 figs. 106 ANNALS OF THE SOUTH AFRICAN MUSEUM LIST OF SPECIES SM Station Numberof Number Individuals Suborder GAMMARIDEA Family Acanthonotozomatidae Dikwa acrania Griffiths . ; ‘ : eg ae ; 86 2 Family Ampeliscidae Ampelisca anomala Sars. : ; : : : P } 86 10 Ampelisca brevicornis (Costa) . A : : : ; : 69 1 Ampelisca byblisoides K. H. Barnard Wer oe Ee, Oe 53 4 103 1 Ampelisca palmata K. H. Barnard . es ee Me ee 53 3 Byblis anisuropus Stebbing . ; ; : , : ; : 86 1 Byblis gaimardi (Kroyer) .. ane an Fi eee 60 1 109 3 Family Amphilochidae Hoplopleon medusarum K. H. Barnard . ; : ; =) 103 1 Unguja yaya Griffiths . ; ; : : : : : : 86 1 Family Corophiidae Chevalia aviculae Walker . ; : : : : : ’ 86 77 103 2 Concholestes armatus Griffiths . . . . 2 Ts hae 31 3 78 31 86 4 103 34 109 13 Gammaropsis afra Stebbing ene Set ast Ly ean 86 4] 91 3 103 32 109 2 Photis uncinata K. H. Barnard . 5 , : : : : 86 26 Unciolella spinosa Griffiths ath Me alt , Neh Ml 60 1 69 1 86 8 109 1 Family Dexaminidae Atylus homochir Haswell . , : ; ; E é J) £09 1 Lepechinella occlo J. L. Barnard at Pee ae i ecauh esc 60 1 Family Eusiridae : ns ie ee Pe es 86 9 _ Dautzenbergia mee (Chevreux) : . ‘ 5 : 60 1 107 4 Family Gammaridae Maera inaequipes (Costa) i Oe ae 86 6 Maera mastersi (Haswell) . ; : . : ; : : 86 2 Family Haustoriidae Urothoe elegans Bate . : : : : : : 53 1 Urothoides inops J. L. Barnard . . : : s ; : 86 1 Family Ischyroceridae Cerapus tubularis Say . : , : ; ; : , 5-408 26 Family Leucothoidae 3 Leucothoe dolichoceras K. H. Barnard . $ 5 ; F 103 2, Leucothoe spinicarpa (Abildgaard) . : E : : ; 86 12 THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES Family Liljeborgiidae Liljeborgia epistomata K. H. Barnard Family Lysianassidae Amaryllis macrophthalma Haswell Bathyamaryllis conocephala (K. H. Barnard) . Hippomedon longimanus (Stebbing) . Hippomedon onconotus (Stebbing) Izinkala fihla gen. et sp. nov. Lepidepecreum clypeatum Chevreux . Lepidepecreum twalae Griffiths . Lysianassa ceratina Valettiella castellana gen. et sp. nov. Family Oedicerotidae Perioculodes longimanus (Bate & Westwood) . Family Phoxocephalidae Heterophoxus opus Griffiths Family Podoceridae Laetmatophilus purus Stebbing Family Stegocephalidae Stegocephaloides australis K. H. Barnard Family Stenothoidae Parametopa grandimana Griffiths Proboloides rotunda (Stebbing) . Superfamily TALITROIDEA Family Phliantidae Plioplateia triquetra K. H. Barnard . Suborder CAPRELLIDEA Family Aeginellidae Eupariambus fallax K. H. Barnard Pseudoprotella phasma (Montagu) Family Phtisicidae Phtisica marina Slaber 107 SM Station Number of Number 86 Individuals y SSS SS So Sa wwe seme ~] 0O — W 108 ANNALS OF THE SOUTH AFRICAN MUSEUM SYSTEMATIC DISCUSSION Family Acanthonotozomatidae Dikwa acrania Griffiths, 1974 Fig. | Dikwa acrania Griffiths, 1974: 266, fig. 2. Material SM 86, 2 specimens Remarks This record is only the second of this interesting species and extends its range considerably to the north. Further drawings of certain appendages are provided here to supplement the original description. These reveal a number of interesting features previously unnoticed. The surface of the mandibular molar is strongly triturative and projects a considerable distance from the body of the mandible, the apical margin is formed into a sharp chitinized ridge. Article 2 of pereiopods 3 and 4 is posteriorly keeled on inner and outer margins, leaving a Fig. 1. Dikwa acrania Griffiths, 1974. Ovigerous female, 4,5 mm. A. Pereiopod 4 with dactyl enlarged. B. Medial view of pereiopod 5. C. Mandible. D. Telson. ; THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 109 deep groove posteriorly, article 4 is acutely produced distally and the dactyl bears small chitinized teeth. The fifth perelopod was erroneously described as missing in the original description. It is, in fact, grossly reduced with article 2(?) apparently fused to, and partially covered by, an enlarged coxa 7; the remainder of the appendage consists of two small articles bearing short anterior spines. This remarkable feature is somewhat similar to that found in Tetradeion (Stegocephalidae) and should be incorporated into the generic definition of Dikwa. Distribution Endemic, Still Bay to Zululand 200-550 m. Family Dexaminidae | Lepechinella occlo J. L. Barnard, 1973 Fig. 2 Lepechinella occlo J. L. Barnard, 1973: 21, figs 8-9. Material SM 60, | specimen SM 86, 9 specimens Remarks The present material differs only marginally from that described by Barnard. Thus in South African specimens coxae | and 2 are more distinctly bifid, the accessory teeth on pereon segments 6 and 7 and pleon segments |-3 somewhat more distinct, and the epimeral setae less marked. These differences cannot be considered of taxonomic significance, particularly in the light of similar size-related differences reported by Barnard (1973). Distribution New Zealand, 721-860 m; Natal 550-810 m. This record is the first from southern Africa. Family Eusiridae Dautzenbergia grandimanus (Chevreux, 1887) Fig. 3 Parapleustes megachir Walker, 1897: 230, pl. 18 (fig. 4). Sympleustes megachir: Stebbing, 1906: 317. Stephensen, 1944: 5, fig. 1. Sympleustes grandimanus: Sexton, 1909: 857, pl. 90 (figs 8-32). Sympleustes (Dautzenbergia) grandimana: K. H. Barnard, 1937: 158. Dautzenbergia grandimanus: J. L. Barnard, 1961: 106. Material SM 60, | specimen SM 107, 4 specimens 110 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 2. Lepechinella occlo J. L. Barnard, 1973. Male, 6 mm. A. Lateral aspect. B. Accessory flagellum. C. Mandible. D-—E. Coxae |, 2. F. Third pleonal epimeron. G. Uropod 3. H. Telson. THE SOUTH AFRICAN MUSEUM'S ME/RING NAUDE CRUISES 11] A Fig. 3. Dautzenbergia grandimanus (Chevreux, 1887). Ovigerous female, 8,5 mm. A. Upper lip. B. Lower lip. C. Mandible. D. Maxilla 1. E. Maxilliped. F—G. Gnathopods 1, 2. H. Uropod 3 and telson. I. Telson. 112 ANNALS OF THE SOUTH AFRICAN MUSEUM Remarks The genus Dautzenbergia has recently been revived by J. L. Barnard (1961) to accommodate species of ‘“Sympleustes’ with a partially cleft telson. Of the three species in the genus D. grandimanus and D. megacheir appear to be indistinguishable. The present material correlates closely with the descriptions given by Sexton (1909) and Stephensen (1944). Distribution North Atlantic, Indian Ocean. This is the first record from southern Africa. Family Haustoriidae Urothoides inops J. L. Barnard, 1967 Fig. 4 Urothoides inops J. L. Barnard, 1967: 23, figs 8-10. Material SM 86, | specimen Remarks There can be no doubt that the present material is synonymous with that illustrated by J. L. Barnard (1967). This unusual species can readily be dis- tinguished by its broad, flat, down-turned rostrum, which covers the anterior end of the body and deflects antenna 2 laterally. Pereiopods 3-5 are strongly developed and have article 2 thickened by glandular tissue. The mouthparts, and particularly the mandibles, are extremely large relative to the body, which is almost as broad as long. Distribution California, 2 700m; Natal 550m. This is the first record of this species from the southern African region. Family Lysianassidae Bathyamaryllis conocephala (K. H. Barnard, 1925) igi) Amaryllis conocephalus K. H. Barnard, 1925: 324. Bathyamaryllis conocephala: K. H. Barnard, 1940: 441. Material SM 61, 2 specimens (including neotype SAM-—A13657) SM 78, | specimen SM 109, | specimen SM 103, fragment SM 60, 2 specimens THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 113 Fig. 4. Urothoides inops J. L. Barnard, 1967. Female, 2 mm. A. Ventral view of rostrum. B. Mandible. C. Lower lip. D. Maxilliped. E. Gnathopod |. F-H. Pereiopods 3, 4, 5. I. Uropod 3. J. Telson. 114 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 5. Bathyamaryllis conocephala (K. H. Barnard, 1925). Female, 5 mm. A. Lateral aspect. B. Head and gnathopod 1. C. Mandible. D. Maxilla 1. E. Maxilliped. F. Gnathopod 2. G-—H. Uropods 2, 3. I. Telson. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES HUES) Remarks This species has hitherto been known only through K. H. Barnard’s somewhat scanty and unfigured description of the holotype female. When an attempt was made to obtain this specimen from Barnard’s type collection it was found to have completely decomposed. A neotype has hence been selected from the present material and is shown in Figure 5. The bevelled off antero-lateral angles of the head clearly identify this species and distinguish it from the allied B. rostrata (Chevreux), in which the lateral lobes of the head are large and vertically truncated. The unusual form of the head in these two species is quite distinct from that in the other members of the genus—B. haswelli (Stebbing) and B. perezii Pirlot. Distribution Endemic, Cape Peninsula to Natal, 680-1 300 m. Izinkala gen. nov. Diagnosis Head and coxa | both very small, largely concealed by enlarged coxa 2; epistome and upper lip inconspicuous; mandible with distinct non-dentate incisor, palp attached distally; maxillae poorly developed; maxilliped lacking plates, palp with minute article 4; gnathopod | slender, minutely subchelate, articles 3 and 5 elongate; gnathopod 2 chelate; uropod 3 reduced, enclosed within urosomite 3, outer ramus with small article 2; telson entire. Type species Izinkala fihla sp. nov. Relationships This remarkable genus appears quite discrete from any known form. The greatly reduced mouthparts are of particular note. A comparable degree of reduction occurs in Kerguelenia Stebbing, but here the mandible lacks both incisor and molar, while the maxilliped retains small plates and a normal palp of four articles. The two genera, however, differ grossly in other respects, such as the form of gnathopod | and relative prominence of coxa |, although other notable similarities with certain species of Kerguelenia occur, for example the enclosure of uropod 3 within urosomite 6 in K. borealis Sars. Izinkala gen. nov. also bears a certain resemblance to various genera in the Lepidepecreum group. Thus species such as L. clypodentatum J. L. Barnard show carinate peduncular — segments in antenna |, similar gnathopods | and 2 and enlarged article 2 of pereiopod 5. Clepidecrella J. L. Barnard lacks certain of these features but has reduced maxillipedal plates, a short uropod 3 and entire telson. Despite these features, /zinkala gen. nov. remains quite distinct with its grossly reduced coxa | and forward projecting, enlarged coxa 2. 116 ANNALS OF THE SOUTH AFRICAN MUSEUM Izinkala fihla sp. nov. Figs 6-7 Description (of ?female 3,2 mm) Head shorter than pereon segment |, post-antennal angles acutely produced; peduncle of antenna | large, article | acutely produced, article 2 overhanging remainder of antenna as a broad lobe with lateral flanges between which distal articles originate, flagellum 4-articulate, article 1 broad, posteriorly fringed with long aesthetascs, accessory flagellum 3-articulate; antenna 2 with article 1 sub- circular, apparently fused to head, gland cone prominent, flagellum 3-articulate; epistome and upper lip inconspicuous; mandible with non-dentate incisor, spine row absent, molar apparently large and smooth; maxillae not satisfactorily resolved, minute; maxilliped completely lacking both inner and outer plates, palp article 4 reduced. Coxa | small and thickened by muscle fibres, completely concealed by large forward-projecting coxa 2, margins of coxae 2-4 of the two sides abutting ventrally, the small gap below the head closed by projecting lobe of article 2 of antenna 1; gnathopod | slender, elongate, articles 3 and 5 each as long as 2, palm slightly oblique, defined by a single strong spine which projects into a pocket formed within the thick, highly chitinized dactyl, which bears sharp transparent lateral flanges; gnathopod 2 chelate; article 2 of pereiopod 3 rotund, article 4 greatly widened posteriorly; pereiopod 4 similar in structure to 3 but longer; pereiopod 5 with article 2 elongate and produced distally as far as tip of article 4, articles 4 and 5 acutely lobed posteriorly. Pleonal epimera | and 2 smoothly rounded, 3 rounded-quadrate; pleon segment 4 with a rounded dorsal carina overhanging pleon segment 5; uropods | and 2 strongly spinose dorsally, rami equal; pleon segment 6 overhanging uropod 3 and telson above and to the sides; uropod 3 very small, outer ramus with a spinose article 2 overhung by dorsal projection of article 1, inner ramus shorter, evenly tapering; telson elongate, entire, bearing strong dorsal and terminal spines. Holotype SAM-A13659, ?female, 3,2 mm. Type locality SM 86, 27°59,5’S 32°40,8’E, 550 m, 22 May 1976. Material SM 86, 4 specimens SM 103, 1 specimen Valettiella gen. nov. Diagnosis Antenna | geniculate, articles 2 and 3 slender and elongate; article | of antenna 2 greatly enlarged; upper lip and epistome inconspicuous; mandibular THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES LF Fig. 6. Izinkala fihla gen. et sp. nov. Female 3,2 mm. A. Lateral aspect. B. Antennae | and 2. C. Mandible. D. Maxilliped E-F. Unidentified mouthparts, probably maxilla 1. 118 ANNALS OF THE SOUTH AFRICAN MUSEUM G Fig. 7. Izinkala fihla gen. et sp: nov. Female, 3,2 mm. A. Gnathopod | with palm enlarged. B. Gnathopod 2. C-—D. Pereiopods 3, 5. E, Lateral view of urosome. F. Uropod 3. G. Telson. THE SOUTH AERICAN MUSEUM'S MEIRING NAUDE CRUISES 119 molar triturative, incisor strongly toothed, palp attached level with molar; inner plate of maxilla | setose medially; coxae | and 2 rectangular and equally elongate; gnathopod |! weakly chelate; gnathopod 2 subchelate; telson short, cleft. Type species Valettiella castellana sp. nov. Relationships The strongly dentate mandibular incisor and unspecialized gnathopods of this genus place it in a distinct group presently comprising Valettia Stebbing, 1888 and Valettiopsis Holmes, 1908. The new form, however, shows features intermediate between those defining Valettia and Valettiopsis, as well as a number of distinct characteristics of its own. Thus Valettia has the outer plate of the maxilliped apically produced and coxae | and 2 similarly short and broad, while Valettiopsis has a normal maxilliped but coxa 2 elongate and almost covering a reduced coxa |. Valettiella gen. nov. not only combines a normal maxilliped with elongate and equal coxa | and 2, but also differs from both the above genera in its unusual geniculate antenna i with elongate peduncle, enlarged article | of antenna 2, widened and castelloserrate article 2 of posterior pereiopods and shortened uropod 3 with reduced inner ramus. Valettiella castellana sp. nov. | Figs 8-9 Description (of ovigerous female, 4 mm) Head as long as first two pereon segments, ocular lobes acute, eyes weak; antenna | with article | elongate, article 2 almost as long as 1, normally folded back into ventral groove of article |, flagellum 9-articulate, accessory flagellum uni-articulate; article 1 of antenna 2 acutely produced into an enlarged shield projecting just below ocular lobe of head, gland cone prominent, flagellum 5-articulate; mandible with 3-articulate palp, incisor with 6 strong teeth, lacinia mobilis with 4 teeth, spine row of 8 spines, molar large and strongly ridged; maxilla | with bi-articulate palp projecting beyond tip of outer plate, palp with 7 terminal spine teeth, outer plate bearing 8 strong serrate spines, inner plate setose medially; inner plate of maxilla 2 strongly setose medially; maxilliped with powerful 4-articulate palp, outer plate apically rounded, bearing eight medial spine teeth. Coxae 1-3 equally elongate, | and 2 with small postero-distal tooth; gnathopod | weakly chelate, article 5 considerably larger than 6; gnathopod 2 with article 3 only slightly elongate, articles 5 and 6 subequal, palm oblique; pereiopods 3-5 progressively longer, article 2 greatly lobed posteriorly, anterior margin spinose, posterior margin strongly castelloserrate. 120 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 8. Valettiella castellana gen. et sp. nov. Ovigerous female, 4mm. A. Lateral aspect. B. Antenna | with accessory flagellum enlarged. C. Antenna 2. D. Mandible. E. Lower lip. F. Maxilla 1. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES U2 B Fig. 9. Valettiella castellana gen. et sp. nov. Ovigerous female, 4 mm. A. Maxilla 2. B. Maxilliped. C. Gnathopod 1. D. Gnathopod 2 with palm enlarged. E. Pereiopod 5. F—H. Uropods 1, 2, 3. I. Telson. | DD ANNALS OF THE SOUTH AFRICAN MUSEUM Pleon segments |, 2, 4 and 5 each bearing a single medio-dorsal tooth, pleonal epimera quadrate postero-distally; rami of uropods | and 2 equal, strongly spinose dorsally and apically; uropod 3 considerably shorter than | and 2, outer ramus with a small article 2, inner ramus about half size of outer and closely appressed to its medio-ventral surface; telson short, cleft, each lobe bearing a large terminal spine and three small setae. Holotype SAM-A13658, ovigerous female, 4 mm, unique. Type locality SM 86, 27°59,5’S 32°40,8’E, 550 m, 22 May 1976. Family Stenothoidae Parametopa grandimana Griffiths, 1974 Fig. 10 Parametopa grandimana Griffiths, 1974: 324, fig. 18. Material SM 86, 13 specimens SM 103, |! specimens Fig. 10. Parametopa grandimana Griffiths, 1974. Male, 4,5 mm. A. Gnathopod 1. B. Gnathopod 2. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES IZ Remarks The above material includes adult males of up to 4,5 mm which show gnathopods somewhat different to those described by Griffiths (1974). In terminal males the palm of gnathopod | is defined by two small spines and the dactyl bears three setae. Article 6 of gnathopod 2 is somewhat more elongate than is typical of smaller males and the palm is undefined, while the dactyl becomes greatly thickened terminally. Distribution Endemic, Still Bay to Natal 200-680 m. ACKNOWLEDGEMENTS This project was made possible through the generosity of the staff of the South African Museum, who not only lent the author the material but also provided excellent library facilities. Iam indebted to the South African Council for Scientific and Industrial Research for financial support. My thanks also to Mrs S. Hardman who kindly typed the manuscript. REFERENCES BARNARD, J. L. 1961. Gammaridean Amphipoda from depths of 400 to 6000 meters. Galathea Rep. 5: 23-128. BARNARD, J. L. 1967. Bathyal and abyssal gammaridean Amphipoda of Cedros Trench, Baja California. Bull. U.S. natn. Mus. 260: 1-205. BARNARD, J. L. 1973. Deep-sea Amphipoda of the genus Lepechinella (Crustacea). Smithson. Contr. Zool. 133: 1-31. BARNARD, K. H. 1925. Contributions to the crustacean fauna of South Africa. 8. Further additions to the list of Amphipoda. Ann. S. Afr. Mus. 20: 319-380. BARNARD, K. H. 1937. Amphipoda. Scient. Rep. John Murray Exped. 1933-34 5: 131-201. BARNARD, K. H. 1940. Contributions to the crustacean fauna of South Africa. 12. Further additions to the Tanaidacea, Isopoda, and Amphipoda with keys for the identification of hitherto recorded marine and freshwater species. Ann. S. Afr. Mus. 32: 381-543. GRIFFITHS, C. L. 1974. The Amphipoda of southern Africa, Part 4. The Gammaridea and Caprellidea of the Cape Province east of Cape Agulhas. Ann. S. Afr. Mus. 64: 251-336. Louw, E. 1977. The South African Museum’s Cruises on the R/V Meiring Naude off the east coast of southern Africa. 1. Station data, 1975, 76. Ann. S. Afr. Mus. 8: 147-159. STEBBING, T. R. R. 1906. Amphipoda 1. Gammaridea. Tierreich 21: I-XXXIX, 1-806. SEXTON, E. W. 1909. Notes on some Amphipoda from the north side of the Bay of Biscay: Families Pleustidae and Eusiridae. Proc. zool. Soc. Lond. 1909: 848-879. . STEPHENSEN, K. 1944. Crustacea Malacostraca, VIII: (Amphipoda IV). Danish Ingolf-Exped. 3: 1-51. WALKER, A. O. 1897. On some new species of Edriophthalma from the Irish Seas. J. Linn. Soc. 26: 226-232. 6. SYSTEMATIC papers must conform to the International code of zoological nomenclature (particularly Articles 22 and 51). Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov., etc. An author’s name when cited must follow the name of the taxon without intervening punctuation and not be abbreviated; if the year is added, a comma must separate author’s name and year. The author’s name (and date, if cited) must.be placed in parentheses if a species or subspecies is transferred from its original genus. The name of a subsequent user of a scientific name must be separated from the scientific name by a colon. Synonymy arrangement should be according to chronology of names, i.e. all published scientific names by which the species previously has been designated are listed in chronological order, with all references to that name following in chronological order, e.g.: Family Nuculanidae Nuculana (Lembulus) bicuspidata (Gould, 1845 Figs 14-15A Nucula (Leda) bicuspidata could, get: O Sil Leda plicifera A. Adams, 1856: Laeda bicuspidata Hanley, 1859: ae: pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b). Nucula largillierti Philippi, 1861: 87. Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. Note punctuation in the above example: comma separates author’s name and year semicolon separates more than one reference by the same author full stop separates references by different authors figures of plates are enclosed in parentheses to distinguish them from text-figures dash, not comma, separates consecutive numbers Synonymy arrangement according to chronology of bibliographic references, whereby the year is placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable. In describing new species, one specimen must be designated as the holotype; other speci- mens mentioned in the original description are to be designated paratypes; additional material not regarded as paratypes should be listed separately. The complete data (registration number, depository, description of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.: Holotype SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973. Note standard form of writing South African Museum registration numbers and date. 7. SPECIAL HOUSE RULES Capital initial letters (a) The Figures, Maps and Tables of the paper when referred to n the text e.g. °... the Figure depicting C. namacolus ...’; ‘. . . in C. namacolus (Fig. 10)...” (b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded by initials or full names e.g. DuToit but A.L.du Toit; Von Huene but F. von Huene (c) Scientific names, but not their vernacular derivatives e.g. Therocephalia, but therocephalian Punctuation should be loose, omitting all not strictly necessary Reference to the author should be expressed in the third person Roman numerals should be converted to arabic, except when forming part of the title of a book or article, such as “Revision of the Crustacea. Part VIII. The Amphipoda.’ Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial capital letter, provided the same generic name is used consecutively. Name of new genus or species is not to be included in the title: it should be included in the abstract, counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts. CHARLES GRIFFITHS THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES PART 6 AMPHIPODA 4 \/ =a VOLUME 74 PART 5 JANUARY 1978 | ISSN 0303-2515 MUS. adit ZOOL. LIPRARY APR 2 281078 HARVARD UNIVERSI ry ANNALS OF THE SOUTH AFRICAN MUSEUM CAPE TOWN INSTRUCTIONS TO AUTHORS 1. MATERIAL should be original and not published elsewhere, in whole or in part. 2. LAYOUT should be as follows: (a) Centred masthead to consist of Title: informative but concise, without abbreviations and not including the names of new genera or species Author’s(s’) name(s) Address(es) of author(s) (institution where work was carried out) Number of illustrations (figures, enumerated maps and tables, in this order) (b) Abstract of not more than 200 words, intelligible to the reader without reference to the text (c) Table of contents giving hierarchy of headings and subheadings (d) Introduction (e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents (f) Summary, if paper is lengthy (g) Acknowledgements (h) References (i) Abbreviations, where these are numerous 3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced with 2,5 cm margins all round. First lines of paragraphs should be indented. Tables and a list of legends for illustrations should be typed separately, their positions indicated in the text. All pages should be numbered consecutively. Major headings of the paper are centred capitals; first subheadings are shouldered small capitals; second subheadings are shouldered italics; third subheadings are indented, shouldered italics. Further subdivisions should be avoided, as also enumeration (never roman numerals) of headings and abbreviations. Footnotes should be avoided unless they are short and essential. Only generic and specific names should be underlined to indicate italics; all other marking up should be left to editor and publisher. 4. ILLUSTRATIONS should be reducible to a size not exceeding 12 < 18 cm (19 cm including legend); the reduction or enlargement required should be indicated; originals larger than 35 x 47 cm should not be submitted; photographs should be rectangular in shape and final size. A metric scale should appear with all illustrations, otherwise magnification or reduction should be given in the legend; if the latter, then the final reduction or enlargement should be taken into consideration. All illustrations, whether line drawings or photographs, should be termed figures (plates are not printed; half-tones will appear in their proper place in the text) and numbered in a single series. Items of composite figures should be designated by capital letters; lettering of figures is not set in type and should be in lower-case letters. The number of the figure should be lightly marked in pencil on the back of each illustration. 5. REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: ‘Smith (1969) describes...’ ‘Smith (1969: 36, fig. 16) describes...’ ‘As described (Smith 1969a, 1969b; Jones 1971)’ ‘As described (Haughton & Broom 1927)...’ ‘As described (Haughton et al. 1927)...’ Note: no comma separating name and year Dagination indicated by colon, not p. names of joint authors connected by ampersand et al. in text for more than two joint authors, but names of all authors given in list of references. (b) Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, b, etc. to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a). For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list o, Scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number (only if independently paged) in parentheses, pagination (first and last pages of article). Examples (note capitalization and punctuation) BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140. FISCHER, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. gén. 74: 627-634. Koun, A. J. 1960a. Ecological notes on Genus (Mollusca: Gastropoda) i in the Trincomalee region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320. Konn, A. J. 19606. Spawning behaviour, eee masses and feral development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 51. THIELE, J. 1910. Mollusca: B. Calveieechece? Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. (continued inside back cover) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 74 Band . January 1978 Januarie Rant ee Deck THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES PART 7 MARINE ISOPODA By BRIAN KENSLEY Cape Town Kaapstad The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular intervals as material becomes available Obtainable from the South African Museum, P.O. Box 61, Cape Town Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM word uitgegee in dele op ongereelde tye na beskikbaarheid van stof Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad OUT OF PRINT/UIT DRUK 1, 201-3, 5-8), 3(1-2, 4-5, 8, t.—p.i.), 5(1-3, 5, 7-9), 6(1, t.—p.i.), 711-4), 8, 911-2, 7), 10(1-3), 11(1-2, 5, 7, t—p.i.), 15(4—5), 24(2), 27, 311-3), 32(5), 33 Price of this part/Prys van hierdie deel R3,10 Trustees of the South African Museum © Trustees van die Suid-Afrikaanse Museum 1978 ISBN 0 908407 34 3 Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk., Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES PART 7 MARINE ISOPODA By BRIAN KENSLEY South African Museum, Cape Town (With 21 figures and 1 table) [MS. accepted 9 August 1977] ABSTRACT Excluding the Anthuridea, twenty-one species of isopods are recorded from deep water off the Natal coast, including fourteen new records. The latter include one new genus, Spinarcturus, and ten new species. These are Microarcturus biserialis, M. youngi, Neastacilla longispina, Spinarcturus natalensis, Excirolana bicornis, Cirolana caeca, Serolis brinki, Haploniscus gernekei, Eurycope glabra, and Ilyarachna wolffi. Five endemic South African species are also recorded, as well as Bathycopea typhlops, premouyy only known from the coast of Ireland. CONTENTS PAGE Introduction : : ; : VTS Species list . , : é Rape Me Systematic disenssion! ; : ; 128 General discussion . : : Wee | 'S)5) Acknowledgements . ; 4 ee boy) References . 4 : ‘ : geri, INTRODUCTION The present paper forms part of a series, based on material collected by the South African Museum during the 1975, 1976 cruises off the Natal coast on the R/V Meiring Naude. For the scope of, and the background to this pro- gramme, as well as for all station data, the reader is referred to Louw (1977). The Anthuridea, of which about seven species were collected, are not dealt with, and will form the subject of a future paper. The following abbreviations are used throughout this paper: SAM—South African Museum catalogue number; SM—Meiring Naude Station numbers; TL—total length; ovig.—ovigerous; juv.—juvenile. 125 Ann. S. Afr. Mus. 74 (5), 1978: 125-157, 21 figs, 1 table. 126 ANNALS OF THE SOUTH AFRICAN MUSEUM 3° 10 20 30. 40 50 B22 10° 20° 30° 40° 0" skh 10 20! | Ig 40° | g : SES Bh gs 50 ae 0 Sa ee Kosi Bay 56 A (oan) 27° 2 Ly Eh “4 5 3 6, 28 oy) , 10 eee ro) 63 > ae on 12 Gio tt 2 : " as loo Qh 65 | °70 4 : y | | | 15 78 1g [+20 30 6 77 17 75 % Ih i) °73 80 : 40 40 | Bo 2 “e 2 88 50) 0 ? 26 87 (Neal | 28 ee AR 28° | | Ses | 31 ky) go 089634 10° 10 ott on * 597 93 92 6 9°37 36 a 20 “38 639 194 191 105 102 Fa 3d “103 fo 197196 “108 ie i6 Q 4) 199 0 48 Richards B " dj Icnards Day mn 50’ 12 } 48 47. 2 m 48 49 14 10 “113 50 ho 15 20 20' 30h 80! 40) Sh 40 | 45 e J 450. a gy, DURBAN Y] Yi 50! 3 io ay 30 “40 50" 32° oo 2 30 10" 50 33° 0 0 Fig. 1. Map showing Meiring Naude 1975, 1976 stations. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES Ail SPECIES LIST Material not identified to specific level is either damaged or immature. SM Station 3d ojo) ovig. 22 Juvs SUBORDER VALVIFERA Family Arcturidae * Microarcturus biserialis sp. nov. ; ; , OY 3 — <= = Microarcturus oudops (Barnard) . : : . 103 2 4 4 — * Microarcturus youngi sp. Nov. . : ; an 86 12 5 3 +50 103 8 10 — 37 * Neastacilla longispina sp. nov. . ; : 5 &© 2), — 3 3 * Spinarcturus natalensis gen. et sp. nov. . 5 ti) 4 DD, 3 D 103 ol — — 2 SUBORDER FLABELLIFERA Family Cirolanidae *Excirolana bicornis sp. nov. ‘ : : 5 68 4 6 7 *Cirolana caeca sp. nov. ‘ ; ; : eS 2 4 4 — Cirolana imposita Barnard . ; ‘ : AG 1 — — 23 1 — — — Family Sphaeromatidae *Bathycopea typhlops Tattersall . ' : 5 108 2 3 1 — Family Serolidae *Serolis brinki sp. nov. . Sy Pe a, : 5» 108 2 3 1 — Family Cymothoidae Nerocila sp. . : : : : : : 5 OF — — — 1 SUBORDER GNATHIIDEA Family Gnathiidae Gnathia sp. . : : : : ‘ : 5 sh) +20 + 20 +20 +20 SUBORDER ASELLOTA Family Stenetriidae *Stenetrium abyssale Wolff . : : : . 86 1 — 1 — Stenetrium dalmeida Barnard : : ; a SO 1 —- — — Stenetrium dagama Barnard Family Haploniscidae * Haploniscus gernekei sp. nov. . : 3 . 86 3 3 — — 103 9 4 — —- Family Ischnomesidae Ischnomesis sp. j : i : : ; 4 6 2 fragments Family Eurycopidae *Furycope glabra sp. nov. . , : ‘ ny 60) — 1 = — 103 1 — — — Family Dendrotionidae * Acanthomunna spinipes (VanhOffen) . P eX) 4 8 3 — 103 y) 1 — — Family Ilyarachnidae *Ilyarachna wolffi sp. nov. . : : : TP SO 4 _- _ == 103 2 — — — Ilyarachna sp. : ; : é f 3 . 103 — Z, — — *New Record 128 ANNALS OF THE SOUTH AFRICAN MUSEUM SYSTEMATIC DISCUSSION Family Arcturidae Microarcturus biserialis sp. nov. Fig. 2 Description Body elongate, slender, widest at pereionites II and III; head fused with pereionite I, anterior margin concave, anterolateral corners acute, eyes lacking, one pair small submedian tubercles anterior to submedian pair dorsal spinose processes; lateral margin of head incised. Coxae of pereionites II and III acute, of pereionites III to VII with two or three smaller points. Each pereionite possessing dorsal submedian pair of spinose processes and broader apically acute lateral process. Pleon consisting of three fused segments plus pleotelson, former each marked by a pair of submedian spines; pleotelson apically acute, with acute lateral tooth at about midpoint. Antennule not reaching midpoint of second antennal peduncle segment. Three distal segments of antennal peduncle becoming progressively longer. Mandibles, maxillae and maxilliped typical of the genus. Pereiopod I shorter than following pereiopods, with numerous serrate spine-like setae. Pereiopods II to IV slender, all segments with elongate simple setae on ventral surface. Pereiopods V to VII more robust, dactyli, propodi, carpi, meri and distal part of ischia with dense ventral pile of short setules, propodi and carpi with few fringed ventral spines. Penial process with distal third of rami separate, tips truncate. Pleopod 1 with endopod about half length of exopod, distally rounded, bearing simple setules; exopod apically curved, with only two distal fringed setae; about twenty simple spines on outer margin, broad furrow on anterior face. Pleopod 2 rami similar, distally rounded/truncate, stylet on endopod only slightly longer than ramus, proximally stout, tapering distally, with small spines on inturned distal flanges. Material Holotype SAM-A15467 1 3 TL5,9 mm SM 109 28°41,0’S 32°36,8’E 1 300 m 233 TL5,9 mm SM 109 Remarks The structure of the male pleopods | and 2, the mouthparts, and pereional appendages leave no doubt that the present species belongs to the genus Microarcturus, in spite of the lack of eyes and the relatively short exopod of pleopod 1. The dorsal sculpturing resembles that found in the males of M. ornatus Kensley, 1975, and M. similis (Barnard), but neither of these has THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 129 Boe F Fig. 2. Microarcturus biserialis sp. nov. A. 3 dorsal view. B. @ lateral view. C. Pleopod 1 g. D. Maxilliped. E. Pleopod 2 3. F. Pereiopod I. G. Pereipod II. H. Pereiopod VII. I. Penis. 130 ANNALS OF THE SOUTH AFRICAN MUSEUM the double row of spines as well developed on the last three pereionites. The specific name ‘biserialis’ is derived from the double row of prominent dorsal spines on the pereion and pleon. Microarcturus oudops (Barnard) Fig. 3 Neoarcturus oudops Barnard, 1914: 214, pls 18c, 19b; 1920: 397; 1940: 508. Nordenstam, [YB Ils. Previous records Off Cape Point. Material SAM-A15472 2 gg 492 4 ovig. 22 SM 103 Remarks Barnard (1914) based the new genus Neoarcturus on several features, including the non-geniculate body, the non-elongate fourth pereionite, distinct coxae, pleon of four fused segments, and the antennal flagellum of three articles. All these features apply equally to Microarcturus. Kensley (1975) characterized Fig. 3. Microarcturus oudops (Barnard) Pleopod 1 3. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 131 Microarcturus by incised lateral margins of the head, dorsolateral eyes, first pereionite completely fused to head, two or three pleonal segments anterior to the pleotelson, and the first pleopod of the male with the inner ramus more than half the length of the outer. All these features are found in M. oudops. Further, although Barnard characterized Neoarcturus as being blind and possessing three pairs of oostegites, four pairs of oostegites are present, as are eyes; these latter are, however, very weakly pigmented. There is thus no justification for placing oudops in a genus separate from Microarcturus. The relative length of the inner ramus of the first pleopod of the male cannot be used as a criterion for generic separation as this shows a range from short in M. biserialis to fairly long in M. youngi. The first pleopod of this species is refigured. Microarcturus youngi sp. nov. Fig. 4 Description Female. Body with numerous small tubercles; widest at pereionites II and III. Head with anterolateral corners subacute, with submedian ridges. Eyes large, lateral. Coxae indistinct, of pereionite I ventrally directed, tridentate; coxae of pereionites II and III expanded laterally, convex in outline; coxa of pereionite IV rectangular. Pereionites V to VII shorter and narrower than preceding ones. Pleon consisting of one indistinct and two distinct segments plus pleotelson. Latter broad, terminally subacute, with rounded lateral tooth. Antennule reaching to midpoint of second antennai peduncle segment. Antennal peduncle 4-segmented, two basal segments together equal in length to third segment, latter slender, slightly shorter than fourth segment; flagellum of two articles with slender terminal spine. Mandibles, maxillae and maxilliped typical of the genus. Pereiopod I shorter than following pereiopods, with numerous serrate spine-like setae. Pereiopods II to IV relatively slender, with numerous slender setae. Pereiopods V to VII more robust, propodi, carpi and meri with dense very short setae on ventral surface plus several widely spaced spines, those on propodus barbed. Male. Body parallel-sided, cylindrical, pereionites varying in sculpture from granular to almost smooth. Pleon with pair of large subconical processes at base. Penial rami fused for three-quarters of length, tips diverging. Pleopod 1 basis with rounded proximal flange, and fourteen peg-like processes; inner ramus shorter than outer, distally rounded/truncate; outer ramus with apex produced and bent, numerous spines on outer margin, broad furrow on anterior face, ten plumose setae distally. Pleopod 2 with stylet of endopod stout, cylindrical, slightly convoluted apically. ANNALS OF THE SOUTH AFRICAN MUSEUM 1132 Li, wit” Fig. 4. Microarcturus youngi sp. nov. A. 2 dorsal view. B. 3 lateral view. C. Penis. D. Maxilliped. E. Pleopod 1 g. F. Antennule. G. Pereiopod VII. H. Pereiopod II. I. Pleopod 1 ¢. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 133 Material Holotype SAM-A15465 1 ovig. 2 TL 5,8 mm SM 86 = 27°59,5’S 32°40,8’E 550m Allotype SAM-A15465 1¢ TL6,0mm SM 86 12 gs 592 3ovig. 92 SO0juvs SM 86 SAM-A15466 8 gg 1092 37juvs SM 103 Remarks M. youngi is obviously closely related to the group of species including M. ornatus Kensley, M. dayi Kensley, and M. similis (Barnard), in which the basic plan and sculpture of the males and females are similar. Each, however, may be distinguished by details of the sculpture. The most distinctive feature of M. youngi is the very prominent pair of bosses at the base of the pleotelson in the male. The species is named for Mr D. Young, first officer of the R/V Meiring Naude. Neastacilla longispina sp. nov. Figs 5-6 Description Female. Body slender, elongate. Head and first pereionite fused, fusion indicated by shallow furrow; eyes large, lateral; pair of dorsal spines and few Fig. 5. Neastacilla longispina sp. nov. A. Ovigerous @ lateral view. B. ¢ lateral view. 134 ANNALS OF THE SOUTH AFRICAN MUSEUM small tubercles present. Pereionites I and II with small dorsal spine and spinose lateral process. Pereionite II with single very strong dorsal spine, smaller spine between dorsal spine and spinose lateral process. Pereionite IV elongate, with pair of strong submedian dorsal spines at midpoint, and single slightly hooked median dorsal spine on posterior margin. Pereionite V with few small tubercles and spinose coxa. Pleon consisting of three fused pleonites plus pleotelson, segments indicated by shallow furrows; spinose lateral process on third pleonite. Pleotelson with distal half tapering, apically acutely rounded. Antennule consisting of 3-segmented peduncle, basal segment with small rounded dorsal process; flagellum of single article reaching beyond basal antennal segment. Antenna longer than the total body length, basal segment short, with spinose distal process; following three segments very elongate, slender, flagellum Vines Fig. 6. Neastacilla longispina sp. nov. A. Pereiopod VII. B. Pereiopody (Ge Pereiopod II. D. Pleopod 2 g. E. Antennule 3. F. Pleotelson 3. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 135 short, of three articles, terminal article very short. Mouthparts typical of the genus. Pereiopod I shorter than following pereiopods, dactylus distally rounded/ truncate, with strong terminal serrate spine; all segments bearing numerous simple setae; serrate spines also present on dactylus and propodus. Pereiopods II to IV slender, elongate, dactyli absent, remaining segments bearing elongate simple setae. Pereiopods V to VII becoming shorter posteriorly, more robust than preceding pereiopods, dactyli with two short apical claws, numerous fine setules on all segments except basis, especially concentrated on ventral surfaces. Male. Pereionite IV more elongate than in female; head and pereionites with few scattered tubercles; no elongate dorsal spines as in female. Antennular flagellum relatively longer than in female. Antenna with second peduncular segment with small scattered tubercles. Pleotelson with slight point at lateral midpoint. Pleopod | with rami of equal length, with elongate distal plumose setae; exopod with three elongate proximal setae, endopod with proximal indentation. Pleopod 2 rami of equal length, with distal elongate plumose setae; endopod with stylet at base of inner margin extending well beyond rami, apically acute and slender. Material Holotype SAM-A15459 1 3 TL10,5 mm SM 86 27°59,5’S 32°40,8’E 550 m | Allotype SAM-A15459 19° TL8,3mm SM 86 Ig 292) 3 yuvs SM 86 Remarks The present species bears little resemblance to the three species of Neastacilla hitherto recorded from South African waters, but most closely resembles Astacilla attenuata Hale, 1946, from New South Wales. N. Jongispina differs from the Australian species especially in possessing dorsal elongate spines on the pereion, and in having more elongate antennae. The specific name is derived from the dorsal spines of the female. Spinarcturus gen. nov. Diagnosis Arcturid possessing a geniculate body; first pereionite not fused with the head; dorsolateral eyes; antennae shorter than the total body length; fourth pereionite not elongate; anterior four pairs of pereiopods unspecialized, not differentiated from the posterior three pairs; pleopod | of the male relatively unspecialized, not furrowed on anterior face; stylet of pleopod 2 of the male apically simple. Type species of the genus Spinarcturus natalensis. The generic-name is derived from the numerous spines of the body. 136 ANNALS OF THE SOUTH AFRICAN MUSEUM Spinarcturus natalensis sp. nov. Figs 7-8 Description Male. Body elongate, geniculate, with numerous short fine setules entrapping debris. Head with concave anterior margin, not fused with pereionite J. Latter bearing pair of slender spines. Pereionites II to IV with two pairs of dorsolateral spines. Pleon consisting of three fused segments, first pleonite unarmed, second pleonite with one pair dorsolateral spines, third pleonite with pair strong proximolateral spinose processes, rest of segment smooth, convex, terminally bluntly rounded. Coxae not dorsally visible. Eyes dorsolateral. Antennule 4-segmented, terminal article elongate, bearing aesthetascs. Antenna shorter than body, peduncle consisting of two short proximal segments and two more elongate distal segments, plus flagellum of three articles, slender terminal spine present; all segments with scattered setae. Mandible with tridentate incisor, tridentate lacinia, four setae in row, broad truncate molar process with four stout spines on ventral margin. Maxilla | with three fringed setae on inner ramus, several simple spines on outer ramus. Maxilla 2 with both lobes of outer ramus tipped with two sparsely fringed Fig. 7. Spinarcturus natalensis gen. et sp. nov. A. 3 dorsal view. B. Pleopod 1 g. C. Pleopod 2 3. D. Penis. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 137 setae, inner ramus with about ten fringed setae. Maxilliped with several fringed setae on four distal palp segments and on a endite. Pereiopod I shorter than following pereiopods, dactylus with strong terminal spine, smaller secondary spine, and several fringed spines, propodus with many fringed spines; carpus, merus, and ischium with few ventral fringed spines and numerous fine short setules. LIZZ Ss Za. Fig. 8. Spinarcturus natalensis gen. et sp. nov. A. Pereiopod VII. B. Maxilla 2. C. Mandible. D. Uropod apex. E. Pereiopod I. F. Maxilla 1. G. Maxilliped. H. Antennule. I. Antenna. 138 ANNALS OF THE SOUTH AFRICAN MUSEUM Pereiopods II to IV essentially similar to pereiopods V to VII, latter with more elongate propodi and carpi. Perelopods more or less covered with fine short setules and adhering debris. Rami of penis united almost to apex, latter tapered, finely setose. Pleopod | relatively unspecialized, with proximal indentation in endopod, both rami apically rounded, with numerous plumose setae. Pleopod 2 both rami apically rounded, with plumose setae; stylet of endopod sabre-shaped, almost twice length of ramus, apically acute. Uropod with strong ridge near outer margin, outer ramus small, triangular, inner ramus less than half width of outer, with two subapical spines. Female. Similar to male, but pereionites II to IV wider; broodpouch formed by four pairs of oostegites. Material Holotype SAM-A15473 1 3 TL 6,0 mm SM 86 27°59,5’S 32°40,8’E 550 m Allotype SAM-A15473 1 ovig. 9 TL 8,2 mm SM 86 SOS Z2ZLQ Lowe 2ws SMS SAM-A15474 1g 2juvs SM 103 Remarks Deciding on the generic position of the present species gives rise to some difficulty. Pereionite IV is not elongate, which rules out Astacilla, Neastacilla, Arcturella, Arcturina, Arcturopsis, Parastacilla and Arcturinoides. The head is not fused with the first pereionite, which eliminates Pleuroprion, Antarcturus and Jdarcturus. The antenna is shorter than the body, and has a flagellum of three articles, which rules out Antarcturus, Arcturus, and Dolichiscus. Pleopod 1 in the male is not specialized or furrowed as in Austroarcturus, Microarcturus, Antarcturus and Holidotea. The elongate stylet of the second pleopod of the male is reminiscent of Arcturinoides and Arcturina but does not have a trifid apex as in these latter. The first pleopod with its proximal indentation again resembles that of Arcturina and Arcturinoides. The pereiopods, however, are not differentiated as they are in these genera. The creation of a new genus for this material would thus seem to be warranted. The specific name is derived from the Province of Natal. Family Cirolanidae Excirolana bicornis sp. nov. Figs 9-10 Description Male. Body width about half length, widest at pereionites II and VI. Head with dorsolateral eyes; two elongate slightly curved apically rounded ‘horns’ anterior to eyes, length of ‘horns’ varying with total length. Frontal lamina distally rounded, dorsally visible between antennal bases, proximally narrowed. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 139 —————>——. } \) Ze Fig. 9. Excirolana bicornis sp. nov. A. 6 dorsal view. B. Mandible. C. Maxilla 1. D. Maxilla 2. E. Maxilliped. F. Head of ¢ in lateral view. Clypeus acutely pointed in lateral view. Pereionites unsculptured except for faint, impressed line. All coxae pointed, those of pereionites IV to VII being elongate-acute. Pleotelson triangular, distal margin dentate, apically acute. Antennal peduncle 3-segmented, third segment elongate; flagellum of eight to nine articles. Antenna longer than antennule, peduncle 4-segmented, terminal segment longest, flagellum of fourteen articles. Mandible with 3-segmented palp, middle segment nearly three times length - of basal segment; incisor 4-dentate; setal row of fourteen to sixteen short 140 ANNALS OF THE SOUTH AFRICAN MUSEUM spine-like setae; molar process with numerous small teeth on upper surface. Maxilla 1 with three setae on inner ramus, ten dentate and simple spines on apex of outer ramus. Maxilla 2 with several plumose setae on inner ramus, inner lobe of outer ramus with five setae, outer lobe with three setae. Maxillipedal endite narrow, armed with five plumose setae and single coupling hook; palp 5-segmented, third segment longest and broadest, all segments setose. Pereiopod I with sensory spines on ventral margin of propodus, carpus and merus, carpus very short. Pereiopod VII armed with clusters of serrate spines on propodus, carpus, merus and ischium. Fig. 10. Excirolana bicornis sp. nov. A. Antenna. B. Antennule. C. Pereiopod I. D. Pereiopod VII. E. Pleopod 2 d. _ F. Apex of telson. G. Uropod. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 14] Pleopod 2 with stylet attached to base of endopod, extending well beyond both rami, apically tridentate. Uropod with inner ramus broader than outer, margins serrate, apically narrowed, with several setae and few spines on outer margin. Female. Similar to male, but lacking ‘horns’ on head. Material Holotype SAM-A15469 1 3 TL (excl. horns) 5,6mm SM 86 27°59,5’'S 32°40,8’E 550 m Allotype SAM-A15469 1 @ (with oostegites) 5,5imm SM 86 Paratypes SAM-A15469 3 gg 592 7juvs SM 86 Remarks Sexual dimorphism in Excirolana has been recorded only in one other species, E. kumari Bowman, 1971, from Malaysia. In this species the male bears a pair of triangular horns at the dorsal posterolateral corners of the head, two slender lateral horns on pereionites I and II, plus a mediodorsal spine on pereionite II. The distinctive pair of horns anterior to the eyes on the head of the male (from which is derived the specific name), immediately separates E. bicornis from E. kumari. ; Cirolana caeca sp. nov. Figs 11-12 Description Body width slightly more than half length, strongly convex dorsally, widest at pereionites IV and V, coxae well developed, leaf-shaped; no dorsal sculpture other than impressed line on pereionite VII. Pleon consisting of five segments plus pleotelson, lateral margin of first pleonite overlapped by coxae of last pereionite; lateral margin of fifth pleonite overlapped by fourth; pleotelson wider than long, distally rounded, distal margin crenulate. Eyes absent. Antennule shorter than antenna, third peduncular segment longest, flagellum of five articles. Antenna with terminal peduncular segment longest, flagellum of eleven articles. Mandibular palp 3-segmented, middle segment longest, bearing simple and plumose spines; incisor of four broad teeth; setal row of ten curved spine-like setae; molar process with marginal row of teeth. Maxilla 1 inner ramus short, with three apical plumose setae; outer ramus with several dentate and simple spines. Maxilla 2 inner ramus with five plumose setae, inner lobe of outer ramus with four simple setae, outer lobe with three. 142 ANNALS OF THE SOUTH AFRICAN MUSEUM Maxillipedal endite short, with one coupling hook and four plumose setae; palp 5-segmented, middle segment broadest and longest. Pereiopod I shorter than following pereiopods, carpus short and triangular, ventral margins of propodus, carpus, and merus bearing few sensory spines. Pereiopod VII with serrate spines on ventral and distal margins of segments. Penial processes digitiform. Pleopod 2 male with stylet on inner ramus longer than rami, apically with slender spine and subterminal blunt tooth. Uropod with both rami distally narrowly rounded, inner ramus broad, with sparse plumose setae. Fig. 11. Cirolana caeca sp. nov. A. ¢ dorsal view. B. Maxilliped. C. Maxilla 1. D. Maxilla 2. E. Mandible. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 143 Material Holotype SAM-A15468 1 ¢ TL 3,9 mm SM 78 27°31,6’S 32°50,0’E VSO) tom Allotype SAM-A15468 1 ovig.2 4,2 mm SM 78 Paratypes SAM-A15468 1 3 TL 3,2 mm 5 99 TL 2,8 mm-4,0 mm 3 ovig. 22 TL 3,6 mm-4,0 mm SM 78 Remarks This very distinctive species differs markedly in body shape from other blind species of Cirolana, such as C. californiensis Schultz and C. poissoni Monod. The specific name is derived from the lack of eyes. Fig. 12. Cirolana caeca sp. nov. A. Antenna. B. Antennule. C. Pleopod 2 3. D. Uropod. E. Apex of telson. F. Pereiopod I. G. Pereiopod VII. 144 ANNALS OF THE SOUTH AFRICAN MUSEUM Family Sphaeromatidae Bathycopea typhlops Tattersall Fig. 13 Bathycopea typhlops Tattersall, 1905: 12, 65, pl. 3. Loyola e Silva, 1971: 216, figs 2-3. Schultz, IO7Bs LIS Previous record Off County Kerry, Ireland. Material SAM-A15461 2 33 399 lovig. 2 3,9-5,3 mm SM 103 Remarks This is only the second record of this remarkable species. Tattersall (1905) recorded B. typhlops from five stations off the Irish coast, in depths ranging from about 400 to 830 metres, and in all cases the bottom substrate was fine sand. The length of the adult male and female from Ireland was 5,0 mm, agreeing well with the present material. No differences in proportions or in the appendages could be detected between Tattersall’s and Loyola e Silva’s figures and descriptions, and the present specimens. The huge distance between these two records can only be explained by a lack of sampling of the appropriate substrate. Fig. 13. Bathycopea typhlops Tattersall, dorsal view. Family Serolidae Serolis brinki sp. nov. Figs 14-15 Description Male. Body longer than wide, lacking spines and tubercles on dorsum. Head with anterolateral angles acute, slightly produced, anterior margin THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 145 Fig. 14. Serolis brinki sp. nov. medially emarginate, eyes entirely absent. Coxal plates marked off by sutures on pereionites II to IV, coxal plate of pereionite VII not extending beyond pleural plates of second and third pleonites. Pleotelson with very slight rounded median ridge, more strongly indicated proximally, becoming obsolete distally; small lateral point at uropodal insertion. Antennular peduncle longer than flagellum, latter of twenty articles. Antennal peduncle about equal in length to entire antennule, flagellum of fourteen articles. Left mandible with lacinia tridentate, right mandible with incisor somewhat expanded, with two strong spines and rounded tubercles; lacinia process distally dentate. Maxillipedal palp 3-segmented, distal segment well developed. Pereiopod I, palm of propodus with alternating closely packed slender bifid and broad finely setulose setae; distal margin of carpus with two short setae and few fine setules. Pereiopod II with double row of eight stubby, bristled setae on palm, distal margin with several simple setae. Pereiopod VII slender, elongate, with setae on all segments except basis. Pleopod 2 with distal portion of endopodal stylet about three times longer than proximal portion. Uropod with exopod well developed, oval, shorter than endopod. Colour pattern (when alive): only anterior half of dorsum pigmented red-grey; posterior half white. Pigment present on antennal and antennular peduncles, with white dapples on head and terga. Head and anterior four pereionites with broad hyaline margins. 146 ANNALS OF THE SOUTH AFRICAN MUSEUM Material Holotype SAM-A15460 1 3 TL 8,6mm SM 103 28°31,7’S 32°34,0’E 680 m Remarks This blind species of Serolis closely resembles two previously recorded deep-water species, viz. S. vemae Menzies, 1962, from the North and South Atlantic, and S. menziesi Hessler, 1970, from the South Atlantic off Brazil, but differs from both. Fig. 15. Serolis brinki sp. nov. A. Inner view of mandibles. B. Maxilliped. C. Pleopod 2 ¢. D. Pereiopod I. E. Spines on palm of propodus, pereiopod I. F. Pereiopod II. G. Pereiopod VII. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 147 S. menziesi does not have a lateral spine on the pleotelson, the uropodal exopod is minute and the pleotelson possesses a triangular flattened proximal area. In S. vemae the uropodal exopod is also much smaller than in S. brinki, the pleotelson also possesses a flattened proximal area, but lacks a lateral spine. S. brinki is the first species of the genus recorded in South African waters, and is named for Dr C. van der Merwe Brink, President of the South African Council for Scientific and Industrial Research. Family Stenetriidae Stenetrium abyssale Wolff Stenetrium abyssale Wolff, 1962: 25, figs 1-5. Previous records Kermadec Trench (north-east of New Zealand) and Tasman Sea, 4510- 4540 m. Material SAM-A15470 1 ovig.2 5,3mm 13 3,0mm SM 86 SAM-A15471 12 5.4mm 4 g¢ 5,0 mm-5,8 mm SM 103 < Remarks Wolff (1962) gives the dimensions of 9,9 mm TL ¢ and 8,6 mm TL Q. The present specimens, including an ovigerous female, are all less than 6 mm in length. This is the only difference from the Kermadec material, the appendages and body shape agreeing exactly with Wolff’s description. Family Haploniscidae Haploniscus gernekei sp. nov. Fig. 16 Description Body about twice longer than wide. Anterolateral margins of pereionites and pleotelson with broad hyaline border. Lateral margin of pleotelson continuous with pereion. Distal margin of pleotelson convex, distolateral corners not produced beyond medial margin. Rostrum acute, slightly upturned, with small tubercle on each side of base; frontal margin excavate on either side of rostrum. Lateral margin of pereionite IV squarely truncate. Pleopod | in male with rami fused, distal margin straight, with seven setae on each side, distolateral portion tapering to narrowly rounded apex. Uropods dorsally visible, extending almost to level of medial pleotelsonic margin. Material Holotype SAM-A15454 1 ¢ TL 2,8 mm SM 103 28°31,7’S 32°34,0’E 680 m 148 ANNALS OF THE SOUTH AFRICAN MUSEUM Allotype SAM-A15454 192 SM 103 8 gg 3 22 SM 103 SAM-A15455 3 33 3°22 SM 86 Remarks H. gernekei differs from H. tricornis Menzies, 1962, which it closely resembles, in the antennular flagellum (six articles in H. tricornis, five in H. gernekei, and the relative lengths of the articles differing); the first pleopod of the male, the distolateral angles being more produced than in H. tricornis; in the uropod extending at least to the level of the medial pleotelsonic margin or slightly beyond, and in lacking a lateral ridge on the dorsum of the pleotelson. The species is named for Mr D. Gerneke, previously of the South African Museum, for his invaluable assistance on both the Meiring Naude cruises. E Fig. 16. Haploniscus gernekei sp. nov. A. Holotype dorsal view. B. Maxilliped. C. Antennule. D. Apex of pleopod 1 ¢. E. Pleopod 2 3. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 149 Family Eurycopidae Eurycope glabra sp. nov. Figs 17-18 Description Body glabrous, head and first four pereionites narrower than rest of body. First four pereionites narrow, anterolateral angles acute; pereionite V broader than VI, lateral corners rounded; pereionite VII wider than VI, distal margin slightly concave. Pleotelson of single segment, broader than long, distally broadly rounded, uropods just visible dorsally. Rostrum distally rounded, with eight setae. Basal antennular segment broad, with inner distal angle produced, second segment narrow; flagellum of about fifteen articles. Mandibular palp with third segment strongly rounded, with several setae and fringed spines; molar process broad, truncate, with five marginal setae. Maxilla | with several simple curved spines distally, inner ramus with single strong distal seta. Maxilla 2 slender, with several elongate distal setae. Epipodite of maxilliped with strong rounded process on outer margin; three basal palp segments broad, second segment longest, with two spines on outer distal angle; two distal segments narrow, setose; endite narrower than basal palp segments. Pereiopod I very slender, remaining pereiopods missing. B Fig. 17. Eurycope glabra sp. nov. A. Holotype dorsal view. B. Pleopod 1 g. C. Maxilliped. D. Epipod of maxilliped. 150 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 18. Eurycope glabra sp. nov. A. Left mandible. B. Maxilla 1. C. Maxilla 2. D. Antennule. E. oe 2 3. F. Uropod. G. Rostrum. H. Pereiopod I. THE SOUTH AFRICAN MUSEUM’S MEIJRING NAUDE CRUISES 15] Pleopod | in male narrow, elongate, rami distally rounded, with sub- terminal process on outer margin. Pleopod 2 with outer margin strongly convex. Uropod biramous, inner ramus four and a half times longer than outer. Operculum of female with strong median crest, forming spine-like process proximally. Material Holotype SAM-A15452 1 ¢ TL 3,6 mm SM 103 28°31,7’S 32°34,0’E 680 m Allotype SAM-A15453 192 TL 3,3 mm SM 60 27°09,6'S 32°58,2’E 800-810 m Remarks From Wolff’s key (1962: 144) to the species of Eurycope the present species would seem to resemble £. antarctica Vanhoffen, 1914, but this species does not have acute antero-lateral corners of the first four pereionites, and the pleotelson is not distally as broadly rounded; further, the uropodal exopod is relatively longer, the rostrum not as narrow as in the present material. The specific name derives from the completely smooth dorsum, entirely lacking in sculpture. Family Dendrotionidae Acanthomunna spinipes (Vanhoffen) Fig. 19 Mormomunna spinipes Vanhoffen, 1914: 569, figs 100, 101. Acanthomunna spinipes: Menzies, 1962: 174. Wolff, 1962: 65. Previous records Antarctic (Gauss Station 8.1I.1903) 385 m. Material SAM-A15463 4 g¢ TL2,5mm 8 99 3 ovig. 92 TL4,4mm SM 86 SAM-A15464 2 gg 12 SM 103 Remarks The presence of a uropodal peduncle, pereionites I to III separate, the head not fused with pereionite I, maxilliped palp segments all slender, the presence of a mobile lacinia in the left mandible, an apically truncate molar process, coxae on pereionites II to VII, an antennular flagellum of at least six articles, pereiopods with a single claw, and dorsolaterally inserted uropods, all place the present specimens in the family Dendrotionidae. The presence of eyes indicate the genus Acanthomunna (Wolff, 1962: 65). The present material agrees almost exactly with Vanh6ffen’s description of A. spinipes, especially in the structure of the first and second pleopods of the male. While the generally spinose pereion and pleon, and the elongate antennae and pereiopods also 152 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 19. Acanthomunna spinipes (Vanhoffen). A. Dorsal view. B. Maxilliped. C. Pleopod | 3. D. Pleopod 2 3. E. Right mandible. F. Maxilla 1. G. Maxilla 2. H. Operculum 9. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 153 agree well, unfortunately the spectacularly spinose uropods of A. spinipes are absent in all the Natal material. In one specimen, the base of the uropodal peduncle was still in situ. The relatively large insertion for the uropods and the sturdy remnant mentioned indicate a massive uropod of the type figured by Vanhoffen. Family Ilyarachnidae Ilyarachna wolffi sp. nov. Fig 20 Description Body glabrous, tapering posteriorly. Head with transverse convex portion and broad lateral areas. Pereionite I not as broad as II, both laterally acute. Pereionites II to IV increasing in length, coxae of pereionites posterior to II rounded; posterior margin of pereionite V concave; VI triangular, VII narrower. Pleotelson of two segments, terminal segment longer than broad, distally narrowly rounded. Antennular basal segment broad, outer distal angle somewhat produced, second segment narrow; flagellum of about fifteen articles. Mandibular palp slender, 3-segmented, middle segment with two fringed spines, shorter terminal segment with four spines; incisor process a single rounded tooth; molar process narrow, with three terminal setae; strong ridge on inner face of mandible. Maxilliped with second palp segment very broad and long, third segment triangular, two distal segments short and narrow; endite less than half width of second palp segment, epipodite broad, distally rounded. Pleopod | in male curved, rami distally narrowly rounded, setose. Uropods missing. Material Holotype SAM-A15457 1 3 TL 2,9 mm SM 86 27°59,5’S 32°40,8’E 550 m 3 g3 SM 86 SAM-A15458 2 33 SM 103 Remarks Three species closely resemble present material. (See Wolff 1962: 94.) I. triangulata Menzies, 1962, is very similar in body proportions but lacks the acute coxae of the first two pereionites, the mandibular palp is also very different. I. affinis Barnard, 1920, has acute coxae on the first four pereionites, rather than just the first two. I. crassipes Barnard, 1920, has rounded coxae of the first two pereionites, while the pleon is about as long as wide. The species is named for Dr Torben Wolff, for his contributions to isopod biology and taxonomy. 154 ANNALS OF THE SOUTH AFRICAN MUSEUM ZZ = Sree m— Fig. 20. I/yarachna wolffi sp. nov. A. Holotype dorsal view. B. Pleopod 1 g. C. Maxilliped. D. Antennule. E. Maxilla 1. F. Maxilla 2: G. Mandible. H. Pleopod 2 dg. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 155 TIlyarachna sp. le, ZI Description Body markedly spinose. Head with eight spines. Pereionites I to IV with two rows of small spines and several lateral spines; pereionites V to VII with single row of spines. Pleotelson of two segments, terminal segment with several small granules plus single strong pair of spines. Material SAM-A15462 299 TL3,0 mm SM 103 Remarks As the uropoda and all the pereiopods are lacking from both specimens, specific status is not given, even though a species of //yarachna with a double row of small spines on the anterior four pereionites does not seem to have been recorded. This species is obviously related to such spinose species as I. spinosissima Hansen, I. argentinae Menzies, I. gurjanovae Menzies, and I, multispinosa Menzies GENERAL DISCUSSION Of the 21 species of isopod recorded, 15 were taken from either station SM 86 or SM 103, 7 species being common to both stations. The nature of the bottom at both stations was very similar, and as 9 of the 15 species caught are new, it may fairly be said that this is a habitat not previously sampled. A Fig. 21. Ilyarachna sp. °. 156 ANNALS OF THE SOUTH AFRICAN MUSEUM comparison of the two stations (Table 1) and the isopods caught at them will give some idea of the characteristics of this ecotype. The differences in the catch between the stations can to some extent be explained by the difference in fishing ability of the gear used. The heavy dredge, with its 100 cm by 30 cm mouth, would be able to pick up much larger organisms and rocks than could the light biological dredge (see Menzies 1962, fig. 1) with its narrow 100 cm by 10 cm mouth. Thus Alcyonaria and Gorgonacea with attendant isopods such as Neastacilla would be caught only by the heavy dredge, while some of the smaller debris-dwelling isopods of the upper few centimetres of substrate such as Eurycope, Serolis and Microarcturus would be caught by the light dredge, but perhaps lost through the wider mesh of the heavy dredge. TABLE | A comparison between two stations and the isopods caught at them. SM 86 SM 103 Depth 550 metres 680 metres Gear Heavy dredge Light biological dredge Bottom Fine hard mud, overlain with hetero- Fine hard mud, overlain with hetero- pod and pteropod shells, coral pod and pteropod shells, coral fragments, numerous foraminifera, fragments, numerous foraminifera, sponge fragments and spicules sponge fragments and spicules Non-Isopod Hydroids, gorgonaceans, solitary Hydroids, small echinoderms, pycno- organisms corals, glass sponges, pycnogonids, gonids, amphipods, small crabs, caught small crabs, cumaceans, amphipods, _— small gasteropods and bivalves. ostracodes, polychaetes, small gasteropods and bivalves Isopods Acanthomunna spinipes Acanthomunna spinipes caught Anthurids Anthurids — Bathycopea typhlops Excirolana bicornis — — Eurycope glabra Gnathia sp. = Haploniscus gernekei Haploniscus gernekei Ilyarachna wolffi Ilyarachna wolffi — Ilyarachna sp. Microarcturus youngi Microarcturus youngi = Microarcturus oudops Neastacilla longispina — — Serolis brinki Spinarcturus natalensis Spinarcturus natalensis Stenetrium abyssale Stenetrium abyssale Stenetrium dalmeida — Stenetrium dagama Stenetrium dagama The affinities of this fauna are not easily apparent, especially with 52 per cent being described as new. The presence of a Serolis, and Stenetrium abyssale and Acanthomunna spinipes indicate possible Antarctic/subantarctic affinities, while five endemic species (Microarcturus oudops, Stenetrium dagama, Cirolana imposita, and two easily recognizable anthurids Leptanthura laevigata and Mesanthura catenula) show the expected South African component. The unexpected presence of Bathycopea typhlops can only be explained by the paucity of collecting in this type of habitat, especially off the west African coast. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES Sy ACKNOWLEDGEMENTS I should like to thank the crew of the R/V Meiring Naude and my colleagues of the South African Museum who assisted in the collecting of the material described here. I am grateful to Dr T. E. Bowman of the Smithsonian Institution, and Prof. J. H. Day of the University of Cape Town, who read the manuscript and made many useful criticisms and suggestions. REFERENCES BARNARD, K. H. 1914. Contributions to the Crustacean fauna of South Africa. 3. Additions to the marine Isopoda, with notes on some previously incompletely known species. Ann. S. Afr. Mus. 10: 325-442. BARNARD, K. H. 1920. Contributions to the Crustacean fauna of South Africa. 6. Further additions to the list of marine Isopoda. Ann. S. Afr. Mus. 17: 319-438. BARNARD, K. H. 1940. Contributions to the Crustacean fauna of South Africa. 12. Further additions to the Tanaidacea, Isopoda and Amphipoda, together with keys for the identification of the hitherto recorded marine and freshwater species. Ann. S. Afr. Mus. 32: 381-543. Bowman, T. E. 1971. Excirolana kumari, a new tubicolous isopod from Malaysia. Crustaceana 20: 70-76. HALE, H. M. 1946. Isopoda—Valvifera. Rep. B.A.N.Z. antarct. Res. Exped. 4: 163-212. HESsLeR, R. R. 1970. A new species of Serolidae (Isopoda) from bathyal depths of the equatorial Atlantic Ocean. Crustaceana 18: 227-232. KENSLEY, B. 1975. Marine Isopoda from the continental shelf of South Africa. Ann. S. Afr. Mus. 67: 35-89. Louw, A. E. 1977. The South African Museum’s Meiring Naude cruises. Part 1. Station Data 1975, 1976. Ann. S. Afr. Mus. 72: 147-159. LoyYoLa E SILVA, J. 1971. Sobre os generos Ancinus Milne Edwards, 1840 e Bathycopea Tattersall, 1909, da colecao U.S. Nat. Mus. Archos Mus. nac., Rio de J. 54: 209-223. MENZIES, R. J. 1962. The isopods of abyssal depths in the Atlantic Ocean. Vema Res. Ser. 1: 84-206. NorDENSTAM, A. 1933. Marine Isopoda of the families Serolidae, Idotheidae, Pseudidotheidae, Arcturidae, Parasellidae, and Stenetriidae mainly from the South Atlantic. Further zool. Results Swed. Antarct. Exped. 3 (1): 1-284. SCHULTZ, G. A. 1973. Ancinus H. Milne Edwards in the New World (Isopoda, Flabellifera). Crustaceana 25: 267-275. TATTERSALL, W. M. 1905. The marine fauna of the coast of Ireland. Part V. Isopoda. Scient. Invest. Fish. brch Ire. 1904 (2): 1-90. VANHOFFEN, E. 1914. Die Isopoden der Deutschen siidpolar-Expedition 1901-1903. Dt. Siidpol.- Exped. 14: 447-598. Wo rr, T. 1962. The systematics and biology of bathyal and abyssal Isopoda Asellota. Galathea Rep. 6: 1-320. - nd ed 7 _ ee _ Sara ive a auc eee haan! 6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature (particularly Articles 22 and 51). Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov., etc. An author’s name when cited must follow the name of the taxon without intervening punctuation and not be abbreviated; if the year is added, a comma must separate author’s name and year. The author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is transferred from its original genus. The name of a subsequent user of a scientific name must be separated from the scientific name by a colon. Synonymy arrangement should be according to chronology of names, i.e. all published scientific names by which the species previously has been designated are listed in chronological order, with all references to that name following in chronological order, e.g.: Family Nuculanidae Nuculana (Lembulus) bicuspidata (Gould, 1845) Figs 14-15A Nucula (Leda) bicuspidata Gould, 1845: 37. Leda plicifera A. Adams, 1856: 50. Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b). Nucula largillierti Philippi, 1861: 87. Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. Note punctuation in the above example: comma separates author’s name and year semicolon separates more than one reference by the same author full stop separates references by different authors figures of plates are enclosed in parentheses to distinguish them from text-figures dash, not comma, separates consecutive numbers Synonymy arrangement according to chronology of bibliographic references, whereby the year is placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable. In describing new species, One specimen must be designated as the holotype; other speci- mens mentioned in the original description are to be designated paratypes; additional material not regarded as paratypes should be listed separately. The complete data (registration number, depository, description of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.: Holotype SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973. Note standard form of writing South African Museum registration numbers and date. 7. SPECIAL HOUSE RULES Capital initial letters (a) The Figures, Maps and Tables of the paper when referred to in the text e.g. *... the Figure depicting C. namacolus ...’; *. . . in C. namacolus (Fig. 10)...’ (b) The prefixes of prefixed surnames in all languages, when used in the text, it not preceded by initials or full names e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene (c) Scientific names, but not their vernacular derivatives e.g. Therocephalia, but therocephalian Punctuation should be loose, omitting all not strictly necessary Reference to the author should be expressed in the third person Roman numerals should be converted to arabic, except when forming part of the title of a book or article, such as ‘Revision of the Crustacea. Part VIII. The Amphipoda.’ Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial capital letter, provided the same generic name is used consecutively. Name of new genus or species is not to be included in the title: it should be included in the abstract, counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts. BRIAN KENSLEY THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES PART 7 MARINE ISOPODA VOLUME 74 PART 6 JANUARY 1978 | | ISSN 0303-2515 MUS. COMP. ZOOL. | LIBRARY APR 28 1978 HARVARD UNIVERSITY ug 22 ap NM CAPE TOWN INSTRUCTIONS TO AUTHORS 1. MATERIAL should be original and not published elsewhere, in whole or in part. 2. LAYOUT should be as follows: (a) Centred masthead to consist of Title: informative but concise, without abbreviations and not including the names of new genera or species Author’s(s’) name(s) Address(es) of author(s) (institution where work was carried out) Number of illustrations (figures, enumerated maps and tables, in this order) (b) Abstract of not more than 200 words, intelligible to the reader without reference to the text (c) Table of contents giving hierarchy of headings and subheadings (d) Introduction (e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents (f) Summary, if paper is lengthy (g) Acknowledgements (h) References (i) Abbreviations, where these are numerous 3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced with 2,5 cm margins all round. First lines of paragraphs should be indented. 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REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: ‘Smith (1969) describes...’ ‘Smith (1969: 36, fig. 16) describes...’ ‘As described (Smith 1969a, 19696; Jones 1971)’ ‘As described (Haughton & Broom 1927)...’ ‘As described (Haughton et al. 1927)...’ Note: no comma separating name and year Dagination indicated by colon, not p. names of joint authors connected by ampersand et al. in text for more than two joint authors, but names of all authors given in list of references. (b) Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, b, etc. to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a). For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list o, scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number (only if independently paged) in parentheses, pagination (first and last pages of article). Examples (note capitalization and punctuation) BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140. FISCHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. gén. 74: 627-634. Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320. : Konn, A. J. 1960b. Spawning. behaviour, egg masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51. THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. (continued inside back cover) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 74 Band January 1978 Januarie Part 6 ~~ Deel Bb owing NS THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS By N. A. H. MILLARD Cape Town Kaapstad The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular intervals as material becomes available Obtainable from the South African Museum, P.O. Box 61, Cape Town Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM word uitgegee in dele op ongereelde tye na beskikbaarheid van stof Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad OUT OF PRINT/UIT DRUK 1, 2(1-3, 5-8), 31-2, 4-5, 8, t.-p.i.), 51-3, 5, 7-9), 6(1, t.—p.i.), 71-4), 8, 911-2, 7), 10(1-3), 11(1-2, 5, 7, t.—p.i.), 15(4—5), 24(2), 27, 31(1-3), 32(5), 33 Price of this part/Prys van hierdie deel R3,90 Trustees of the South African Museum © Trustees van die Suid-Afrikaanse Museum 1978 ISBN 0 908407 33 5 Printed in South Africa by ; In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk., Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS By N. A. H. MILLARD South African Museum, Cape Town (With 9 figures, 8 tables and 2 appendices) [MS. accepted 9 August 1977] ABSTRACT The geographical distribution of 251 species of hydroid Cnidaria round the coast of southern Africa is analysed by means of a system of radial sectors. It is concluded that there is no distinct west coast fauna as such, but that a boundary zone exists at approximately 31°S latitude between an east coast region with mainly tropical affinities and a temperate west—plus— south coast region with a reduced tropical component and a large endemic population. The main centre of the endemic population is the Agulhas Bank. The relationships of the tropical species are mainly with the Western Indian Ocean Province and the Indo-West-Pacific Region. Deep-water species (45) which occur below 400 m are of a mixed nature and show no clear relationship with other parts of the world. Deep-water species are classified into stenobathic/ eurybathic and stenothermic/eurythermic categories, and their distribution relative to water temperatures is discussed. CONTENTS PAGE Introduction : ‘ : : ; i 5 ; sn LESS) Methods . : ; ‘ : : t j E nel LOO Faunal boundaries. é i ; : : . 164 The tropical component . ; ; 5 : ; 169 The temperate component : ; : : : wat tel © The endemic component . ; F ; ‘ G : 173 The cosmopolitan and scattered components ; ‘ 174 Distribution and depth . ; : ; K ; ae tS Estuarine and brack-water species . : 2 : mel 9 Conclusion . : : 5 ; ‘ ; : ‘ : 182 References . : : ; ; k : ; oh LSS) Appendix 1 F : : : : 5 4 : 4 185 Appendix 2 : ; : q : , i 5. IBS INTRODUCTION This paper is an attempt to analyse the distribution of the hydroid fauna of the coast of southern Africa as documented in a recent monograph (Millard 1975). Since the latter went to press a few more taxonomic papers on, or lists of, hydroids have appeared, namely Arnaud ef al. (1976), Beurois (1975), Calder (1975, 1976), Cooke (1975), Cornelius (1975a, 1975b), Mergner & Wedler (1977), Millard & Bouillon (1975), Millard (1977a) and Watson (1975). Among 159 Ann. S. Afr. Mus. 74 (6), 1978: 159-200, 9 figs, 8 tables, 2 appendices. 160 ANNALS OF THE SOUTH AFRICAN MUSEUM these are records which extend the range of some South African species and they have been taken into account in the analysis which follows, as also have records from Buchanan (1957), Michel (1974), Blanco (1967, 1968, 1973, 1974) and Blanco & Bellusci de Miralles (1972a, 19726), which have only recently become available to the author. Within the borders of southern Africa, here taken to be the 20° parallel of south latitude, several new distribution records from deep water off the Natal coast and some new records for the country were published by Millard (19775). There are also scattered identifications by the author, as yet unpublished, which are listed in Appendix 1. The following changes in name have been adopted here: Antennella africana Broch, 1914, to A. quadriaurita Ritchie, 1909, fide Millard CIS Vilo2 123). Eulaomedea calceolifera (Hincks, 1871) to Campanularia calceolifera Hincks, 1871, fide Cornelius (1975a: 254).* Lafoea fruticosa (M. Sars, 1851) to Lafoea dumosa (Fleming, 1820) fide Cornelius (I97Sbe 385). Obelia bicuspidata Clarke, 1875, to O. bidentata Clarke, 1875, fide Cornelius (1975a: 260). Any zoogeographical analysis is aimed at pinpointing population changes within an area, and, if possible, relating them to physical or chemical factors and determining relationships with other areas. Such an analysis is subject to certain unavoidable weaknesses—the assignment of species to categories or components is largely subjective; numbers of species within categories are liable to change as more records are forthcoming (for instance some of the South African endemics may in time be expected to turn up outside our boundaries); and numbers of species may be influenced by different intensities’ of collecting in different areas or by misidentifications. The longer we wait the more reliable will be the results, but results, if only interim ones, are needed now. The larger the number of species involved the more meaningful will be the conclusions. One must bear in mind that it is the main trends which are significant rather than the details. METHODS The first step in the analysis was to remove all doubtful records and incomplete identifications. This left a total of 251 species. The first objective was an analysis of distribution within the borders of southern Africa, and for this some system of comparison was necessary. When dealing with the littoral zone it is a comparatively simple matter to convert the distance along the coast to a straight line, a method with obvious practical * Cornelius pointed out that the type species of Campanularia is Sertularia verticillata Linnaeus, 1758, and not Sertularia volubilis Linnaeus, 1758, as previously thought (Millard 1975: 203). Since S. verticillata is a branched species the diagnosis of Campanularia given previously (Millard 1975: 203) must be altered to include branching forms, and Eulaomedea sunk in Campanularia. 16] THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS “(SL61) PALT[UAL WOLZ “somtpeso] SUIDOT[OD UTP SY} SUIMOYS BOLIfY UJOYyINOS JO dey “{ “SI ce vet l€ O€ 6% 82 él StL vl el Zl 2 X A 7 AY ow yueg seujn ? 7 Gee .. 1 uopuo7 yse3 UeACH YL Z Aeg 294309 Ay SUYOT 3S }410d If elovecaicnnen JONIAONd 3dVvd yB1ng1005 oburdis).-° TWIVN * : E keg spreyoy os > ie eee ‘y SOUBT) Payey e19N7] 4S : wd ae axe} e6ue|UN s} eoeyu ese auequeyu| cel ee Nee ewes 8e Ze of 9€ SE St ve ve ce As le 10} 67 8Z Le dijyGuey aded yulog aded 97 GZ Aeg asjey UMO}SUOWIS A\jaao}D B4saquazinjw £4 oupnpue)) jeesyapno Keg sdwes uanoyeg VIASNINAd ddVO JHL VOINsV - LSAM HLNOS Ve 162 ANNALS OF THE SOUTH AFRICAN MUSEUM advantages, and this has been done by Stephenson (1947), Stephenson & Stephenson (1972), Day (1967) and others. However, when dealing with the sub- littoral area as well, this is clearly not possible for a continent of the shape of southern Africa. After much consideration the following arbitrary method was devised, in which the coastal waters were divided into thirty-six sectors of approximately equal area (Fig. 2). The central point for the sectors was chosen as the intersection of the latitude line of 20°S and the longitude line of 24°E, a point which is approxi- mately mid-way between the west and east coastlines. From this central point radii were drawn at 5° intervals, starting with one through the Cape Peninsula, and then working outwards in both directions. The sectors thus constructed were numbered 1-36. The reason for placing one radius (Number 14/15) through the Cape Peninsula is that Cape Point is generally recognized as an approximate zoogeographical boundary between the cold Benguela water on the west and the warmer Agulhas water on the east, and might be expected to have some significance. It so happens that another radius (Number 15/16) conveniently passes through Cape Agulhas, the most southerly point of Africa, and a third (Number 23/24) through the boundary between Transkei and Natal. ae DG / ES Ox 15° 20° 25° 30° 35° 40 20) 35 40 Fig. 2. Map of southern Africa illustrating the sector system which was used to analyse the distribution. THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 163 It must be emphasized that this is an arbitrary method, and obviously could not be applied to all coastlines, but it is felt that it might form a useful working method and basis for comparison with other southern African marine groups. It has the advantage that each sector covers an approximately equal area of sea and ‘straight-line’ length of coast and represents a cross-section through all depths. Since the sectors are roughly normal to the coast and the currents tend to follow the coast, each sector is affected in a similar way and population differences related to water temperature should be highlighted. It is felt that the advantages of this method at least outweigh the disadvantages (and impracticabilities) of the ‘straightening-out’ method of Stephenson. The second objective was an analysis of the world-wide distribution of the southern African species, and for this the species were delegated to components defined as follows: TROPICAL Circumtropical. Present in the tropics of all three oceans, but may spread north or south into warm temperate seas as well. Indo-West-Pacific Region. Stretching from the Suez Canal across the Indian Ocean and the East Indies into the Western Pacific as far as Hawaii and Easter Island. Includes northern Australia, Madagascar and East Africa. May spread into warm temperate seas as well. Western Indian Ocean Province. The western section of the Indo-West- Pacific Region, bounded in the north by the Persian Gulf, and including Madagascar and surrounding islands. TEMPERATE. (In a trial analysis it was found impossible to differentiate between cold temperate and warm temperate fauna when assessing world distribution.) Antitropical (bipolar). Present in the temperate zones of all three oceans, and present in both the Northern and the Southern hemisphere, but not in the tropics except perhaps just at the edge. Southern. Restticted to the temperate waters of the Southern hemisphere, but often in the subantarctic islands as well, and rarely reaching the Antarctic. Atlantic. Restricted to the northern and/or southern temperate zones of the Atlantic, but absent from the tropics. May spread further north into the subarctic. ENDEMIC OTHER Cosmopolitan. Present in all three oceans, and occurring in both tropical and temperate seas. Scattered. Present in various regions, but not sufficiently widespread to be included in Cosmopolitan. Does not fit any other category. 164 ANNALS OF THE SOUTH AFRICAN MUSEUM A list of species, together with their components and distribution within thirty-four sectors, is given in Appendix 2. (Sectors 35 and 36 have been omitted as no records were available from this region.) FAUNAL BOUNDARIES It has long been recognized that the marine fauna of southern Africa includes both tropical elements, predominating on the east coast, and temperate elements, predominating on the south and west coasts. Most authorities, following Stephenson (1947) and Stephenson & Stephenson (1972), have recognized a separate east coast (tropical or subtropical) fauna, a south coast fauna (warm temperate) and a west coast fauna (warm or cold temperate) for the littoral region. These earlier views are summarized by Briggs (1974), who includes the east coast in the Western Indian Ocean Province of the Indo-West- Pacific tropical region; the south coast in the Agulhas Province of the South African warm temperate region; and the west coast as a separate South West Africa Province of the South African warm temperate region. Opinions have differed as to the position of the boundary between the east and south coast, and whether the west coast represents a separate faunal province, and if so whether it should be regarded as cold temperate or warm temperate. The distribution of the faunal components of the continental shelf and the extent of the faunal regions is largely dependent on the ocean currents round the South African coast, in general the warm southward-flowing Agulhas Current being responsible for the southerly extension of tropical forms on the east coast, and the cold northward-flowing Benguela Current being responsible for the more northerly distribution of cool-water forms on the west coast. It is not intended to discuss these current systems in any detail here since accounts - are available in most textbooks of marine biology (a concise general account is given by Stephenson & Stephenson (1972) and more recent contributions are summarized by M. J. Penrith (1976)). In order to determine whether there is any region of the coast where a marked change in the hydroid fauna occurs, the range of the tropical, temperate and endemic components was plotted separately (Fig. 3). (The range of species was used in preference to presence/absence, in order to minimize differences in collecting intensity.) This figure emphasizes the tropical nature of the east coast fauna, for there is a strong tropical component at the northern end which gradually drops away, until beyond St Helena Bay on the west coast (Sector 14) only one species remains for a short distance. The rate of disappearance is greatest at the boundary of Sectors 23/24, and is also high in the region of Sectors 13/14/15. The temperate and endemic components are both highest on the south coast, falling away fairly evenly to the north-east, but more sharply to the north-west in the region of Sectors 13/14/15. A similar trend appears when the range of species is plotted as a percentage of the total range (Fig. 4). It is thus possible to distinguish two ‘zones of change’, though they are not as marked as might be expected. The first (Sectors 23/24) provides a convenient NUMBER OF SPECIES 20 10 50 40 30 4 20 10 80 70 60 50 40 30 20 10 THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS a 2) = Cc > o aos & iS) © jaa) c— Ni] GS a = “oD NOES 2 = eos EB oc tn ag > ra) ® ® a 2 S 3 S55 Beno 5 = im jOO re a { { tts t 4 { TEMPERATE ENDEMIC TROPICAL 5 "10 15 20 “95 SECTORS «I|nhaca 30 165 Fig. 3. The distribution of the temperate component (above), the endemic component (centre) and the tropical component (below). From range of species. 166 ANNALS OF THE SOUTH AFRICAN MUSEUM boundary between the east and south coast fauna. It is located between 30 and 32°S latitude, i.e. the stretch of coast between the Bashee River in the south and Durban in the north. For the discussions which follow, the boundary between Sectors 23 and 24 is used (31°S). This is more or less in agreement with con- clusions drawn from other faunal groups. Stephenson & Stephenson (1972) suggest a boundary zone between Port St Johns and Qolora, 1.e. at about 32°S, for the intertidal fauna (the form of the graph of the tropical component in Figure 3 of this paper is very similar to that for the intertidal fauna in Stephenson & Stephenson 11972, ie. 8:26); Je 1 Ba Smith G949) and M.-L. Penrith (1970) suggest the Great Kei River at 32°42’S for the fishes, and Day (1967) suggests the Bashee River at 32°15’S for the benthic Polychaeta. TEMPERATE 20 owe 40 0-0 20 ° ENDEMIC : RANGE 80 60 TROPICAL PERCENTAGE 40 20 SECTORS Fig. 4. The distribution of the temperate, endemic and tropical components, from range of species as a percentage of the total range. Sectors 1-7 have been omitted since the total number of species is less than twenty and the percentages unrealistic. THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 167 At the boundary of Sectors 23/24 the monthly minimum surface temperature near the coast does not fall below 16°C, the maximum does not rise above 27°C and the mean ranges from 21 to 24°C (U.S. Naval Oceanographic Office 1967). This is approximately the region where the Agulhas Bank begins to widen and force the warm Agulhas Current offshore. Further south the latter is separated from the coast by a cooler counter-current flowing northwards. The usefulness of the second ‘zone of change’ in the south-westerly corner of the Cape Province (Sectors 13/14/15) is more debatable. It could be used to divide the temperate region, which stretches from north of 20°S on the west coast to 30-32°S on the east coast, into two provinces, if warranted by the fauna. To test this an arbitrary boundary was placed at Cape Point (Sector 14/15) and the composition of species on the two sides of it compared with one another and with the east coast (Table 1). (Cape Point was used as a boundary by Stephenson (1947, fig. 3), Ekman (1953) and Briggs (1974).) It may be noted from Figures 3 and 4 that there is no evidence of a change in fauna at Cape Agulhas (Sector 15/16), the most southerly point of Africa). TABLE | Analysis of fauna of west coast, as compared with that of the south and east coasts (actual records, not range). East coast (Sectors 24-36) West coast South coast (Sectors 1-14) | (Sectors 15-23) Component No. of he No. of VA No. of Sfp species species species Tropical 8 9,6 30 19,0 eZ 39,6 Temperate . 13 IS7/ 20 Ora 12 6,6 Endemic to area : : 2 2,4 D)\ 13,3 7 933 Endemic to more than one area 25 30,1 4| D9 30 16,5 21 1353 28 15,4 Tes) 15,8 23 12,6 Cosmopolitan Scattered Total It is apparent from Table 1 that the west coast fauna consists mainly of endemic, cosmopolitan and scattered species. The tropical species diminish in number round the coast from east to west; only 8 species remain on the west coast and these are all absent north of St Helena Bay. The temperate species of the west coast are fewer in number than those of the south coast. Only 2 of them (Clavopsella nayvis and Gonothyraea loveni, both from Table Bay harbour and probably introduced by ships), and possibly a third (Sarsia eximia), are restricted to the west coast. Only 2 rare species are endemic solely to the west coast (Myriothela tentaculata and Eudendrium ritchiei), whereas 21 are endemic to the south coast and 17 to the east coast. Of the cosmopolitan and scattered components only 3 (Acryptolaria crassicaulis, Hybocodon unicus and Hydro- dendron gracilis) have been recorded from the west coast alone. 168 ANNALS OF THE SOUTH AFRICAN MUSEUM In total, thus, the west coast has only 8 unique species (9,8 %), of which only 2 (2,4°%) are endemic to the area. Briggs (1974) requires at least 10 per cent endemic species for the designation of a separate province, and on this basis the west coast of South Africa does not qualify, at least for the hydroids. The hydroid fauna of the west coast seems to be mainly an impoverished south coast fauna. Although Day (1967) accepted a separate west coast element for the littoral polychaets, he could find no marked difference between the south and west coasts for the sublittoral ones, and ascribed this to the fairly uniform water temperature on the bottom of the continental shelf (he gave figures of 12—14°C at 100 m from Port Elizabeth to Liideritz, but the records collated by Hulley (1972) show a somewhat lower range of 8-13°C at 100 m between 20°S on the west coast and Port Elizabeth on the south coast). On the other hand there is a difference in the surface water temperature on the two sides of Cape Point (Day gives a range of 15-20°C for the south coast and 12—15°C for the west coast, but under certain conditions and for certain periods the south coast temperature may drop to as low as 11°C (during local upwelling) and rise as high as 24°C, while that of the west coast may drop to 9°C (M. J. Penrith 1976)). In this respect 1t might be noted that all hydroids are sublittoral in the sense that they cannot withstand drying and survive in the littoral zone only in damp overhangs or open runnels. None of this really explains why there should be a diminution of hydroid species on the west coast and practically no endemic population of cold- temperate forms. Ekman (1953) associates the poverty of the benthos of the ‘Namaqua coast’ with the hydrogen sulphide resulting from frequent upwelling and red tides. Briggs (1974) suggests that certain species are prevented from” spreading westwards round Cape Point by the absence of high summer temperatures for reproduction (he gives a February mean of 16—-18°C for South West Africa as against one of 20—22°C for the south coast, but compare remarks in parenthesis above). M.-L. Penrith & Kensley (1970a) suggest that a region of minimal temperatures between Hondeklip Bay and Liideritz (i.e. north of St Helena Bay) acts as a barrier to the establishment of southern warm- temperate intertidal species further north. The paucity of hydroid records on the west coast may be partly a reflection of the difficulty of collecting on the exposed and inhospitable Skeleton Coast. However, M.-L. Penrith (pers. comm.), who has collected in this area, feels that the poverty is a real one, and perhaps linked to the absence of suitable habitats such as sheltered coves, rock pools, etc., and the preponderance of surf beaches and sandy floors. The harsh conditions and poverty of the rocky-shore fauna are also emphasized by M. J. Penrith (1976). There is no evidence at all of a southward spread of species from tropical west Africa. The southern boundary of the west African tropical region is too far north for consideration in this paper (14 or 15°S: Briggs 1974; 18-22°S: Hulley 1972). M.-L. Penrith & Kensley (19706) found the first of the west African littoral species at Rocky Point (18°59’S). THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 169 In conclusion the south and west coast sublittoral areas are considered to represent a single temperate province for the hydroid fauna, which might be termed the Agulhas/Namaqua Province, the centre of the endemic population being on the Agulhas Bank. THE TROPICAL COMPONENT Of the total number of 251 species, 77 (30,7 °%%) can be classified as tropical. Of these, 20 are circumtropical, 24 Indo-West-Pacific and 33 confined to the Western Indian Ocean. In addition there are 17 endemic species which are confined to the South African east coast, and therefore with tropical tendencies (these were termed ‘subtropical endemics’ by Day (1974)). (Appendix 2.) Ekman (1953) and Briggs (1974) consider that the main centre of origin of the tropical fauna is the Indo-Malayan Region. From here species tend to spread west to the Indian Ocean (and may penetrate into the Red Sea or round southern Africa), and to a lesser extent east to the Eastern Pacific. Species have been classified here as Indo-West-Pacific or Western Indian Ocean even though a few have one or two records outside these areas. Thus, 2 have penetrated into the Mediterranean (Corydendrium parasiticum, Thyro- scyphus fruticosus), 4 have reached the tropical west coast of Africa (Abietinaria laevimarginata, Campanularia africana, Corydendrium parasiticum, Sertularia ligulata), | has reached tropical South America (Dentitheca bidentata), and | has reached southern Australia (Dynamena obliqua). Further spread would produce a circumtropical distribution, but of the species classified as such no less than 13 have failed to reach the east coast of the Pacific, and this applies also to 4 of the cosmopolitans (Table 2). There is therefore support for Briggs’s contention (1974) that the ‘East Pacific barrier’ is one not easily crossed. There is some evidence for a secondary centre of origin in the Western Indian Ocean, for in addition to the seventeen South African subtropical endemics (i.e. present on the east coast only), there are seven Western Indian TABLE 2 List of circumtropical and cosmopolitan species which do not occur on the east coast of the Pacific. Circumtropical Cosmopolitan Diphasia digitalis Antennella secundaria Dynamena quadridentata Cladocoryne floccosa Gymnangium hians Coryne pusilla Halecium dyssymetrum Halecium sessile Hydrodendron caciniformis Idiellana pristis Monostaechas quadridens Plumularia strictocarpa Scandia mutabilis Sertularella diaphana Sertularia distans S. marginata S. turbinata 170 ANNALS OF THE SOUTH AFRICAN MUSEUM Ocean species which, though absent from the Indo-Polynesian area, have yet penetrated into the Red Sea (C/ytia latitheca, Cytaeis nassa, Diphasia heurteli, Halopteris glutinosa, Hydractini kaffraria, Plumularia wasini, Solanderia minima). Some communication between southern Australia and the Western Indian Ocean is indicated, since there are five species common to the two, namely Amphisbetia maplestonei, Crateritheca acanthostoma, Halopteris glutinosa, Sertularella arbuscula and Synthecium dentigerum. These occur nowhere else (except for H. glutinosa which also occurs in the Red Sea) and the likelihood is that the movement has been from west to east with the West Wind Drift. THE TEMPERATE COMPONENT The total temperate component includes 28 species (11,2%), of which 11 are antitropical (bipolar), 10 are restricted to the Southern hemisphere and 7 are restricted to the Atlantic (Appendix 2). These are species which cannot survive the high temperatures of the tropics, although some can spread into the cooler waters of the subpolar regions. The southern species must have evolved in the south, but the antitropical and Atlantic species may have evolved in either the south or the north and must at some time have crossed the tropics. This may have been achieved by equatorial submergence, for Briggs (1974) points out that the tropical surface water is only about 30-40 m deep in the Eastern Atlantic. It may also be due to transportation on the hulls of ships. In all, twenty-one species have been recorded on ships’ hulls or floating objects such as buoys. The fauna of ships’ hulls is a characteristic group of hardy species (Millard 1952), most of them cosmopolitan. Three of the species (all. belonging to the temperate component) are known only in harbour areas in South Africa, and it is suggested that these are recent introductions by ships: Gonothyraea loveni is well established in Cape Town docks, occurring abundantly on pylons, cables, etc., as well as on hulls of ships. It has not been reported from any other part of South Africa. Elsewhere it is known from the North Atlantic, New Zealand, Tasmania and the Argentine. Campanularia calceolifera has been found, and reproducing, in Cape Town docks on a barge and on a raft, neither of which had left the harbour. It must, therefore, be established, though not common. It is not known for certain elsewhere in South Africa, a record from False Bay by Stechow (1925) being of infertile material and thus subject to doubt. The species is well known on European and Mediterranean coasts and also occurs on the east coast of North America. Clavopsella navis is known from a ship’s hull in Cape Town docks, and also from the Kiel Canal (Thiel 1962 as C. quadranularia). The latter is probably the original locality since the colonies there were more abundant and better developed. THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 17] Since all these three species occur both in the South and North Atlantic they must have traversed the tropics, though they have not established them- selves in tropical latitudes so far as is known. They may have been hardy enough to survive a brief passage through warmer waters on ships’ hulls, or they may have died back during transport, leaving the hydrorhiza to regenerate again in suitable temperature conditions. Ralph (1961) also found evidence of the introduction of hydroids by ships’ hulls to New Zealand harbours. It is interesting to compare the relations of the temperate species with the other two southern continents. From Table 3 it can be seen that South Africa has 9 species in common with southern Australasia alone, but only 3 species in common with South America alone. There are 7 species common to all three. TABLE 3 Temperate species common to the southern continents. South Africa and South South Africa and temperate Temperate part of all Australasia South America three southern continents Kirchenpaueria triangulata Phialella turrita Amphisbetia minima Nemertesia ciliata Sertularella striata _ Filellum antarcticum Parascyphus simplex Symplectoscyphus paulensis Gonothyraea loveni Plumularia obliqua Nemertesia cymodocea P. spinulosa Plumutaria filicaulis Sarsia eximia Staurocladia vallentini Sertularella annulaventricosa Tubularia larynx Stereotheca elongata Symplectoscyphus macrogonus (Total: 9) (Total: 3) (Total: 7) This would suggest dispersal eastward from the Agulhas Bank. Beurois (1975) emphasized the role of eastward drifting kelp in the dispersal of attached forms to St Paul and Amsterdam Islands, and Arnaud et al. (1976) discussed the transport of South African fauna to St Helena on kelp. If the list in Table 3 is extended to include the cosmopolitan and scattered components and those tropical forms which have spread south of the Tropic of Capricorn, the relation- ship between South Africa and Australasia is more marked (34 species in common with Australasia alone, 10 with South America alone, and 26 in all three). The distribution of those South African species which have antarctic or subantarctic affinities is given in Table 4. In all, 41 species are listed, a relatively small number (16,3 % of the total), and of these half (21) are cosmopolitan and this tends to mask the relationships. Without the cosmopolitan species only 20 (8,0°%) have antarctic affinities and of these 11 are classified as temperate. The absence from South Africa of certain common antarctic genera, such as Staurotheca, Tulpa, Grammaria and Silicularia, is very striking. On the whole there is very little evidence of relationship with the Antarctic. 172 ANNALS OF THE SOUTH AFRICAN MUSEUM TABLE 4 Distribution of species with antarctic/subantarctic affinities. The Magellan Region includes Gauss Station, the Falklands and South Georgia. The Kerguelen Group includes Marion, Prince Edward and Crozet Islands. Be 2 ae 3 2 | Seen Species Component 2 e=| c = § gee se 8 =e) ps) ae 5 An a 24 ee ees Acryptolaria conferta Cosmopolitan x x A. crassicaulis Scattered x x Aglaophenia p. pluma Cosmopolitan < x Amphisbetia minima Temperate (Antitr.) < x A, operculata Cosmopolitan x x x Antennella quadriaurita Scattered x A. secundaria Cosmopolitan x Bimeria vestita Cosmopolitan x Bougainvillia macloviana Scattered x < Campanularia crenata Cosmopolitan x C. integra Cosmopolitan x Clytia hemisphaerica Cosmopolitan x x C. paulensis Scattered x x Coryne pusilla Cosmopolitan x Dynamena cornicina Cosmopolitan x< Filellum antarcticum Temperate (Southern) x XK F. serratum Cosmopolitan x x Halecium beanii Cosmopolitan x H. delicatulum Cosmopolitan x x x H. tenellum Cosmopolitan x x x x Hincksella echinocarpa Temperate (Southern) x Hybocodon unicus Scattered x Kirchenpaueria triangulata Temperate (Southern) x Lafoea benthophila Scattered x x x x L. dumosa Cosmopolitan x x x Modeeria rotunda Cosmopolitan x x Nemertesia cymodocea Temperate (Southern) x N. ramosa Scattered x Obelia bidentata Cosmopolitan x x x O. dichotoma Cosmopolitan x x < x O. geniculata Cosmopolitan x % x x Parascyphus simplex Temperate (Antitr.) s< x Phialella turrita Temperate (Antitr.) x Plumutlaria pulchella Seattered x P. setacea Cosmopolitan x x Sertularella leiocarpa Temperate (Southern) ~ S< S. p. polyzonias Cosmopolitan x s< S. striata Temperate (Southern) x Staurocladia vallentini Temperate (Southern) X< < Symplectoscyphus paulensis Temperate (Southern) S x Zygophylax armata Scattered Ne THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS WB, THE ENDEMIC COMPONENT The number of endemic species strictly belonging to the South African coast south of 20°S is 76. However, there are 2 species which also occur immediately north of 20° on the west coast (Gattya humilis and Salacia articulata). 4 species occur only in South Africa and the Vema Seamount (Halopteris pseudoconstricta, Salacia articulata, Sertularella flabellum and S. megista), and it was shown by Millard (1966) that the hydroid fauna of the Vema Seamount is essentially South African (21 of the 23 species also occur in South Africa). If these 6 species are included, the endemics number 82 (32,7 °% of the total of 251). It might be noted, however, that Mergner & Wedler (1977) have recorded Eudendrium deciduum and Sertularella natalensis from the Red Sea with a query. As a comparison the following percentages of endemics have been given for other large marine groups in southern Africa. The figures are not strictly comparable, however, since the boundaries used for the “Southern African Region’ are not always the same. Polychaeta . 107, (Deyn oy) Echinoderms, other than Holothuria 48% (Clark & Courtman-Stock 1976) ASCidians |=) eres | 60 7 ra Mullan nlO7) Bishi sy ee ee 25s A (ME Ivin Snntinel 970; 380Vontvot 1 500 species) The south coast of southern Africa has more endemic species of hydroid than any other region, and the Agulhas Bank seems to be the centre of distri- bution of the endemic population (Figs 3-5). There are only three genera endemic to South Africa, Bicorona, Hydro- corella and Uniscyphus, and all of these are monotypic. However, certain genera have a larger proportion of endemic species than others (Table 5). These are probably genera in which speciation is actively occurring in the South African area. They are also genera in which many of the species are not easily dis- tinguished from one another, and thus pose constant problems to the systematist. Similarly, species with several subspecies are ones likely to be undergoing active speciation. Thus, Thecocarpus flexuosus has four subspecies; the nominal one is TABLE 5 List of South African hydroid genera with the largest number of endemic species and/or subspecies. Number of endemic species Total number of species Genus or subspecies or subspecies in South Africa Cladocarpus 9 13 Corhiza 5 5) Halopteris 4 6 Hydractinia 3 3 Sertularella 14 23 Zygophylax 4 7 NUMBER OF SPECIES 174 ANNALS OF THE SOUTH AFRICAN MUSEUM WEST SOUTH EAST West only West + south AO South only 30 20 10 South+ east O East only West + south+ east 30 SECTORS Fig. 5. An analysis of the distribution of the endemic species, with the total number of species distributed between six components, as in Stephenson (1972). From range of species. tropical Indo-West-Pacific and the other three are endemic to southern Africa. Sertularella polyzonias also has four subspecies; the nominal one is cosmopolitan and two of the others are endemic to southern Africa. Several common genera, on the other hand, have no southern African endemics: Aglaophenia, Amphisbetia, Dynamena, Lytocarpus, Nemertesia, Obelia and Sertularia. THE COSMOPOLITAN AND SCATTERED COMPONENTS The cosmopolitan species number 28 (11,2°%%). They owe their wide distribution to the fact that they are eurythermic, hardy and adaptable. The wide distribution possibly indicates a very old history, in which there has been plenty of time to disperse. | Species classified as scattered or disjunct number 36 (14,3 %). Most of them NUMBER OF SPECIES THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 175 have odd distributions which do not fit any of the other categories. There are two possibilities. Either these are newly evolved eurythermic species which are in the process of spreading rapidly and likely to become cosmopolitan, or they are ‘peripheral relicts’ of fairly primitive forms which were once more widely distributed. Among the scattered species three are interesting because outside southern Africa they are limited to the west and east coasts of the tropical Atlantic (Clytia hummelincki, Zyzzyzus solitarius) or to the western tropical Atlantic only (Cladocarpus tenuis). Three others have a similar distribution but also extend further north and/or east (Aglaophenia latecarinata, A. pluma dichotoma, Symplectoscyphus amphoriferus). Cosmopolitan and scattered species with antarctic/subantarctic affinities are included in Table 4. DISTRIBUTION AND DEPTH Figure 6 shows that there are more species present in the shallow water (0-100 m) than at any other level, and although there have been fewer samples from deep water it is not expected that more extensive collecting will alter the picture to any great extent. The shallow water is obviously the optimum zone for hydroids. Most of the species in the littoral area are at the upper edge of 180 160 140 120 100 80 60 40 20 © 100.2250 500 750 1000 1250 1500 DEPTH (m) Fig. 6. The distribution of species according to depth. From range of species. 176 ANNALS OF THE SOUTH AFRICAN MUSEUM their depth range and able to survive only under overhangs and in rock pools and crevices. Only 24 of the 98 littoral species have not been found below low water of springs. Deep-water species are those which occur over the edge of the continental shelf, and Ekman (1953) places an arbitrary upper boundary for the deep-sea fauna at 400 m. The number of hydroids known from below this depth round the South African coast is relatively small (45 = 17,9%), and only eight of these reach | 000 m (abyssal). The species are listed in Table 6. These species can be divided into two arbitrary categories on the basis of their depth range both in South Africa and the rest of the world: Stenobathic: those that are restricted to water deeper than 200 m. Eurybathic: those that occur above and below 200 m. These two categories are distributed between the different geographical components as follows: Stenobathic Eurybathic Total bropicall, smc psihae pes we ee Cae 5 5 10 Temperate . l 4 5 Endemic 5 6 1] Cosmopolitan 0 2) 9 Scattered I 9 10 Mkovtalt nee Feu? Poe eh teenie i 12 33 45 All the (12) stenobathic species are in reality cold-water species frequenting water below about 16°C (Ekman 1953, table 42), and this includes five so-called ‘tropical’ species. These species are thus also stenothermic. Their limiting factor for distribution may be depth or temperature or a combination of both. Some of the eurybathic species are also stenothermic. This applies to the following tropical and scattered species whose records within the tropics are never shallow, and which are thus restricted to cold water: Most shallow record in tropics (m) Acryptolaria rectangularis (tropical) 228 Cladocarpus tenuis (scattered) 185 Cryptolarella abyssicola (scattered) 4 560 Symplectoscyphus amphoriferus (scattered) 161 Zygophylax sibogae (scattered) Only one tropical record: depth not given. To the same category of eurybathic/stenothermic species might be added the endemic and temperate eurybathic species, at any rate those restricted to the south-west coast of South Africa and the east coast below 200 m: Cladocarpus sinuosus (endemic): minimum east coast record 400 m. Hincksella echinocarpa (temperate): minimum record Kerguelen 41 m, minimum east coast record | 610 m. THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 177 TABLE 6 South African deep-water species. SB—stenobathic (never recorded above 200 m); EB-—eurybathic (occurring at all depths); ST—stenothermic (cold-water species); ET—eurythermic (occurring at all temperatures). Depth range Depthrange Component Species Component in South outside South _ of depth| Africa (m) Africa (m) temperature Acryptolaria conferta Cosmopolitan 64-1200 0-4400 EB/ET A, crassicaulis Scattered 2835 109-767 EB/ET A. rectangularis . Tropical 110-1200 228-600 EB/ST Antennella quadriaurita Scattered 0-825 0-400 EB/ET Branchiocerianthus imperator Tropical 730 200-5290 SB/ST Campanularia hincksii Cosmopolitan 9-450 0-811 EB/ET Cladocarpus distomus . Tropical 292-2200 55-1362 EB/ET C. dofleini Tropical 425-550 1019 SB/ST C. inflatus Endemic 495 — SB/ST C. millardae Tropical 200-595 5020 SB/ST C. natalensis Endemic 400-900 — SB/ST C. sinuosus . Endemic 183-680 — EB/ST C. tenuis Scattered 495 185-550 EB/ST Clytia gravieri Tropical 0-528 04243 EB/ET Cryptolarella abyssicola Scattered 100-2740 745-4970 EB/ST Filellum serratum Cosmopolitan 24—900 0-650 EB/ET Garveia crassa Tropical 625-900 741 SB/ST Halecium beanii . Cosmopolitan 0-780 0-1134 EB/ET H. sessile Cosmopolitan 425-430 0-752 EB/ET H. tenellum . : Cosmopolitan 0-900 0-730 EB/ET Peat glutinosa . Tropical 0-550 1-80 EB/ET H. polymorpha Tropical 0-900 0-89 EB/ET H. tuba Endemic 11-550 — EB/ET Hincksella echinocarpa Temperate 1610-2200 41-109 EB/ST Hydrocorella africana Endemic 0-500 — EB/ET Kirchenpaueria triangulata Temperate 111-1207 550-585 EB/ST Lafoea benthophila Scattered 425-430 672-3237 SB/ST L. dumosa Cosmopolitan 60-920 0—2200 EB/ET Modeeria rotunda Cosmopolitan 70-550 0-1240 EB/ET Nemertesia antennina Scattered 425-430 0-1779 EB/ET N. ramosa Scattered 11-700 0-872 EB/ET Plumularia prfonbritni Endemic 440 SB/ST P. mossambicae . Endemic 110-550 — EB/ET P. setacea Cosmopolitan 0-430 0-752 EB/ET Sertularella ieecanoas Temperate 200-900 183-672 EB/ST Stegolaria geniculata . Tropical 775-850 253-910 SB/ST Symplectoscyphus amphoriferus Scattered 550-900 161-1256 EB/ST S. arboriformis Endemic 10-420 = EB/ET S. paulensis . Temperate 347-1200 400-672 SB/ST Uniscyphus fragilis Endemic 360-420 — SB/ST Zygophylax africana . Endemic 137-850 — EB/ST Z. armata Scattered 48-440 183 EB/ET Z. brownei . Temperate 400-550 20-752 EB/ET Z. inconstans Endemic 360-450 =< SB/ST Z. sibogae Scattered 88-900 146-550 EB/ST Summary: Cosmopolitan . 9 Scattered 10 Tropical 10 Temperate 5 Endemic 11 Total . 45 178 ANNALS OF THE SOUTH AFRICAN MUSEUM Kirchenpaueria triangulata (temperate): minimum record Agulhas Bank 111 m, minimum east coast record 350 m. Sertularella leiocarpa (temperate): minimum record Tristan 183 m, minimum east coast record 280 m. Zygophylax africana (endemic): minimum east coast record 400 m. For the (10) species in this eurybathic/stenothermic category the limiting factor is obviously temperature rather than depth. The remaining (23) species are truly eurybathic and eurythermic, for they can occur in any temperature and at any depth. As might be expected, the majority (13) are cosmopolitan or scattered. Hedgpeth (1957) suggested that all the deep-sea benthic population will eventually prove to be cosmopolitan, but it is clear from this analysis that geographically cosmopolitan species are not necessarily eurythermic. The data for a selected group of 21 species were examined to see whether the depth distribution round the coast bore any relation to changing water temperatures. These were the species recorded from a minimum of 10 sectors and from depths extending below 100 m. Only 7.of the species so tested showed indications of an increasing minimum depth as one passes round the coast from west to east and then north; only 5 showed indications of an increasing maximum depth (Fig. 7, Table 7). This might be interpreted as an inability in a few species to survive in the warmer surface and shallow waters of the east coast. It is not surprising that none of these species belongs to the tropical component. Cladocarpus 1s a characteristic deep-water genus, and 12 of the 13 South African species extend down below 100 m, 9 below 200 m, 3 below 500 m and 7 do not occur in water shallower than 200 m. TABLE 7 The relationship between depth and distribution. A. The minimum recorded depth in metres; B. The maximum recorded depth in metres. Sectors Species ee ——— Component 0-5 6-10 11-15 16-20 21-24 25-30 31-35 A. Antennella quadriaurita -— 35 0 9 50 ~=100 — Scattered Halecium delicatulum . — — 0 4 AP MNO 55 Cosmopolitan Halopteris tuba. : — —— 22 11 Di 70 110 Endemic Salacia articulata . é 0 25 0 10 0 18 110° Endemic Sertularella flabellum . — — PS 10 27 124 — Endemic Sertularella megista . — —— 0 10 DBT 49 110 Endemic Symplectoscyphus arboriformis . . 0 10 11 11 40 DA — Endemic B. Antennella quadriaurita — 40 Te Os Ae SZS — Scattered Salacia articulata . y 0 23 SO CSS) ses EVaxeleinnvle Sertularella flabellum . — — 99 200 Dilieuene oe — Endemic Sertularella megista . — — 139 6 200) N64 219% 347 SEndennic Symplectoscyphus arboriformis . ; — 20 35.1120 90 420 — Endemic THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 179 WEST SOUTH EAST Salacia articulata DEPTH (m) Sertularella megista 20 25 30 SECTORS Fig. 7. The distribution according to depth of three species which show tendencies towards greater depths in warmer waters. Actual records. —_ 1 ) 10 ESTUARINE AND BRACK-WATER SPECIES Although many hydroids penetrate into the mouths of estuaries, especially on rocky substrata, only two species are truly estuarine and have not been found in the open sea. These are Bimeria fluminalis, which grows on mangrove roots in Richards Bay and St Lucia estuaries, and Hydractinia kaffraria, which is a commensal on the shell of Nassa kraussiana and occurs in estuaries from the Breede River mouth on the south coast to Durban on the east coast. The Limnomedusan, Ostroumovia inkermanica, is known from several brack-water lakes on the east coast, the hydroid from Lagoa Poelela, Mozam- bique, in a salinity of 8 parts per thousand, both hydroid and medusa from Nhlange Lake, Zululand, in 3—4 parts per thousand, and the medusa alone from 180 ANNALS OF THE SOUTH AFRICAN MUSEUM A @ Lovenella chiquitita @ Cladocarpus leloupi A Sertularella natalensis B Corhiza scotiae Salacia articulata Symplectoscyphus macrogonus Sertularella leiocarpa Fig. 8A-—D. Distribution of six representative endemic species: A. Lovenella chiquitita (west coast + False Bay), Cladocarpus leloupi (south coast) and Sertularella natalensis (east coast). B. Corhiza scotiae (west + south coast). C. Synthecium hians (south + east coast). D. Salacia articulata (west + south + east coast). E-F. Distribution of two representative temperate species: E. Symplectoscyphus macrogonus (west + south coast). F. Sertularella leiocarpa (west + south + east coast), also steno- thermic. THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 181 A B Lytocarpus philippinus Dynamena crisioides D Sertularia turbinata Sertularella arbuscula E Hebella scandens Amphisbetia operculata Fig. 9A-D. Distribution of four representative tropical species from east coast: A. Lytocarpus philippinus (to just north of Durban). B. Dynamena crisioides (to East London). C. Sertularia turbinata (to Mossel Bay). D. Sertularella arbuscula (to Saldanha Bay). E-F. Distribution of two representative cosmopolitan species: E. Hebella scandens (St Helena Bay to Mozambique). F. Amphisbetia operculata (Lideritz to Natal). 182 ANNALS OF THE SOUTH AFRICAN MUSEUM Lake St Lucia North in 10,4-10,8 parts per thousand. CONCLUSION The composition of the southern African hydroid fauna is summarized in Table 8, whence it can be seen that there is a mixture of endemic, tropical, temperate and cosmopolitan species in that order of abundance. The presence of a large endemic population, the particular distribution of some of the Western Indian Ocean tropical species (p. 169), and the apparent proliferation of subspecies in certain genera (p. 173), all point to an active evolutionary centre in the South African area. It is clear that there has also been a heavy invasion of tropical species from the Indo-West-Pacific region, and these species form the major component of the fauna of the east coast. Regarding the temperate and cosmopolitan species, and considering (1) the low relationship of the South African temperate fauna to that of the antarctic and subantarctic regions on the one hand, and to that of South America and Australasia on the other, (11) the practically certain role of ships in the introduction of at least three species out of 251 (1,2 %) to South Africa within a period of some 300 years (p. 170), TABLE 8 Summary of analysis of geographical components of South African hydroid fauna (from actual records, not range). West-plus-south East coast: Whole of coast: South Africa: Sectors 1-23 Sectors 24-34 Sectors 1-34 Component (171 species) (182 species) (251 species) No. Percentage |No. Percentage | No. Percentage TROPICAL Circumtropical 10 5,8 18 9,9 20 8,0 Indo-West-Pacific . 7 ARI SS a 23) 12,6 +39,6 | 24 9,6 >30,7 W. Indian Ocean 16 9,4 31 17,0 33 Sil! TEMPERATE Antitropical 10 5,8 5 27 11 4,4) Southern . 8 4,7 ~13,4 5 2 Os 10 4,0 -11,2 Atlantic 5 2,93) 2 ips f 2,8 ENDEMIC To onearea only .. Bye OSS 17 9,3 e520 4 D0STe) To the whole of South 38,0 25,8 325] Africa 20" S15 30F J oliG.s 30 12,0 UNCLASSIFIED Cosmopolitan Phe NDING 28: PANY Boe wiles Scattered . 28 - 16.4 (72 wa) | 23) ON ees eee THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 183 (iii) the fact that drifting weed and flotsam has probably performed a similar role throughout the ages, it is hardly necessary to evoke the Theory of Continental Drift to explain the distribution. : REPERENCES ARNAUD, F., ARNAUD, P. M., INTES, A. & Le Loeurr, P. 1976. Transport d’Invertébrés benthiques entre l’Afrique du Sud et Sainte Hélene par les laminaires (Phaeophyceae). Bull. Mus. natn. Hist. nat. Paris (3) 384, Ecol. gén. 30: 49-55. Beurois, J. 1975. 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(Clavopsellidae nov. fam.), ein neuer Hydroidpolyp aus der Ostsee und seine phylogenetische Bedeutung. Z. morph. Okol. Tiere 51: 227-260. U.S. NAVAL OCEANOGRAPHIC OFFICE. 1967. Monthly charts of mean, minimum, and maximum sea surface temperature of the Indian Ocean. Spec. Publ. nav. oceanogr. Off., Washington 99: 1-48. Watson, J. E. 1975. Hydroids of Bruny Island, southern Tasmania. Trans. R. Soc. S. 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Si AIO’ SU VLEL am Oro 8 YOY S&S vy & € | HLddid 1 SYOLOAS AAXODALVIO qaANVN (ponuNuods) Z XIGNddd VY 199 THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS (L061 x x < XK X poe191}e9S ‘aIyory) Viouup xpjdydoscZ ysvoo aA+S+M €c6l Xx | x x x :onumepugd ‘MOY vuDIIAfo xvi] dydo3sdZ ySvOd J | x > OrMmopu”q LLOL ‘PIII Sijisv4f snyddosiuyQ 9S8I Xx ueyodowsog ‘KPRIQOW BjNI14Jnu sisdoj144n I, 1 T = | ySvod | Aa+S+M x x x 1X s O1Wepu_ €S6] “TOMA MasdDM Y1ADINGN T, jesidoinue OBL] ‘Iopuelos x x x :o1eV1odwis ], 3 SULA xuduvy piavjnqn [ “OB MM Puy (O€8T :peordory, ‘rodsq) snsoo1naf snyddoasoady f, “ed MM PUT 8061 x x « XX x :peordory ‘ualle AA Sypnbap snyddosoady J (1S8I x x & x po10}e9S5 ‘ysnq) snsouuof sndsvd0d0y J, ySBO9 § 7961 “PLEIN x : IWOpu” sniojjaquin *f sndsvo0zay J, ysvoo A+$ (8S6I x x < & POruepugqd ‘pARIINAL) Supyos «f sndsv2020y T, yseoo sg (ZL8I ‘tonedusysit yy) x :Ormepugq sniafiunjd -f sndavz02ay [, a ——_— “ed MA PUL (918 ‘xnoinowrT) x x xX xX X x :jesrdo1ry, snsonxapf -f sndavoozay J, “ed M Purl (Ss “ >jeordo1y, ‘ysnq) smjnipoyep snduvz0zay J. Oeq' M ‘PUT (ZS81 SK :peordory, “ysng) S14Jsodidaaqg Sndivd02ay [ yseoo q+S x x x &X x so1wWepugq LS6I ‘PIII SuD1y wnioayjuds SO PUL MA (ZG)! x :yeordory, ‘SIAiee Wniasijuap wniaayiuds UIOYINOS €Z6] ‘MOYIEIS XO Ex OX :o}e10dwis [, sisuajnod snyddosojoajdud gy ureyyNos (8761 “OOSTIG24L) xX | X Xx x Xx :o}eIoduwiay, snuosoiopu snyddosojoajdud sg ysvoo a+StM (068 ‘touuRI1e) x x XxX x es x SOWIE pu_ stumsofisoqup snyddosojoajdud gy awl ANNALS OF THE SOUTH AFRICAN MUSEUM 200 x XX YS D pA p pu s Hiddid 1 peisi1e95 pe1sness ysvoo J :oTWiepug }Ssv00 | :olumepuq }Ssvoo | sorummepugq onuelyy rojerlodwoy, (9061 ‘UdIIV AA) S114Dj]1]0S SnzAzZZAZ SI6T ‘PILI av3oqis xvjAydoscz 8S6I ‘PIPTINAL wnjngipunful xpojAydosahz LLOL “PARTIAL Supjsuooui xojdydosazZ SS6T “PIPIINA pdsvr0Uulluas xojdydosaZ vol ‘pielfig lauMoig xvjdydosdzZ As SE GE WE OS GE KG 1K NG SCG | EG ES NG OG Cl BY AL OW SIL el CEU OL © 8 4 OY GS vw & G SYOLOAS I AAXOVDALVO qANVN (ponuljuod) 7 xIGNdddV 6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature (particularly Articles 22 and 51). Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed by the appropriate Latin (not English) abbreviation, ‘e.g. gen. nov., sp. nov., comb. nov., syn. nov., etc. An author’s name when cited must follow the name of the taxon without intervening punctuation and not be abbreviated; if the year is added, a comma must separate author’s name and year. The author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is transferred from its original genus. The name of a subsequent user of a scientific name must be separated from the scientific name by a colon. Synonymy arrangement should be according to chronology of names, i.e. all published scientific names by which the species previously has been designated are listed in chronological order, with all references to that name following in chronological order, e.g.: Family Nuculanidae Nuculana (Lembulus) bicuspidata (Gould, 1845) ae Figs 14-15A >» Nueula (Leda) bicuspidata Gould, 1845: 37. Leda plicifera A. Adams, 1856: 50. Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b). Nucula largillierti Philippi, 1861: 87. _ Leda‘bicuspidata: Nicklés, ‘1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. Note punctuation in the above example: : § » < comma/separates author’s name and-year ' » ‘semicolon separates’ more than one reference by the same author full stop separates references by different authors figures of plates are enclosed in parentheses to distinguish them from text-figures dash, not comma, separates consecutive numbers Synonymy arrangement according to chronology of bibliographic references, whereby the year is placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable. In describing new species, one specimen must be designated as the holotype; other speci- mens mentioned in the original description are to be designated paratypes; additional material not regarded as paratypes should be listed separately. The complete data (registration number, depository, description of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.: Holotype SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973. Note standard form of writing South African Museum registration numbers and date. 7. SPECIAL HOUSE RULES Capital initial letters (a) The Figures, Maps and Tables of the paper when referred to in the text e.g. ‘... the Figure depicting C. namacolus ...’; ‘. . . in C. namacolus (Fig. 10)...’ (b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded by initials or full names e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene (c) Scientific names, but not their vernacular derivatives e.g. Therocephalia, but therocephalian Punctuation should be loose, omitting all not strictly necessary Reference to the author should be expressed in the third person Roman numerals should be converted to arabic, except when forming part of the title of a book or article, such as “Revision of the Crustacea. Part VIII. The Amphipoda.’ Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial capital letter, provided the same generic name is used consecutively. Name of new genus or species is not to be included in the title: it should be included in the abstract, counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts. N. A. H. MILLARD THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS —— if Fa 0) | ~ VOLUME 74 PART in DECEMBER 1977 ISSN 0303-2515 i iM . MUS. COMP. ZOOL. l Lime a — i NOV 4 94cC°N A FIV A P) WMIWaReSi ry ANNALS } OUTH AFRICAN OF THE : MUSEUM CAPE TOWN INSTRUCTIONS TO AUTHORS 1. MATERIAL should be original and not published elsewhere, in whole or in part. 2. LAYOUT should be as follows: (a) Centred masthead to consist of Title: informative but concise, without abbreviations and not including the names of new genera or species Author’s(s’) name(s) Address(es) of author(s) (institution where work was carried out) Number of illustrations (figures, enumerated maps and tables, in this order) (b) Abstract of not more than 200 words, intelligible to the reader without reference to the text (c) Table of contents giving hierarchy of headings and subheadings (d) Introduction (e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents (f) Summary, if paper is lengthy (g) Acknowledgements (h) References (i) Abbreviations, where these are numerous 3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced with 2,5 cm margins all round. First lines of paragraphs should be indented. Tables and a list of . legends for illustrations should be typed separately, their positions indicated in the text. All pages should be numbered consecutively. Major headings of the paper are centred capitals; first subheadings are shouldered small capitals; second subheadings are shouldered italics; third subheadings are indented, shouldered italics. Further subdivisions should be avoided, as also enumeration (never roman numerals) of headings and abbreviations. Footnotes should be avoided unless they are short and essential. Only generic and specific names should be underlined to indicate italics; all other marking up should be left to editor and publisher. 4. ILLUSTRATIONS should be reducible to a size not exceeding 12 x 18 cm (19 cm including legend); the reduction or enlargement required should be indicated; originals larger than 35 x 47 cm should not be submitted; photographs should be rectangular in shape and final size. A metric scale should appear with all illustrations, otherwise magnification or reduction should be given in the legend; if the latter, then the final reduction or enlargement should be taken into consideration. All illustrations, whether line drawings or photographs, should be termed figures (plates are not printed; half-tones will appear in their proper place in the text) and numbered in a single series. Items of composite figures should be designated by capital letters; lettering of figures is not set in type and should be in lower-case letters. The number of the figure should be lightly marked in pencil on the back of each illustration. 5. REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: ‘Smith (1969) describes...’ ‘Smith (1969: 36, fig. 16) describes .. .’ “As described (Smith 1969a, 1969b; Jones 1971)’ “As described (Haughton & Broom 1927)...’ ‘As described (Haughton et al. 1927)...’ Note: no comma separating name and year Dagination indicated by colon, not p. names of joint authors connected by ampersand et al. in text for more than two joint authors, but names of all authors given in list of references. (b) Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, b, etc. to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a). For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list o, scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number (only if independently paged) in parentheses, Paeieee (first and last pages of article). Examples (note capitalization and punctuation) BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. FISCHER, P. —H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140. FIscHER, P.-H. , DUvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. Zen. 74: 627-634. Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320. Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51. THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. (continued inside back cover) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 74 ~ Band December 1977 Desember Part 7 Deel PAST AND PRESENT DISTRIBUTION OF SOME RODENT AND INSECTIVORE SPECIES IN THE SOUTHERN CAPE PROVINCE, SOUTH AFRICA: NEW INFORMATION By D. M. AVERY Cape Town Kaapstad The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular intervals as material becomes available Obtainable from the South African Museum, P.O. Box 61, Cape Town Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM word uitgegee in dele op ongereelde tye na beskikbaarheid van stof Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad OUT OF PRINT/UIT DRUK il, AO, GO), SID, ZG, 8, hoot), Ses, S, FD), 6(1, t.-p.i.), 771-4), 8, 9(1-2, 7), 10(1-3), 11(1-2, 5, 7, t.-p.i.), 15(4-5), 24(2), 27, 3111-3), 32(5), 33 Price of this part/Prys van hierdie deel R1,80 Trustees of the South African Museum © Trustees van die Suid-Afrikaanse Museum 1977 ISBN 0 908407 28 9 Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk., Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap PAST AND PRESENT DISTRIBUTION OF SOME RODENT AND INSECTIVORE SPECIES IN THE SOUTHERN CAPE PROVINCE, SOUTH AFRICA: NEW INFORMATION By D. M. AVERY South African Museum, Cape Town (With 3 figures and 2 tables) [MS. accepted 11 August 1977] ABSTRACT Micromammalian material from archaeological sites and modern owl roosts in the Cango valley and Gansbaai area of the southern Cape Province provides new information concerning the past and present distribution of certain species of rodents and insectivores. Modern material from the Cango valley provides evidence of a link between widely separated popula- tions of several species and shows Praomys natalensis (multimammate mouse) to be co-existing with P. verreauxi (Verreaux’s mouse). The archaeological evidence indicates that Saccostomus campestris (Cape pouched mouse) and Steatomys pentonyx (Cape fat mouse) arrived in the southern Cape only at the beginning of the Holocene about 10000 years BP and that P. natalensis perhaps entered the Cango valley with the advent of farming in the area around 200 BP. On the other hand, Tatera afra (Cape gerbil), Gerbillurus paeba (pygmy gerbil) and possibly Otomys unisulcatus (bush Karoo rat) disappeared from the Cango valley some time after about 1 600 BP. CONTENTS PAGE Introduction : ; sc 20h Present distribution . ae DOS Past distribution ; Lo DOF Conclusion . : : e208 Acknowledgements . . 208 References . i : ware 209 INTRODUCTION The evidence put forward below became available during the course of a study of micromammalian remains from archaeological sites and comparable modern owl roosts in the southern Cape Province. Two areas are considered, the Cango valley (3322AC) near Oudtshoorn, and the Gansbaai area (3419CB) on the Walker Bay coast. For the Cango valley (Fig. 2) material was available from the archaeological site Boomplaas A (33°23’S 22°11’E) which represents well in excess of 40 000 years of accumulated deposit during the Upper Pleisto- cene and Holocene (Deacon & Brooker 1976: 211) and from five modern owl roosts. Two of these, Boomplaas B and C which are adjoining rock-shelters, are within 100 m of Boomplaas A and in the same cliff. Two others, Nooitgedacht A 201 Ann. S. Afr. Mus. 74 (7) 1977: 201-209, 3 figs, 2 tables. 202 ANNALS OF THE SOUTH AFRICAN MUSEUM B Praomys verreauxi C Steatomys pentonyx P. natalensis 5 F Otomys laminatus G Dasymys incomtus Fig. 1. Distribution of some rodent and insectivore species showing the relationship of the Cango valley (C) and Gansbaai (G) to existing ranges. A is based on Meester & Lambrechts (1971) and B and D-G are based on Davis (1974). DISTRIBUTION OF SOME RODENT AND INSECTIVORE SPECIES 203 Fig. 2. The position of the sites in the Cango valley. BPA, BPB and BPC = Boomplaas A, B and C; NGA and NGB = Nooitgedacht A and B; OG = Osgat. Metric approximations of 200 ft contours are given. and B (33°22’S 22°10’E) which are also adjoining rock-shelters, are situated 2,5 km north-west of Boomplaas, whilst the fifth site, Osgat (33°22’S 22°11’E), lies 1,3 km north-east of Nooitgedacht and 5 km north of Boomplaas. For the Gansbaai area (Fig. 3) archaeological material was available from Die Kelders 1 (34°32’S 19°22’E), covering a period from about 80000 BP to 45000 BP (years before present) during the Middle Stone Age (MSA) and from about 2000 BP upwards during the Late Stone Age (LSA) (Tankard & Schweitzer 1974: 367; Tankard 1976: 155), and from Byeneskranskop | (34°35’S 19°28’E) which was occupied from about 12000 BP upwards (F. R. Schweitzer pers. comm.) and which lies some 9,5 km south-east of Die Kelders. The modern sample was taken from Byeneskranskop 2, a rock-shelter situated just below the archaeological site and set in the same cliff. Total minimum numbers of individuals represented in the various samples are given in Table 1. It appears from Davis’s (1974) distribution maps that little collecting or recording has been undertaken in the two areas discussed here, although Grindley et al. (1973) collected owl pellets from near Stanford some 15 km north of Gansbaai. For this reason the recent information has proved useful both for ANNALS OF THE SOUTH AFRICAN MUSEUM 204 “UOAIZ OIC SINO}UOD 3J ONT JO SUOTeUIxOIdde sIN}OY “7 pue | doysuvrysouskg = JNA ‘I SOppey ad = INd "BOI IBEQSULDH OY} Ul Sous OY} JO UOTSOd OY], “¢ “SI Aeg JOHICM DISTRIBUTION OF SOME RODENT AND INSECTIVORE SPECIES 205 consolidating known ranges and for filling in gaps in others less well known. Remains from archaeological contexts provide some indication of when various species arrived in or, in some cases, disappeared from the two areas. PRESENT DISTRIBUTION The presence of many of the species listed in Table 1 was predictable and in these cases the evidence merely adds another locality to an already clearly established range. In some cases, however, the sparseness of the existing record adds significance to the present information. Acomys subspinosus (Cape spiny mouse), Saccostomus campestris (Cape pouched mouse), Dendromus melanotis (grey pygmy climbing mouse), D. mesomelas (chestnut climbing mouse), Crocidura flavescens (red musk shrew), C. cyanea cyanea (reddish-grey musk shrew), Suncus varilla (lesser grey dwarf shrew) and, to some extent, Praomys verreauxi (Verreaux’s mouse) fall into this category. The evidence confirms the ranges of A. subspinosus and S. campestris and also the eastward extent of the range of P. verreauxi as given in Davis (1974: 158, 163, 160 respectively). In the case of D. melanotis, D. mesomelas (Davis 1974: 166, 168) and C. flavescens (Meester 1963: 34) confirmation is provided for the existence of links between populations in the south-west and others in the south-east. For S. varilla the present information provides not only confirmation of similar linking but also constitutes a southwards extension of the known range given by Meester & Lambrechts (1971: 11) (Fig. 1A). Further to this, and even more important, is another group of species which has been discovered in the Cango valley. For this group evidence of links between isolated populations had not previously been forthcoming. Dasymys incomtus (shaggy swamp rat), Mystromys albicaudatus (white-tailed rat), Otomys laminatus (laminate vlei rat), and O. saundersae (Saunders’s vlei rat) comprise this group (Fig. 1D-G). It is interesting to note that Davis (1974: 166 et seq.) predicted that the gap was more apparent than real in several cases. Most of these species are not frequently caught by owls (Vernon 1972: 119). Assuming this to represent roughly the actual proportions of animals present, it is likely that still more records will become available with more intensive work. Steatomys pentonyx (Cape fat mouse) was also unexpected in the Cango valley but the situation regarding Steatomys spp. is complicated by the fact that this genus has long been in need of revision (Davis 1962: 73). Currently the south-western Cape population is referred to S. pentonyx and the more northern material to at least two other species. It is thought most likely that the present material represents a considerable eastward extension of the range of S. pentonyx (Fig. 1C). Praomys natalensis (multimammate mouse) has not been thought to occur further west than about 24°E in the southern Cape, the more westerly area being occupied by P. verreauxi (Verreaux’s mouse) (Fig. 1B). Indeed, Davis (1974: 160) doubted Pocock’s identification of P. natalensis from Oudtshoorn, considering it more likely to be referable to P. verreauxi. However, material ANNALS OF THE SOUTH AFRICAN MUSEUM 206 / q}00} SUIySeT SMel— poyenuerayipun ‘ ¥ 66L 9b SO8T 66 El Lp 6c 09% LOS ses VvOL 8Il Ve poyuesoides S[eNpPIAIPUL JO STOqUINU VINUUTUTU [20], a =—= =e — — x x — x = x ossnowuop oded S1IDJNIO SnAniydvAyH x x x x — x x x x x x osnoul ploy podins ound stuopqouyy aa = = = = — — — — — — yel YOR SNJIDA SNIJOY x x x x x x x x x x x OSNOUI S.XNVITIOA IXNDAAAAA “I ae == — — x x x x x nae = OSNOU d}VWUWIeUINI [NAY SisuajojoU SAUODAT x ae x x aa x x x x x x esnoul JIeMG Saplojnuiu SnPT x —= x x = x x = = x x yer durems Asseys Snjwooul skudspg x = x = Xx x Xx x x Xx x yer yoo enbewen, sisuanbouipu sduoyjap x -— x x x x x x x x x osnow Aurds sdeD snsouidsqns sduoop aa ca = = = = = == ei P, i 0 0 O 3) a eee | Nee 1 t eae UE | Si, Se, St represent internal, external and terminal spines, or setae respectively. The number of setae is shown in arabic numerals and spines in roman numerals. The first leg has four less spines and one less seta than the remainder. The fourth leg has a fringe of fine hairs on the medial margins of the second basipod segment and the first exopod segment. All the setae on the swimming legs are plumose, and the setae on the endopods are much more slender than the remainder. The fifth pair of legs of the female (Fig. 1D) are uniramous, three-segmented and symmetrical. The second segment bears an outer seta and the third segment bears a stout terminal spine and a subsidiary basal spine. The terminal spine is not articulated but fused to the third segment and it is finely barbed distally. Male Length 1,80-1,85 mm. Prosome (Fig. 2C, F) somewhat different from that of the female, with the head more evenly tapered but with a cervical groove like that of the female. Forehead shallowly sloped. Rostral filaments absent. Eyes not prominent and dark brown. All five pedigerous segments distinct but fourth and fifth partly fused. Posterior angles of metasome rounded and not produced. Urosome five-segmented of which the first is largest. Caudal rami (Fig. 2E) symmetrical and about five times as long as wide. The urosomes of several males were characteristically twisted to the right. Left antenna | (Fig 2A-B) fifteen-segmented and similar to that of the female. Right antenna | (Fig. 2D) geniculate, with fifteen separate segments, two of which are beyond the articulation. Segments 1-6, 19-21 and 22-25 fused. Prominent setae appear on proximal segments and on segments 14, 16, 18 and 21. The development of the specialization of the geniculate antenna may be observed in the right antenna 1 of a stage V male copepodite (Fig. 4H). A NEW SPECIES OF TORTANUS FROM SOUTH AFRICA 223 —, = S& SS WS 7 —jy \ EEE EEE =a Fig. 2. Tortanus capensis sp. noy. Male. A-B. Left antenna. C. Dorsal view. D. Right antenna. E. Caudal rami. F. Lateral view. Segments 11-12 and 23-25 are fused at that stage, but segments 18 to 22 are free and there are prominent spines on segments 18 and 20. It would appear that the spine on segment 18 forms the serrated lamella on that segment of the geniculate antenna of the adult male. Male fifth legs (Fig 3A—C) uniramous and asymmetrical with the right leg three-segmented and the left leg four-segmented. The first segment of the right leg is swollen and produced into a rounded projection at its medial distal corner. 224 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 3. Tortanus capensis sp. nov. Male. A. Fifthlegs. B. Right fifth leg. C. Left fifth leg. Female. D. Antenna. E. Mandible and palp. F. First maxillary. G. Second maxilla (end of one seta enlarged). H. Maxilliped. The second segment bears a large bilobed medial projection near its distal end, which bears two setae. The terminal segment is in the form of a sub-chelate claw, strongly chitinized near the tip and bearing two medial setae, and two tiny bristles near its tip. The first segment of the left leg is short. The second segment is longer and curved and bears a medial and an outer seta. The third segment is strongly curved tapers distally, and has an angular swelling proximally bearing a medial A NEW SPECIES OF TORTANUS FROM SOUTH AFRICA DUDES \ ea Fig. 4. Tortanus capensis sp. nov. Juveniles. A. Stage IV female, dorsal view. B. Fifth legs, stage IV female. C. Fifth legs stage V female. D. Stage IV male, dorsal view. E. Fifth legs, stage IV male, tip enlarged. F. Stage V male, dorsal view. G. Fifth legs, stage V male. H. Right antenna I, stage V male. seta. The terminal segment is even more strongly curved and bears a finely serrate terminal spine, two outer marginal spinules, two inner marginal setae, and a large number of tiny bristles. Spermatophore 300-380 pm long. Juvenile stages (Fig. 4A—-H) Copepodite stages including stages four and five of both males and females were obtained. The changes that occur in the development of the urosome and the fifth legs particularly are apparent in the figures. Lengths: Stage IV female 226 ANNALS OF THE SOUTH AFRICAN MUSEUM 1,45 mm, Stage V female 1,75 mm, Stage IV male 1,12 mm, Stage V male 1,50 mm. The ornamentation of the swimming legs of the Stage IV and Stage V copepodites is represented in Tables 2 and 3. TABLE 2 Copepodite Stage IV. Ornamentation of swimming legs. Protopod Endopod Exopod P, | 0 0 0 6 2 1 1 0 5 I I P, 1 0 0) 0 6 2; 1 1 I 6. ALLY AE P; 1 0 0 0 5 D I 1 I 5 i If P, I 0 0 0 4 D 1 1 I 5) YT Ui Si, Se, St represent internal, external and terminal spines, or setae respectively. The number of setae is shown in arabic numerals and spines in roman numerals. TABLE 3 Copepodite Stage V. Ornamentation of swimming legs. Protopod Endopod Exopod 1 Ze 1 Z 1 2, 3 Si Se Si Se Si Se Si St Se Si Se Si Se Si St Se P, I 0 O 3) 0) SN ART io ke O 2 etna Ug | P, he 0 O 30 Sri De el Ua | eek eed UCP edt P, LEO 0 0 3 O Bae hal ioe | 1 ean ef +. Tk SF P, iL @ 0 O 3-00 Bi.) Wate al he ye List 53> lle cED Si, Se, St represent internal, external and terminal spines, or setae respectively. The number of setae is shown in arabic numerals and spines in roman numerals. Remarks Tortanus capensis is apparently the twenty-first species of this peculiar genus of copepods to have been described. The genus Corynura was established by Brady (1883) for two species obtained by the Challenger expedition in the Philippine islands. This generic name which was preoccupied was replaced by the name TJortanus (from tortus and anus) by Giesbrecht & Schmeil (1898). The following species are now included in this genus: Tortanus barbatus (Brady), 1883 (=denticulatus Giesbrecht); Tortanus gracilis (Brady), 1883; Tortanus forcipatus (Giesbrecht), 1889; Tortanus recticauda Giesbrecht, 1889; Tortanus discaudatus (Thompson & A. Scott), 1897; Tortanus setacaudatus Williams, 1906; Tortanus murrayi A. Scott, 1909; Tortanus brevipes A. Scott, 1909; Tortanus (Atortus sub. gen. nov.) tropicus Sewell, 1932; Tortanus derjugini Smirnov, 1935 (see Brodsky 1950); “Tortanus sp.’ Colefax, 1940 Um Dakin & A NEW SPECIES OF TORTANUS FROM SOUTH AFRICA UE Colefax 1940); Tortanus longipes Brodsky, 1948; Tortanus vermiculus Shen, 1955; Tortanus spinicaudatus Shen & Bai, 1956; Tortanus denticulatus Shen & Lee, 1963; Tortanus compernis Gonzales & Bowman, 1965; Tortanus rubidus Tanaka, 1965; Tortanus giesbrechti Jones & Park, 1968; Tortanus scaphus Bowman, 1971; Tortanus lophus Bowman, 1971; and Tortanus capensis sp. nov. (S77). Tortanus capensis is the first species of the genus to have been found anywhere round the coasts of Africa, south of the Red Sea. It appears to be most closely related to Tortanus recticauda from the Red Sea (Giesbrecht 1889) and Tortanus murrayi, scaphus and lophus from the East Indies (A. Scott 1909; Bowman 1971), but it differs from each of them in many characters. Tortanus capensis may be most readily distinguished by the fifth legs of either the male or the female, which are quite distinct from those of any of the described species of Tortanus. Specimens of Tortanus capensis from the Swartkops estuary on the south- east coast of South Africa showed a few minor differences in comparison with the type material from Langebaan Lagoon. In the two-segmented urosome of the female and the character of the male fifth legs Tortanus capensis might be placed in Steuer’s group 3 which he established in his 1926 revision of the genus Jortanus. This is equivalent to Sewell’s (1932) subgenus Afortus (Bowman 1971). ACKNOWLEDGEMENTS I thank Mrs S. Grindley and Mr T. Wooldridge for assistance and the South African National Committee for Oceanographic Research for financial support. REFERENCES Bowman, T. E. 1971. Tortanus scaphus and Tortanus lophus, new Pacific planktonic copepods, with notes on Tortanus murrayi (Calanoida, Tortanidae). Pacif. Sci. 25 (4): 521-528. Brapy, G. S. 1883. Report on the Copepoda collected by H.M.S. Challenger during the years 1873 to 1876. Challenger Rep. 1873-76 8: 1-142. Bropsky, K. A. 1948. T. Jongipes sp. nov. from the Sea of Japan. Proc. Pacif. Sci. Res. Inst. Fish Oceanogr. 26: 80. (In Russian.) Bropsky, K. A. 1950. Calanoida of the far eastern and polar seas of the U.S.S.R. Tabl. anal. faune U.R.S.S. Zool. Inst. Acad. Sci. 35: 1-442. (In Russian.) Dakin, W. J. & CoLeFAx, A. N. 1940. The plankton of the Australian coastal waters of New South Wales. Publ. Univ. Sydney (Zool.) 1 (1): 1-215. GIESBRECHT, W. 1889. Elenco dei Copepodi pelagici. Atti. R. Ac. Lincei (4) 5: 24-29. GIESBRECHT, W. & SCHMEIL, O. 1898. Copepoda. 1. Gymnoplea. Tierreich 6: 1-169. GONZALES, J. G. & BOwMan, T. E. 1965. Planktonic copepods from Bahia Fosforescente, Puerto Rico, and adjacent waters. Proc. U.S. nat. Mus. 117: 241-303. JONES, E. C. & Park, T. S. 1968. A new species of Tortanus (Calanoida) from Pago Pago Harbour, American Samoa. Crustaceana (Suppl. 1): 152-158. Scott, A. 1909. The Copepoda of the Siboga expedition. Siboga Exped. Monogr. 29 (A): 1-323. SEWELL, R. B. S. 1932. The Copepoda of Indian seas. Calanoida. Mem. Indian Mus. 10: 400-402. 228 ANNALS OF THE SOUTH AFRICAN MUSEUM SHEN, C. J. 1955. On some marine crustaceans from the coastal water of Fenghsien, Kiangsu Province. Acta. zool. Sin. 7: 75-100. (In Chinese & English.) SHEN, C. & Bal, S. 1956. The marine Copepoda from the spawning ground of Pnueumatophorus japonicus (Houttuyn) off Chefoo, China. Acta. zool. Sin. 8: 177-234. SHEN, C. J. & Lee, F. S. 1963. The estaurine copepoda of Chiekong and Kaikong Rivers, Kwangtung Province, China. Acta. zool. Sin. 15: 571-596. STEUEUR, A. 1926. Revision der Copepodengattung Tortanus Giesbr. Boll. Soc. Adriatica Sci. Nat. 29: 49-69. TANAKA, O. 1965. The pelagic copepods of the Izu region, Middle Japan. Systematic account XIII. Parapontellidae, Acartiidae and Tortanidae. Publ. Seto mar. biol. Lab. 12: 379-408. THompson, I. C. & Scott, A. 1897. On the plankton collected continuously during the two traverses of the North Atlantic in the summer of 1897. Trans. Liverpool biol. Soc. 12: 80. WILLIAMS, L. W. 1906. Notes on marine Copepoda of Rhode Island. Amer. Nat. 40 (447): 639-660. 6. SYSTEMATIC papers must conform to the International code of zoological nomenclature (particularly Articles 22 and 51). Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov., etc. An author’s name when cited must follow the name of the taxon without intervening punctuation and not be abbreviated; if the year is added, a comma must separate author’s name and year. The author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is transferred from its original genus. The name of a subsequent user of a scientific name must be separated from the scientific name by a colon. Synonymy arrangement should be according to chronology of names, i.e. all published scientific names by which the species previously has been designated are listed in chronological order, with all references to that name following in chronological order, e.g.: Family Nuculanidae Nuculana (Lembulus) bicuspidata (Gould, 1845) Figs 14-15A Nucula (Leda) bicuspidata Gould, 1845: 37. Leda plicifera A. Adams, 1856: 50. Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b). Nucula largillierti Philippi, 1861: 87. Leda bicuspidata: Nicklés, 1950: 163, fig. 301; ‘1955: 110. Barnard, 1964: 234, figs 8-9. Note punctuation in the above example: comma separates author’s name and year semicolon separates more than one reference by the same author full stop separates references by different authors figures of plates are enclosed in parentheses to distinguish them from text-figures dash, not comma, separates consecutive numbers Synonymy arrangement according to chronology of bibliographic references, whereby the year is placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable. In describing new species, one specimen must be designated as the holotype; other speci- mens mentioned in the original description are to be designated paratypes; additional material not regarded as paratypes should be listed separately. The complete data (registration number, depository, description of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.: Polehvpe SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973. Note standard form of writing South African Museum registration numbers and date. 7. SPECIAL HOUSE RULES Capital initial letters (a) The Figures, Maps and Tables of the paper when referred to in the text _ e.g. ‘... the Figure depicting C. namacolus...’; ‘. . . in C. namacolus (Fig. 10)...’ (b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded by initials or full names e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene (c) Scientific names, but not their vernacular derivatives e.g. Therocephalia, but therocephalian Punctuation should be loose, omitting all not strictly necessary Reference to the author should be expressed in the third person Roman numerals should be converted to arabic, except when forming part of the title of a book or article, such as ‘Revision of the Crustacea. Part VIII. The Amphipoda.’ ‘ Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial capital letter, provided the same generic name is used consecutively. Name of new genus or species is not to be included in the title: it should be included in the abstract, counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts. JOHN R. GRINDLEY A NEW SPECIES OF TORTANUS (CRUSTACEA, COPEPODA) FROM SOUTH AFRICA | rt SA VOLUME 74 PART 10 JANUARY 1978 | | | | MUS. COMP. Oe ee LIZRARY APR 28 19/8 ARV AR UNIVERSITY ~ANNALS OF THE SOUTH AFRICAN MUSEUM CAPE TOWN INSTRUCTIONS TO AUTHORS 1. MATERIAL should be original and not published elsewhere, in whole or in part. 2. LAYOUT should be as follows: (a) Centred masthead to consist of Title: informative but concise, without abbreviations and not including the names of new genera or species Author’s(s’) name(s) Address(es) of author(s) (institution where work was carried out) Number of illustrations (figures, enumerated maps and tables, in this order) (b) Abstract of not more than 200 words, intelligible to the reader without reference to the text (c) Table of contents giving hierarchy of headings and subheadings (d) Introduction (e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents (f) Summary, if paper is lengthy (g) Acknowledgements (h) References (i) Abbreviations, where these are numerous 3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced with 2,5 cm margins all round. First lines of paragraphs should be indented. Tables and a list of legends for illustrations should be typed separately, their positions indicated in the text. All pages should be numbered consecutively. Major headings of the paper are centred capitals; first subheadings are shouldered small capitals; second subheadings are shouldered italics; third subheadings are indented, shouldered . italics. Further subdivisions should be avoided, as also enumeration (never roman numerals) of headings and abbreviations. Footnotes should be avoided unless they are short and essential. Only generic and specific names should be underlined to indicate italics; all other marking up should be left to editor and publisher. 4. ILLUSTRATIONS should be reducible to a size not exceeding 12 x 18 cm (19 cm including legend); the reduction or enlargement required should be indicated; originals larger than 35 x 47 cm should not be submitted; photographs should be rectangular in shape and final size. 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REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: ‘Smith (1969) describes...’ ‘Smith (1969: 36, fig. 16) describes...’ “As described (Smith 1969a, 19696; Jones 1971)’ ‘As described (Haughton & Broom 1927)...’ ‘As described (Haughton et al. 1927)...’ Note: no comma separating name and year Dagination indicated by colon, not p. names of joint authors connected by ampersand et al. in text for more than two joint authors, but names of all authors given in list of references. (b) Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, b, etc. to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a). For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list o, Scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number (only if independently paged) in parentheses, pagination (first and last pages of article). Examples (note capitalization and punctuation) BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140. FiscHER, P.-H., DuvAL, M. & RAFFy, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. gén. 74: 627-634. Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320. : Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51. THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. (continued inside back cover) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 74 Band January 1978 Januarie Part 10 #£4Deel NOTES ON THE SCORPION FAUNA OF THE CAPE PART 3 SOME OBSERVATIONS ON THE DISTRIBUTION AND BIOLOGY OF SCORPIONS ON TABLE MOUNTAIN By FE. B. EASTWOOD Cape Town Kaapstad The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular intervals as material becomes available Obtainable from the South African Museum, P.O. Box 61, Cape Town Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM word uitgegee in dele op ongereelde tye na beskikbaarheid van stof Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad OUT OF PRINT/UIT DRUK 1, 2(1-3, 5-8), 31-2, 4-5, 8, t.—p.i.), 51-3, 5, 7-9), 6(, t.—p.i.), 7-4), 8, 911-2, 7), 101-3), 11(1-2, 5, 7, t.—p.i.), 15(4—5), 24(2), 27, 31(1-3), 32(5), 33 Price of this part/Prys van hierdie deel R2,30 Trustees of the South African Museum © Trustees van die Suid-Afrikaanse Museum 1978 ISBN 0 908407 31 9 Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk., Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap NOTES ON THE SCORPION FAUNA OF THE CAPE PAR 3 SOME OBSERVATIONS ON THE DISTRIBUTION AND BIOLOGY OF SCORPIONS ON TABLE MOUNTAIN By E. B. EASTWOOD South African Museum, Cape Town (With 12 figures and 4 tables) [MS. accepted 18 August 1977] ABSTRACT A survey of the scorpion fauna of Table Mountain on the Cape Peninsula was carried out. Four of the six species recorded from this area were found. Distribution and habitat preferences are discussed with reference to topography and vegetation. Some observations on behaviour in the field and laboratory are recorded. A regional key is given. CONTENTS _ PAGE Introduction : ; : : ; : : PS) Description of the study area . : ; A an 230 Materials and methods . ; : ; 4 e723) Distribution and habitat . ; ; : ; le 3 7) Scorpion shelters : : : ; : : 3 236 Coexistence : : 3 : f : : eA) Scorpion predators . é : : , : Fae Al Some observations on behaviour . d : We Key to the scorpions of Table Mountain . . 244 Conclusions 4 : , : : , : e246 Acknowledgements . : ‘ . , . 5 Vay References . ; ; i 3 f 4 5 ot ead INTRODUCTION Six species in two families of scorpions have been recorded from Table Mountain. Opisthophthalmus capensis (Herbst) was recorded by Pocock (1896), Purcell (1899) and Lawrence (1955); Uroplectes insignis Pocock by Pocock (1890, 1896), Kraeplin (1894, 1899), Purcell (1901) and Hewitt (1918); U. lineatus (Koch) was reported by Purcell (1901) and Kraeplin (1908); U. variegatus (Koch) by Pocock (1896) and Hewitt (1918), and Parabuthus capensis (Ehrenberg) by Hewitt (1918) and Eastwood (19775). A juvenile specimen from Signal Hill was identified as U. vittatus (Thorell) by Penther (1900) but it is very likely that this record pertains to U. lineatus. The work on Cape scorpions by early taxonomists was confined to morphological and geographical considerations. With later descriptions of new forms, a reappraisal of scorpion taxonomy has become very complex without the relevant background on ecology and behaviour. A survey was carried out by regular collecting over a period of one year. 229 Ann. S. Afr. Mus. 74 (10), 1978: 229-248, 12 figs, 4 tables. 230 ANNALS OF THE SOUTH AFRICAN MUSEUM To complement field observations, live scorpions were kept in the laboratory. Reports are given on diversity and distribution, coexistence, habitats and shelters, and where possible notes are given on behaviour. A regional key was constructed for field or laboratory use. DESCRIPTION OF THE STUDY AREA Location Table Mountain forms the northern end of the mountainous ridge of the Cape Peninsula and lies approximately 33°57’S 18°25’E. Topography, geology and soils Table Mountain forms a plateau of Table Mountain Sandstone, which is surrounded by cliffs giving way to gentle slopes. The southern end is deeply dissected by the Disa stream forming the Orangekloof subarea. The sandstone overlies a layer of Malmesbury shale which in turn rests on Cape Granite. The last two formations give rise to the gentle slopes (Moll & Campbell 1976), the areas with which this study is primarily concerned. Soil types are dependent on the parent material, and those derived from the sandstone are usually acidic sands, the granite and shale derived soils contain more clay and are less acidic. The lower slopes are covered with sandstone debris which give rise to soils of mixed origin (Moll & Campbell 1976). Climate Table Mountain lies in the Winter Rainfall Region characterized by hot, dry summers and cool, wet winters. The topography plays an important part in the climate. Rainfall for the different subareas are as follows: Summit and eastern slope . : . 1420-1780 mm Western, northern slopes and Siena Hill RN: 889 mm Southern subarea : ; ; 1227 mm (from Moll & Campbell 1976) Temperature differences over the whole area are not as pronounced. Vegetation The most extensive plant community is macchia or fynbos, consisting of Proteaceae, Ericaceae, Restionaceae, Geraneaceae and other families. There are also small areas of natural forest communities as well as extensively cultivated alien forest vegetation consisting of Pinus spp., Eucalyptus spp., and Acacia spp. mainly on the eastern slopes and lower northern slopes. There are also areas of Hakea sp. on the western slopes. The classification of plant communities is given in Figure 1. MATERIALS AND METHODS Six study subareas were defined: 1. North-facing slopes 4. East-facing slopes 2. West-facing slopes 5. Plateau subarea 3. Southern subarea (Orangekloof) 6. Signal Hill subarea 231 NOTES ON THE SCORPION FAUNA OF THE CAPE +7404 eae ce +t fe: 2 oa - + aoe fg 2 eS Dry proteoid Oa =) — LS ) (eo) —J) Short restoid Alien forest Map of Table Mountain showing approximate boundaries of plant communities. Big: 1. Each subarea was divided into a number of transects, 0,5 km wide, extend- ing from the base of the mountain to the sandstone cliffs. The table or plateau not divided up in this way. Was soil b) logs and other debris, and search- 9 Collecting was carried out by moving up a transect and collecting samples Records were kept of altitude by turning over surface objects such as rocks ing under the loose bark of dead trees. types and vegetation. D3) ANNALS OF THE SOUTH AFRICAN MUSEUM Several areas were isolated for the more detailed studies of demes or popula- tion aggregates of Opisthophthalmus and Uroplectes. In these areas burrow morphology, run morphology, feeding and coexistence of species were studied. In order to observe the feeding and burrowing behaviour of O. capensis, studies were carried out in the field where possible. In addition a number of specimens were kept in specially prepared boxes in the laboratory to complement field observations of feeding and burrowing. The morphology of burrows was studied by excavation and measurement. The numbers of observed specimens in relation to vegetation types is given in Table 1. TABLE | Comparison of relative numbers of scorpions observed in each of the six study subareas and seven vegetation types. O. capensis U. lineatus U. insignis U. variegatus Study subarea Vegetation type ©=£_ TT oa 2 oF < § © 6 1g North- and west- Dry proteoid OS) Zi Dap Bil S54: facing slopes, and a Signal Hill . . . Lowscrub WS) 3 DS South-facing slopes and Plateau subareas . Tall proteoid ad Sy Plateau subarea . Short restoid 3 East-facing Moist proteoid Dnwe'S slopes subarea. . _- Se ——— Forest scrub- forest Ye 1728 North- and east- facing subareas . Alien forest I Rotale 3 119 156 45 9 DISTRIBUTION AND HABITAT One scorpionid species, viz. Opisthophthalmus capensis, was found on Table Mountain, while other recorded species were buthids. O. capensis has also been found on the lower-lying Cape Flats and recorded randomly in the Cape Peninsula. This and allied species extend into Namaqualand (Eastwood 1977a). Another species, O. macer Thorell, extends from False Bay eastwards to Port Elizabeth (Lawrence 1955). Parabuthus capensis (Ehrenberg) extends from Clan- william in the north to Cape Town, and eastwards to Tulbagh (Eastwood 19775). Another widely distributed species, P. granulatus (Hemprich & Ehrenberg), is found on the Cape Flats to the north and has recently been collected in the suburban areas north of the city. Uroplectes lineatus is reported by Lawrence (1955) to extend from Cape Town eastwards to Mossel Bay, and northwards as far as Tulbagh and Worcester. U. insignis is confined to Table Mountain. U. variegatus, another local species, is recorded from the Cape Peninsula and its immediate neighbourhood, and Langebaan in the north (Lawrence 1955). In the present survey of the Table Mountain study area, O. capensis was located all along the west-facing subarea, the Signal Hill subarea and the NOTES ON THE SCORPION FAUNA OF THE CAPE 233 @ U. lineatus O U. insignis @ U. variegatus O O. capensis 0 V. BRANCO Fig. 2. Map of Table Mountain showing distribution of scorpions collected during the survey. western extremity of the north-facing subarea. There was a tendency for demes to be found in fairly open areas of the dry proteoid communities, i.e. below the 400 m contour, where the slopes were not too steep and ground cover plentiful, but they also occurred in the low scrub communities where the ground cover and substratum were suitable (Fig. 3). U. lineatus is the most widely distributed scorpion on Table Mountain (Fig. 2). As well as occupying widely differing habitats it was found in all the 234 ANNALS OF THE SOUTH AFRICAN MUSEUM me Fig. 3. Typical habitat of Opisthophthalmus capensis, Uroplectes lineatus and U. variegatus. Inthe foreground is the dry proteoid plant community and further back the lowscrub community. subareas under study, and at varying altitudes where ground cover was adequate. The demes were not as obvious as those of O. capensis and in many cases only a few specimens could be found in a large area. This could be accounted for by the immovability of much of the ground cover, or thick scrub and grass, making collection difficult. Thus U. /ineatus was found in the dry proteoid, low scrub, tall proteoid, moist proteoid, short restoid and forest scrub-forest plant com- munities. This species appeared to be most abundant in the dry proteoid communities on the north- and west-facing slopes. Only one specimen was found in alien forest. It will adapt to many situations as shown by the range of habitats it occupies, and is often reported to be found in the homes of residents near Table Mountain and even at the 600 and 700 m contours where access for the collector is allowed only by traverses along rocky ledges. Specimens of U. variegatus were found in the same areas as occupied by O. capensis in the dry proteoid communities (Fig. 2). U. insignis is taxonomically very near U. lineatus. Purcell (1902) regarded it as a distinct species. He recorded specimens from the Constantia and Newlands slopes of Table Mountain (east-facing subarea). In the present survey two substantial demes were located (Fig. 2). These were situated in the forest and scrub forest communities which dominate the east-facing subarea. Specimens were collected from under stones as well as from under the dried bark of Pinus pinaster and P. radiata stumps in areas which are being cleared of alien vege- tation (Fig. 4). NOTES ON THE SCORPION FAUNA OF THE CAPE py) hil Fig. 4. Typical habitat of Uroplectes insignis and U. lineatus in the forest scrub-forest community of the east-facing study subarea. Fig. 5. Run and burrow of Opisthophthalmus capensis. The antechamber is absent in this particular run. 236 ANNALS OF THE SOUTH AFRICAN MUSEUM No specimens of Parabuthus capensis were found in the present survey. There are several specimens in the collection of the South African Museum collected from Devil’s Peak (north-facing subarea) which lies in the dry proteoid community. SCORPION SHELTERS Opisthophthalmus capensis was always found to construct its burrow beneath some surface object. The excavation leading from the opening under the stone or log to the burrow entrance is the run. The structure of the run is dependent on the size and shape of the ground cover and nature of the substratum (Fig. 5). Thus the run varies in length and sometimes has two or even three openings. Just before the burrow entrance proper there is usually a bulge or separate section which could be called an ‘antechamber’ (Fig. 6B). Often food remains are found in this area and its function appears to be a feeding area, and possibly affords freer movement for the scorpion to turn or employ its sting. Excavation of some fifty burrows sometimes revealed food remains in the terminal chamber of the burrow itself. The burrow is usually a simple tunnel having a charac- A Canis) QrOUndlCOVeG.N v.ls 0. 2) Sia fans _—_—_—__——~___ = +> burrow entrance ruin entrance : : : : terminal chamber — direction of slope iS . see eae antechamber - - - °. \ | Fig. 6. A. Cross-section. B. Plan of typical run and burrow of Opisthophthalmus capensis. NOTES ON THE SCORPION FAUNA OF THE CAPE Di. teristically oval entrance penetrating the ground at about 30° from the hori- zontal (Fig. 6A). It often turns to right or left, if there are obstacles in the way, and ends in a very slightly enlarged terminal chamber. Variation is due to the nature of the substratum, size or sex of the inhabitant. Males construct shallower burrows or none at all, living only in a run. Only one specimen occupies each burrow. Ground cover, usually sandstone debris lying on soils of mixed origin, often sandy loam, under which burrows were found varied from 15 to 60 cm in diameter. If the stones lay on a slope the run entrances always faced downhill, ensuring dry runs during rainy weather. Table 2 gives the measurements of fifty burrows. TABLE 2 Measurements in centimetres of fifty burrows of Opisthophthalmus capensis (36 33, 14 99) Min. Max. Mean Run length g ‘ : : 3 ; 2 9,0 27,0 V2, 2 ; : : ; 10,0 22,0 15,3) Run width ¢ f ; : ; ; : 2,6 4,0 3) 2 Q : : ; : : : DS 3,4 3,1 Burrow entrance height ¢ . 4 : f 1,8 Bal 2,9 Q . ; : ; 2,0 3,6 By Il Burrow entrance width ¢. f : ; Dell 32 2,9 Ons : : : Dep 3,0 2,8 Burrow length ¢ . p : ; ; 0,0 20,0 10,6 Oe : ; 3 : ; 10,0 21,0 19,8 Vertical depth of burrows 5. : ; 355) 6,5 S77 Orie ; : 6,0 13,5 11,1 Uroplectes lineatus occupies a wide range of shelters which may be classified as follows:. 1. Beneath suitable stones—no evidence of any run construction, but where concavities form a natural shelter. Beneath stones where a run has been constructed. Beneath debris such as logs and dried organic matter. In rock crevices, or under stones lying on a flat rock surface. Beneath tree bark. A deme of U. lineatus in the south-facing subarea was found suitable for a study of run morphology. The structure of the run is dependent on the nature of both ground cover and substratum (Figs 7-8). The runs vary greatly in length and width (Table 3) and may have one or two entrances. U. lineatus in the east-facing subarea had the same shelters as U. insignis, namely the space between dried bark and stumps of pine trees (Fig. 9), although the two species were not found together under the same cover. Where sandstone he eS) TABLE 3 Measurements in centimetres of thirty runs of Uroplectes lineatus. Min. Max. Mean Length of run 5 : : : ; : 355 OES) 11,3 Width of run : 5 : ; 0,8 22 1,4 Ground cover size (diameter) Y , ; 9,0 55,0 31,0 238 ANNALS OF THE SOUTH AFRICAN MUSEUM \ | Fig. 7. Diagrams showing variation of runs or scrapes of Uroplectes lineatus. Fig. 8. Run of Uroplectes lineatus. Usually the runs are not as well defined as this one. NOTES ON THE SCORPION FAUNA OF THE CAPE 239 Fig. 9. A section of bark removed from a pine stump, showing a specimen of Uroplectes insignis, several isopods and a longicorn beetle. debris formed the ground cover, only one specimen of U. lineatus or U. insignis was found under each stone, whereas up to nine specimens of these species were collected from a single stump. One specimen of U. /ineatus was found in a shallow, wide burrow in the north-facing subarea and not covered by any surface object. Presumably the burrow was excavated by another animal. U. variegatus was found to occupy narrow runs, usually well defined with a small terminal chamber. Some specimens were also found in natural depres- sions beneath stones. Table 4 gives measurements of five runs of U. variegatus. TABLE 4 Measurements in centimetres of five runs of Uroplectes variegatus. Min. Max. Mean Length . “ ; ; s : : : 4,0 13,0 9.4 Width . : d , : : 4 ; 0,75 1,4 0,92 Ground cover size (diameter) . ; : 16,0 48,0 37,0 Parabuthus capensis was not collected during the survey, but has been collected by the author from other parts of the Cape. This species often excavates runs which vary according to the nature of ground cover and substratum and usually consist of a more or less round or oval space beneath the ground cover with one or two entrances. 240 ANNALS OF THE SOUTH AFRICAN MUSEUM COEXISTENCE The following pairs of species were found to coexist: Opisthophthalmus capensis— Uroplectes lineatus, O. capensis—U. variegatus, U. lineatus—U. insignis. In field studies demes or population aggregates were roughly delineated by markers to determine the spatial distribution of their shelters. Of eight demes of O. capensis, five showed a spatial overlap with demes of U. lineatus (Fig. 10) in studies carried out in the west-facing subarea. Specimens of U. variegatus were recorded in association with O. capensis in two of these demes where U. lineatus was absent. In one instance a specimen of U. variegatus had its run under the same rock as O. capensis, their respective entrances Opening on opposite sides. | 400m _ contour low scrub dry proteoid @ U.lineatus ©. O.capesis Fig. 10. Diagram of population aggregates of Opisthophthalmus capensis and Uroplectes lineatus in the west-facing study subarea, showing spatial overlap. In a study of coexistence of scorpions by Williams (1970) two mechanisms were found to permit coexistence: the choice of different-sized prey, and habitat specialization. The first mechanism undoubtedly operates in the case of O. capensis— U. lineatus and O. capensis—U. variegatus. U. insignis and U. lineatus were found under the bark of pine stumps in the same immediate vicinity, but specimens of either species did not share the same shelter. The mechanism permitting coexistence in this case is not quite ‘clear. The record of Parabuthus capensis from Devil’s Peak indicates the prob- ability of the coexistence of this species with U. lineatus. P. capensis from the Citrusdal district has been found in association with U. carinatus (Pocock) and O. pallidipes Koch. NOTES ON THE SCORPION FAUNA OF THE CAPE 241 SCORPION PREDATORS Because of the nocturnal habits of scorpions, predation is not often reported. The author has seen a troop of baboons on the northern slopes of Table Mountain foraging beneath stones. Baboons (Papio ursinus (Kerr)) are well-known predators of scorpions. The bokmakierie (Telophorus zeylonus (Linn.)) and the fiscal shrike (Lanius collaris Linn.) have been reported to prey on Uroplectes lineatus (1. G. Taylor pers. comm.). Owls are undoubtedly the commonest predators of scorpions. The spotted Eagle Owl (Bubo africanus (Tem.)), a local species, is reported to feed on scorpions (Roberts 1940). The Barn Owl (Tyto alba (Scopolt)) feeds on buthids and scorpionids (Vernon 1972). SOME OBSERVATIONS ON BEHAVIOUR The behaviour exhibited in burrow construction has been discussed by Williams (1966) for the Neotropical species Anuroctonus phaeodactylus (Wood) (Vejovidae), and Newlands (1972) reported on the burrowing behaviour of Protophthalmus holmi Lawrence, Opisthophthalmus concinnus Newlands and O. flavescens Purcell, all Ethiopian Scorpionidae. An important adaption of Opisthophthalmus capensis is its ability to con- struct burrows. In the laboratory specimens made burrows by loosening tightly- packed soil with a biting action of the chelicerae. The loose sand was then scraped out backwards by legs I and II which were tucked under the body. Legs III and IV provided the traction while the pedipalps were extended anteriorly and acted as supports to prevent the body tipping forward and occasionally as levers which aided in the backward movement. The cauda was extended straight out behind the body. The soil was transported rapidly backward in a single movement and the soil deposited outside the entrance on a tumulus by a slight flick of the first two pairs of legs so that the soil sprayed out. Occasionally the scorpion moved its load of soil by moving backward for a few centimetres, stopping, and thus pulling the soil backward with each step. With each step there was a pause while the hind legs were moved backward again to repeat the process. Thus two methods were employed to transport the soil: the first was used in exceptionally dry soil where backward movement was smooth and unimpeded; the second method was employed when the soil was damp and therefore more difficult to move, or if the burrow was steep or filled with small stones, twigs and similar obstacles. Several specimens did not burrow but made runs under stones provided for this purpose. Initially the animal would remove the soil in the usual manner as well as forming a broad chamber beneath the stone by pushing the soil with the tail. The curved tail was laid flat on the surface and then extended, thus moving a good volume of soil each time. This tail scraping is reported as being a widespread phenomenon among scorpions (Newlands 1972). It is also reported as a part of burrow maintenance in Anuroctonus phaeodactylus (Wood) by Williams (1966). Field observations showed that immature specimens were adept at burrow- 242 ANNALS OF THE SOUTH AFRICAN MUSEUM ing. Several second or third instar nymphs were found in burrows of 1-2 cm in length under stones. These burrows rarely had the characteristic oval opening and were often nothing more than shallow depressions. The tarsi of O. capensis are adopted for a pelophilous habitat, that is the claws are sharply curved and tarsal setae are not as dense as in O. wahlbergi (Thorell) for example, a psammophilous species. Caudal segment V is heavily sclerotized with numerous setae and well-developed keels to facilitate tail- scraping. Collections made in the field during the winter months (June—Sep- tember) indicate that these scorpions remained deep in their burrows on cold days, but if the sun was hot they were often found at the entrances of their burrows or in the runs, facing outward. This “doorkeeping’ is presumably maintained while the scorpion waits for prey which may seek refuge under stones (Alexander & Ewer 1958). The tail was curled around sideways, parallel to and resting on the substratum so that the telson, which was folded over the dorsal surface of caudal segment V, lay against the lateral side between the sixth and seventh mesosomal segments. From this position the metasoma could easily be brought into action sideways, as had been observed in the laboratory. This sideways movement is obviously more suitable for stinging prey while the scorpion is in its run. The pedipalps were arranged so that the femora were perpendicular to the body and the tibiae and chelae lay parallel to the body. The movable and immovable fingers were kept about 5 mm apart. If the scorpion was just inside the burrow entrance, the chelae may have protruded and were arranged as above. Sometimes the chelae were folded over each other with the dorsal and outer sections facing outwards, forming a sort of shield. This is a typical defensive stance for Scorpionidae (Newlands 1969). When a stone was lifted while a scorpion was in its run it would lift its tail so that the telson lay above the first caudal segment and the body was lifted off the substratum. If further aggravated it would adopt an aggressive stance so that the last two mesosomal segments were curved upward, the metosoma perpendicular to the ground except for the last caudal segment which was held horizontal with the telson above the last mesosomal segment or carapace. Another reaction was retreat, in which case the cauda was brought down backwards and the animal scuttled down its burrow, either backward or forward. The function of the doorkeeping seems to be related to feeding, where the scorpion waits for prey to come into the run. At no time was it observed that the scorpion was an active predator. However, active foraging in the vicinity of burrows is known for Opisthophthalmus (G. Newlands, pers. comm.). Feeding was observed in the field in the runs or antechambers. Feeding behaviour has been discussed for Neotropical scorpions by Baerg (1954) and Hadley & Williams (1968) (Buthidae and Vejovidae) and for the South African Opisthophthalmus latimanus by Alexander (1972). In the field O. capensis was observed feeding on Dorylinus helvolus (Formicidae) and Temnopteryx phalerata (Blattariae). In the laboratory a wide range of prey species were accepted and eaten, while others were ignored. Prey species which NOTES ON THE SCORPION FAUNA OF THE CAPE 243 were accepted were Periplanita americana, Temnopteri phalerata (Blatteriae), Dorylinus helvolus (Formicidae), Holopternia valga (Hemiptera) as well as U. lineatus, isopods, Scarabaeidae, beetle larvae, moths and moth larvae. The following species were either killed or ignored but not eaten: Chilopoda, Diplopoda and Anthia spp. (Carabidae). During excavation of burrows food remains in the form of pellets were sometimes found in the terminal chambers or in the runs. The food remains were identified as small tenebrionid beetles and cockroaches from the indigestible elytras and legs. Thus feeding occurred both in the burrows and the runs. Feeding behaviour was elicited in the laboratory by placing the prey near the scorpion which usually rests in the typical doorkeeping posture. Small prey was firmly grasped in both chelae and the metasoma brought upright. Small prey was not stung and the biting action of the chelicerae began immediately. If the prey was lively but not very large the scorpion brought the telson into play from the side and usually slowly, seeking for a soft spot in which to inject the venom. If the prey was large and aggressive the scorpion initiated offensive behaviour and the telson was brought swiftly over the carapace to deliver a series of stings. The ingestion process lasted from | to 3 hours in the laboratory, depending on the size of the prey. The formation of pellets of food remains was not observed. Ingestion started immediately after capture and in heavily sclerotized prey the starting point was ventrally between head and thorax. The chelicerae moved alternately in and out while the fingers grasped at the food, giving rise to the typical biting action. Sponge-bathing activities were not observed after feeding, nor after burrow- ing in wet soil when the legs and chelicerae became dirty. In humid laboratory environments O. capensis was never seen to approach the dish of water in the cage and it seems likely that water balance is maintained by contact with a wet substratum as reported by Hadley & Williams (1968) in observations at night of desert scorpions. If the cage was kept very dry and water placed in the cage the scorpions would climb over the edge of the dish and dip the chelicerae and pedipalps into the water. At the same time the pedi- palps moved rhythmically up and down. Uroplectes lineatus adopted a very characteristic resting posture while under ground cover. The pedipalps were usually held so that they lay slightly anterior to or lateral to and subparallel to the carapace. The tail was curved over the abdomen dorsally so that the second caudal segment was horizontal to the body and the terminal segments curved around sideways and the telson lay laterally between mesosomal segments V and VI, being folded over against caudal segment V dorsally. In the defensive posture the tail lay behind the body with the telson curved forward so that it lay above the first caudal segment. When agitated this scorpion can strike very rapidly to the side, forward or backward. This species did not use the chelae to grasp an aggressor but only struck out with the tail. In the aggressive posture there was a slight forward stilting. Venom-dripping was associated with aggressive behaviour. In the 244 ANNALS OF THE SOUTH AFRICAN MUSEUM laboratory it was noticed that U. lineatus was able to excavate a run, but the behaviour of digging was not observed. In the east-facing subarea specimens were found in association with Isopoda, geckos, longicorn beetles, Lagria villosa F. (Lagriidae), Pyrrhocoridae, Hemiptera, cockroaches (Temnopteryx phalerata (Sauss)) and U. insignis. In this environment both U. lineatus and U. insignis were observed feeding on cockroaches. Prey capture was not observed here or in the laboratory. Two pregnant female specimens of U. insignis and one of U. lineatus were kept in the laboratory. Parturition was observed in both species which behaved similarly. Parturition behaviour was as follows: the scorpion stilted on the two last pairs of legs prior to delivery; the first two pairs were held under the body so that the distal segments were parallel to the body. This posture was main- tained throughout parturition. The young began passing through the genital opening head first, one by one into the ‘birth basket’ formed by the first two pairs of legs. The young became active after a few minutes and began ascending the mother’s back by way of her walking legs. When all the young had reached the back they assumed a completely random orientation, sometimes in several layers, which continued to the first moult. This random orientation of first instar young is characteristic of the Buthidae (Williams 1969) (Figs 11-12). With the onset of the second instar stage the litter became distinctly more elongate. Duration of first instar stage of U. insignis was 9-12 days with litters of 12 and 13 specimens, and duration of first instar of U. lineatus was 11 days with a litter of 8 specimens. Parturition took place from January to February. KEY TO THE SCORPIONS OF TABLE MOUNTAIN 1. Chelae slender; the dorsal surface not separated by a keel (Family Buthidae). Found in simple runs or scrapes under stones or beneath (mee oe so a 2 — Chelae large, the dorsal curiite Renaitedl into inner anal er bee ione by a smooth keel (Family Scorpionidae). Found in burrows, usually opening under stones oe ie Opisthophthalmus capensis (Herbst) 2. Species large; stridulatory areas on dorsal surface of caudal segments I and II well developed. Posterior dorsal surface of caudal segment II raised to form a subtriangular lip Parabuthus capensis (Ehrenberg) Species small; stridulatory areas on cauda absent .. oe me 3 3. Tergites with short lateral keels and seven black longitudinal lines Uroplectes variegatus (Koch) — Tergites without lateral keels, darkly pigmented Nor iReoay its 4 4. Tubercle below aculeus distinctly laterally compressed; vesicle yellow a 5 Uroplectes lineatus (Koch) — Tubercle below neulene cones bal blunted; vesicle black Uroplectes insignis Pocock NOTES ON THE SCORPION FAUNA OF THE CAPE 245 Maly Fig. 11. First instar of Uroplectes insignis. Note the random orientation typical of the Buthidae. Fig. 12. Second instar of Uroplectes insignis. The young are beginning to leave the mother’s back. 246 ANNALS OF THE SOUTH AFRICAN MUSEUM CONCLUSIONS Four scorpions species were found on Table Mountain and Signal Hill. Two species, Opisthophthalmus capensis and Uroplectes lineatus, were very abundant whereas the other two were rare (U. insignis and U. variegatus). O. capensis was abundant on the western north-facing subarea, the greater proportion occurring in the dry proteoid plant community below the 400 m contour. Fewer specimens occurred in the low scrub communities since these areas are very steep and the substratum and ground cover unstable during the rainy season. U. lineatus was found to be fairly abundant in all subareas studied and has a random distribution, occurring in varied habitats. U. variegatus and U. lineatus were found respectively to coexist with O. capensis, although these two former species were not found to coexist. The fact that all the young pro- duced by two females of U. insignis had the characteristics of this species suggests that it is a true-breeding species which coexists with U. lineatus. Apparently isolating mechanisms have been acquired for this example of sympatric association, but what these are, is not quite clear. The notes on the behaviour make comparison with other species possible. In the burrowing behaviour of O. capensis two methods are employed to transport the soil; the first is similar to the behaviour shown by Protophthalmus holmi Lawrence (as reported by Newlands (1972)), i.e. the soil is transported rapidly backward in a single movement, and the second is like that of O. flaves- cens Purcell and O. concinnus Newlands, 1.e. the soil is transported backward by a series of jerks. Observations of laboratory specimens of O. capensis revealed that the first method was used in exceptionally dry soil where backward move- ment was smooth and unimpeded. The second method was employed when the soil was damp and therefore difficult to move, or if the burrow was steep or filled with small stones, twigs or similar obstacles. Thus O. capensis, which is pelophilous, exhibits burrowing behaviour similar to psammophilous scorpions in certain conditions. The excavation of a run by Parabuthus capensis was described by Eastwood (19775). Part of this process was similar to the behaviour shown by O. capensis when obstacles had to be removed, but loose sand was excavated in a manner characteristic to Parabuthus. The defensive behaviour of O. capensis is the same as the generalized behaviour for Opisthophthalmus as discussed by Newlands (1969). The resting posture of O. capensis prior to feeding differs from that of vejovid scorpions in the position of the metasoma. In O. capensis the metasoma is usually curved forward alongside the body whereas the vejovid scorpions discussed by Hadley & Williams (1968) held the metasoma straight out backwards. Feeding behaviour of O. capensis was similar to that observed for Parabuthus capensis, except that the latter was more reluctant to attack larger and more active prey. Birth activities of Neotropical Buthidae were discussed by Williams (1969) and this was found to be similar for U. insignis and U. lineatus. The first instar young of these two species showed the random orientation on the mother’s NOTES ON THE SCORPION FAUNA OF THE CAPE 247 back which was reported for the genus Centruroides (Williams 1969). Because of the position of Table Mountain, the human impact on its ecology is considerable. Frequent fires cause damage to the floral and faunal ecology. In this study it was observed that in recently burnt areas O. capensis was abun- dant, where U. lineatus was more abundant in adjacent unburned areas. It is obvious that O. capensis is able to withstand a veld fire because of its ability to burrow, whereas a local deme of Uroplectes can be exterminated. The lower slopes of the north-facing and east-facing subareas are covered in plantations of alien forest. In these areas there is very little insect life and consequently few scorpions were found. ACKNOWLEDGEMENTS I thank Dr V. Whitehead for advice and encouragement during this project, and the Director of the South African Museum for providing research and transport facilities. For constructive comments and advice on this and other projects I thank Mr B. Lamoral of the Natal Museum and Mr G. Newlands of the South African Institute for Medical Research. Thanks are also due to Dr V. Whitehead, Messrs A. Byron and S. Rice for help with the photographic work, and the Cape Town City Engineer’s Department and the State Department of Forestry for permits and permission to collect on Table Mountain. REFERENCES ALEXANDER, A. J. 1972. Feeding behaviour in Scorpions. S. Afr. J. Sci. 68: 253-256. ALEXANDER, A. J. & Ewer, D. W. 1958. Temperature adaptive behaviour in the scorpion Opisthophthalmus latimanus Koch. J. exp. Biol. 35: 349-359. BAERG, W. J. 1954. Regarding the biology of the common Jamaican scorpion. Ann. ent. Soc. Am. 47: 272-276. Eastwoop, E. B. 1977a. Notes on the scorpion fauna of the Cape. Part 1. Description of neotypes of Opisthophthalmus capensis (Herbst) and remarks on the O. capensis and O. granifrons Pocock species-groups (Arachnida, Scorpionida, Scorpionidae). Ann. S. Afr. Mus. 72: 211-226. Eastwoop, E. B. 1977b. Notes on the scorpion fauna of the Cape. Part 2. The Parabuthus capensis (Ehrenberg) species-group; remarks on taxonomy and bionomics (Arachnida, Scorpionida, Buthidae). Ann. S. Afr. Mus. 73: 199-214. HADLEY, N. F. & WILLIAMS, S. C. 1968. Surface activities of some North American scorpions in relation to feeding. Ecology 49(4): 726-734. Hewitt, J. 1918. A survey of the scorpion fauna of South Africa. Trans. R. Soc. S. Afr. 6: 89-192. KRAEPLIN, K. 1894. Revision der Skorpione. Jb. hamb. wiss. Anst. 12: 75-96. KRAEPLIN, K. 1899. Scorpiones und Pedipalpi. Das Tierreich 8: 1—265. KRAEPLIN, K. 1908. Skorpione und Solifugen. Ergebnisse e. Forsch. Siidafrika Bd 1, Lief 2. Denksch. med.-naturw. Ges. Jena 13: 247-282. LAWRENCE, R. F. 1955. Solifugae, Scorpions and Pedipalpi, with checklists and keys to South African families, genera and species. S. Afr. anim. Life 1: 152-262. Mo Li, E. J. & CAMPBELL, B. M. 1976. Table Mountain. A conservation and management report. Cape Town: University of Cape Town. NEWLANDS, G. 1969. Scorpion defensive behaviour. Afr. wild Life 23: 147-153. NEWLANDS, G. 1972. Notes on psammophilous scorpions and a description of a new species (Arachnida: Scorpionides). Ann. Transv. Mus. 27: 241-254. 248 ANNALS OF THE SOUTH AFRICAN MUSEUM PeNTHER, A. 1900. Zur kenntnis der Arachnidenfauna Sitidafrikas (Scorpiones). Annln. naturh. Mus. Wien 15: 153-163. Pocock, R. I. 1890. A revision of the genera of scorpions of the family Buthidae, with descrip- tions of some South African species. Proc. zool. Soc. Lond. 1890: 114-141. Pocock, R. I. 1896. A further revision of the species of scorpions belonging to the South African genera Uroplectes, Lepreus and Tityolepreus. Ann. Mag. nat. Hist. (6) 17: 377-393. PURCELL, W. F. 1899. On the species of Opisthophthalmus in the collection of the South African Museum, with descriptions of some new forms. Ann. S. Afr. Mus. 1: 131-180. PuRCELL, W. F. 1901. On some South African Arachnida belonging to the orders Scorpiones, Pedipalpi and Solifugae. Ann. S. Afr. Mus. 2: 137-225. Roserts, A. 1940. Birds of South Africa. Revised by McLachlan, G. R. & Liversidge, R. 3rd ed. 1970. The Trustees of the John Voelcker Bird Book Fund. VERNON, C. J. 1972. An analysis of owl pellets collected in southern Africa. Ostrich 43: 109-124. WILLIAMS, S. C. 1966. Burrowing activities of the scorpion Anuroctonus phaeodactylus (Wood) (Scorpionida: Vejovidae). Proc. Calif. Acad. Sci. 34: 419-428. WILLIAMS, S. C. 1969. Birth activities of some North American scorpions. Proc. Calif. Acad. Sci. 37: 1-24. WILLIAMS, S. C. 1970. Coexistence of desert scorpions by differential habitat preference. Pan-Pacif. Ent. 46: 254-267. 6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature (particularly Articles 22 and 51). Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov., etc. An author’s name when cited must follow the name of the taxon without intervening punctuation and not be abbreviated; if the year is added, a comma must separate author’s name and year. The author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is transferred from its original genus. 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Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51. THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. (continued inside back cover) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 74 +#&Band January 1978 Januarie Part 11 Deel Yia,, S AU /0Ui0. NONI spas NOTES ON THE SCORPION FAUNA OF THE CAPE PART 4 THE BURROWING ACTIVITIES OF SOME SCORPIONIDS AND BUTHIDS (ARACHNIDA, SCORPIONIDA) By E. B. EASTWOOD Cape Town Kaapstad The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular intervals as material becomes available Obtainable from the South African Museum, P.O. Box 61, Cape Town Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM word uitgegee in dele op ongereelde tye na beskikbaarheid van stof Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad OUT OF PRINT/UIT DRUK 1, 2(1-3, 5-8), 3(1-2, 4-5, 8, t.-p.i.), 5(1-3, 5, 7-9), 6(1, t.—p.i.), 711-4), 8, 91-2, 7), 101-3), 11(1-2, 5, 7, t-—p.i.), 15(4-5), 24(2), 27, 311-3), 32(5), 33 Price of this part/Prys van hierdie deel R1,70 Trustees of the South African Museum © Trustees van die Suid-Afrikaanse Museum 1978 ISBN 0 908407 32 7 Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk., Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap NOTES ON THE SCORPION FAUNA OF THE CAPE PART 4 THE BURROWING ACTIVITIES OF SOME SCORPIONIDS AND BUTHIDS (ARACHNIDA, SCORPIONIDA) By E. B. EAsTwoop South African Museum, Cape Town (With | figure and 2 tables) [MS. accepted 23 August 1977] ABSTRACT Burrowing activities of the scorpionid Opisthophthalmus macer Thorell were studied in the field and laboratory, and comparisons made with O. capensis (Herbst). O. karrooensis Purcell, O. pallidipes Koch and O. peringueyi Purcell were found to burrow only in their immature stages. The morphological adaptions of the adults are discussed in relation to their scrape-dwelling habit. The specialized digging behaviour of the buthid Parabuthus planicauda (Pocock) is described. Pregnant females were often found to construct proper burrows prior to parturition. CONTENTS PAGE IiEKOGUCHIONS 9 oe ee we a bat ie Pte ae, Pale DAD DISCUSSION eee eee css sae A OS oh el OO GOnclusiGns) ten) we oe eee ah agar a 25S ACknOWwledSementS es gu.) sacs.) te a el 254 IRNGLGKENCES Sr eae oe et ee a ee 4 INTRODUCTION Opisthophthalmus macer Thorell is a true burrowing scorpionid, distributed from False Bay to Port Elizabeth along the south coast. In this paper the burrow morphology and burrowing behaviour of this scorpion are reported and com- pared to the corresponding activities of O. capensis (Herbst). O. karrooensis Purcell, O. pallidipes Koch and O. peringueyi Purcell all occupy the drier semi-desert regions of the Cape, namely the Karoo and Namaqualand southwards to Piketberg. The young of these species are true burrowers, whereas the adults have in common a scrape-dwelling habit with special morphological adaptions for this particular way of life. Digging behaviour has been reported in the Buthidae and shows certain similarities to Parabuthus planicauda (Pocock) and P. capensis (Ehrenberg). This was illustrated by Pavlovsky (1924) for the transcaspian species Liobuthus kessleri Birula, where the first three pairs of walking legs were used for excava- tion. Lankester (1882) reported on the digging behaviour of the Egyptian species Buthus australis citrina (Hemprich & Ehrenberg). In this case the 249 Ann. S. Afr. Mus. 74 (11), 1978: 249-255, 1 fig., 2 tables. 250 ANNALS OF THE SOUTH AFRICAN MUSEUM pedipalps aided the first three pairs of legs in digging operations. P. planicauda is a widely distributed buthid of the southern Cape, extending from Worcester and Tulbagh in the west to Albany and Cathcart in the east (Lawrence 1955). Under certain conditions this species may construct true burrows. The specialized digging behaviour of this species is discussed in relation to the activities of P. capensis (Ehrenberg). DISCUSSION Opisthophthalmus macer Thorell Two demes of this species were studied. The first was found on the southern slopes of the Kleinriviersberge in a habitat consisting of macchia, namely Proteaceae, Restionaceae, Ericaceae, Geraniaceae and other families. Available ground cover was sandstone and quartzite debris and the substratum was deep, sandy loam rich in organic matter. O. macer was found to burrow only beneath suitable ground cover. Typically a run was constructed with a single entrance, leading to the burrow opening. Often an antechamber was present, the probable function of this being a feeding area. The burrows of females were invariably longer than those of males and often males were found to construct only a run. Table | gives the measurements of twenty burrows and runs. The second deme was located next to the sea in the southern part of Walker Bay. This habitat consisted of rocky limestone slopes with very little soil and low coastal scrub. In this habitat the burrows were generally shorter and none were found with an antechamber in the run. Specimens were also more darkly coloured. Table 1 gives the measurements of ten burrows and runs of the second deme. Burrowing behaviour of O. macer was observed as follows: compacted soil was loosened by the action of the chelicerae and scraped into a heap by the first two pairs of walking legs. The heap was then moved backward by legs I and IJ, which were tucked beneath the body. Legs III and IV provided the traction while the tail was extended, the pedipalps being used as supports. The tarsal claws of O. macer are more sharply curved than those of O. capensis. According to Newlands (1972) this is related to a pelophilous habit. This view could not be substantiated in the present study, since the Kleinriviersberge deme was located in sandy loam and the deme from the coastal habitat was found to burrow in compacted sand, whereas O. capensis from the Cape Peninsula burrowed in sandy loam only. Tail scraping was not observed although the heavily sclerotized fifth caudal segment of O. macer is adapted for this function. The scrape-dwellers, Opisthophthalmus karrooensis Purcell, O. pallidipes Koch and O. peringueyi Purcell Observations of a deme of O. karrooensis showed that although the adults were found to occupy scrapes or runs, or simply depressions beneath stones, NOTES ON THE SCORPION FAUNA OF THE CAPE 251 TABLE | Measurements in centimetres of twenty burrows and runs of Opisthophthalmus macer in a habitat of Cape macchia (Deme 1), and ten burrows in a limestone and coastal scrub habitat (Deme 2). Deme | Deme 2 Min. Max. Mean Min. Max. Mean Run length 8,0 19,0 16,5 6,0 10,0 7,0 3,0 14,0 10,2 Ds) 7,0 5,0 Run width DS) 3) 3,0 Dp 3,0 2,6 Burrow entrance height . Burrow entrance width . Burrow length Vertical depth of burrow +O Os +O Oy +O Os =} OOS tO MON +0 Oy ww i) ww ey we N _ the immature stages constructed burrows characteristic of other groups in this genus. These burrows opened beneath ground cover and the length and breadth varied with the stage of development. The burrows were often simple, straight tunnels varying from 2 to 9 cm in second and third instars to 6 to 17 cm in subadults. The scrape-dwelling habit of adults and burrowing by the young was also evident in populations of O. pallidipes and O. peringueyi from the Clanwilliam district. Table 2 shows the relationship between the mode of shelter and certain morphological characters for ten species of Opisthophthalmus. The pedipalps of these three species are powerful and heavily sclerotized. The chelae of the males in particular are very long and afford frontal and lateral shielding of the opisthosoma (Fig. 1). The mesosoma is broad and distinctly dorsoventrally compressed, and the metasoma slightly more laterally compressed than those species which burrow in all stages. The tail is adapted for tail-scraping operations by being heavily sclerotized. Parabuthus planicauda (Pocock) While collecting this species in a variety of habitats over a very large area it was noticed that pregnant females were usually found in deep, narrow ANNALS OF THE SOUTH AFRICAN MUSEUM Aya MOJING SUNOA pure s}[npe MOJING BUNOA SIdTJOMp-odesos sjjnpe MOJING SUNOA SIO[JOMp-odesOs s}[Npe MOLING SUNOA pue s}[npe ejep ou MOLING SUNOA pue s}[Npe MOLING SUNOA SIO[[JOMp-advss s}[npe © UL poyesuojS Jou © UL poyesuoje £ UI poyesuOyS © UI poyesuO]S JOU £ UI poyesuOys © UI poyesUOCTS JOU © UL poyesuoys possoiduros Aj[e19}e] 10U possorduros Ajjesoyey ApUSITs possorduros Ajjesoyey ATVYSITS posserdu0s Aj[es9yey] JOU _ -passorduro05 Ajjesoyey ATVUSITS possoi1du05 Aj[e1oze] OU possoiduros Ajjesoyey ATWYSITS possoiduros Ajje1yUsAOSIOp A[SUOIS JOU possoidur0s A][e1JUSAOSIOP A[BUOIIS possoiduros A][eIQUSAOSIOP A][SUOI}S posserduo05 A][e1JUSAOSIOP A[SUOI}S JOU possorduros A][e1JUsAOSIOp A[SUOIIS posso1dur0s A][eIJUSAOSIOP A[SUOIS JOU possoiduro0s Aj[eUsAOSIOp A[SUOIS 7 (S10Y.L) 184aq] YOM °C [Jeong 1dansuisad *—C * YoY sadipyjod °C ][210U,L, 42a *—C y]s91nd ppnvI1suo] °C * YOOY snuvuiijv] ‘CO * yjeoIng sisuaoodivy °C MOIING SUNOA pue sj}[Npe £ UL poyesuOja JOU possoiduros Ajje19}e] JOU possorduros A][eIJUSAOSIOP A[SUOI}S JOU yoos0g suosfiudss °C SUNOA JO} B}yep OU SIO][OMp-odeIOs s}[nNpe £ UI poyesuoOyS possordwo0s Ajjesoyey ATWYSI[S possoiduro0s A][VIJUSAOSIOP A[SUOI}S ][901Ndg svsis °C MOIING SUNOA puke s}[Npe © UI po}esuO]s JOU possorduio5 Ajyes9ye] OU possorduro05 A][e1yUSAOSIOP A[SUOI}S JOU e (iSqioH) sisuadvd °C J9}[94S ee) BUWOSPIOIAL PWIOSOSII, PUL LUIOSOIg so1seds “I9}JOYS JO spou pue sIa}ORILYO [BOISOTOYdIoUL UIe}ID Ud9MjOq AIYsUONeIOI 9Y} SuNeoIpur sniupoysydoyssid—E Jo sa1oadg Z AIGVL NOTES ON THE SCORPION FAUNA OF THE CAPE 253 Fig. 1. Pedipalp and prosoma of male specimens of A. Opisthophthalmus karrooensis and B. O. macer. depressions or properly constructed burrows, opening beneath stones. The entrances were oval or round and the burrows varied from 4 to 11 cm in length. Males, non-pregnant females, and young were found in scrapes of varying shapes and lengths. Often specimens occupied natural depressions under stones, where there was no evidence of excavation. Laboratory specimens of pregnant females were observed scraping sand as follows: the tail was curved either sideways or directly over the mesosoma so that the telson lay above the carapace, and the first two pairs of walking legs were used in alternative scraping motions to loosen the sand. The body was then raised by stilting on the hind legs while the first two pairs of legs and sometimes the third pair scraped the sand very vigorously so that it sprayed out behind the scorpion; the pedipalps and fourth pair of legs were used as supports during this operation. Transport of soil from a burrow was not observed. The defensive armature of Parabuthus consists of the well-developed meta- soma and highly toxic venom. It was observed that the terminal chambers of the burrows were wide enough to enable the sting to be used effectively, and the deep narrow depressions were constructed so as to allow the sting to be employed without hindrance. CONCLUSIONS When the burrowing behaviour of O. macer was compared to O. capensis only one difference was noted, namely the scraping of soil into a heap by the first two pairs of legs prior to removal from the burrow in the former species. Eastwood (1978) noted that O. capensis modified its behaviour under different 254 ANNALS OF THE SOUTH AFRICAN MUSEUM substratum conditions. For example, when the soil was dry the scorpion transported it backwards out of the burrow in a single rapid movement, whereas if it was damp or there were stones or other obstacles in the soil, it was removed in a series of backward jerks. Thus it appears that the pattern of behaviour is flexible to accommodate various ecological conditions. It may be that the scraping of soil into a heap prior to transportation is used by O. capensis, but was not observed in this species because the conditions of the soil did not require it. However, this observation illustrates that more detailed information is required to determine significant specific behaviour patterns. The measurements of burrows and runs for the demes occupying two different habitats indicate that burrow and run length are a function of the nature of the habitat, i.e. the Kleinriviersberge specimens had deeper burrows since the substratum consisted of deep, sandy loam, whereas in the coastal habitat of the second deme, specimens constructed shallower burrows in pockets of compacted sand on a limestone outcrop. Generally O. macer and O. capensis show similar trends in burrow construction and behaviour, and more in-depth studies would be necessary to determine significant differences. The scrape-dwelling species of Opisthophthalmus are adapted to their habitat by the development of a flattening of the body and the increased effective- ness of defensive and aggressive armature. That the young of these forms are burrowers indicates that they may have evolved from the true-burrowers. The flattening of the opisthosoma and metasoma and elongation of the chelae are also characteristic of the ischnurinine genus Hadogenes, a lithophilous form, and is a good example of parallel evolution. The burrowing habit of P. planicauda appears to occur only in females prior to parturition. The sand-scraping behaviour of this species is essentially the same as that reported for P. capensis by Eastwood (1977). Removal of soil in large quantities by P. capensis was not observed in P. planicauda. However, this process is obviously necessary for burrow excavation, since the confined space at the entrance would not allow for the posture required for sand-scraping operations. No doubt many other Parabuthus species will be found to burrow and a great deal needs to be studied with respect to functional morphology, burrowing behaviour and the relationship to the type of habitat. ACKNOWLEDGEMENTS I thank Dr V. Whitehead for his advice and help, and the Director of the South African Museum for providing research and transport facilities. REFERENCES EAstwoop, E. B. 1977. Notes on the scorpion fauna of the Cape. Part 2. The Parabuthus capensis (Ehrenberg) species-group; remarks on taxonomy and bionomics (Arachnida, Scorpionida, Buthidae). Ann. S. Afr. Mus. 73: 199-214. NOTES ON THE SCORPION FAUNA OF THE CAPE 255 Eastwoop, E. B. 1978. Notes on the scorpion fauna of the Cape. Part 3. Some observations on the distribution and biology of scorpions on Table Mountain. Ann. S. Afr. Mus. 74: 229-248. LANKESTER, E. R. 1882. Observations on scorpions. Proc. R. Soc. 33: 95-104. LAWRENCE, R. F. 1955. Solifugae, Scorpions and Pedipalpi, with checklists and keys to South African families, genera and species. S. Afr. anim. Life 1: 152-262. NEWLANDS, G. 1972. Ecological adaptions of Kruger Park scorpionids (Arachnida: Scorpionides). Koedoe 15: 37-48. PAVLOVSKY, E. 1924. Zur Ausseren Morphologie der Scorpione. Ezheg. zool. Mus. 25: 125-141. 6. SYSTEMATIC papers must conform to the International code of zoological nomenclature (particularly Articles 22 and 51). Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov., etc. An author’s name when cited must follow the name of the taxon without intervening punctuation and not be abbreviated; if the year is added, a comma must separate author’s name and year. The author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is transferred from its original genus. The name of a subsequent user of a scientific name must be separated from the scientific name by a colon. Synonymy arrangement should be according to chronology of names, i.e. all published scientific names by which the species previously has been designated are listed in chronological order, with all references to that name following in chronological order, e.g.: Family Nuculanidae Nuculana (Lembulus) bicuspidata (Gould, 1845) Figs 14-15A Nucula (Leda) bicuspidata Gould, 1845: 37. Leda plicifera A. Adams, 1856: 50. Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b). Nucula largillierti Philippi, 1861: 87. Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. 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REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: ‘Smith (1969) describes...’ ‘Smith (1969: 36, fig. 16) describes...’ ‘As described (Smith 1969a, 1969b; Jones 1971)’ ‘As described (Haughton & Broom 1927)...’ ‘As described (Haughton et al. 1927)...’ Note: no comma separating name and year Dagination indicated by colon, not p. names of joint authors connected by ampersand et al. in text for more than two joint authors, but names of all authors given in list of references. (b) Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, b, etc. to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a). For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list o, scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number (only if independently paged) in parentheses, pagination (first and last pages of article). Examples (note capitalization and punctuation) BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. FIscHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140. FiscHer, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. gén. 74: 627-634. Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320. Koun, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51. THELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. (continued inside back cover) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 74 Band February 1978 Februarie Part 12 Deel CRETACEOUS FAUNAS FROM ZULULAND AND NATAL, SOUTH AFRICA THE AMMONITE FAMILY LYTOCERATIDAE NEUMAYR, 1875 By WILLIAM JAMES KENNEDY & HERBERT CHRISTIAN KLINGER Cape Town Kaapstad The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular intervals as material becomes available Obtainable from the South African Museum, P.O. Box 61, Cape Town Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM word uitgegee in dele op ongereelde tye na beskikbaarheid van stof Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad OUT OF PRINT/UIT DRUK 1, 2(1-3, 5-8), 3(1-2, 4-5, 8, t.-p.i.), 51-3, 5, 7-9), 6(1, t.—p.i.), 711-4), 8, 9(1-2, 7), 10(1-3), 11(1-2, 5, 7, t.—p.i.), 15(4—5), 24(2), 27, 31(1-3), 32(5), 33 Price of this part/Prys van hierdie deel R6,10 Trustees of the South African Museum © Trustees van die Suid-Afrikaanse Museum 1978 ISBN 0 908407 36 X Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk., Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap CRETACEOUS FAUNAS FROM ZULULAND AND NATAL, SOUTH AFRICA THE AMMONITE FAMILY LYTOCERATIDAE NEUMAYR, 1875 By WILLIAM JAMES KENNEDY Geological Collections, University Museum, Oxford & HERBERT CHRISTIAN KLINGER South African Museum, Cape Town (With 53 figures) [MS. accepted I September 1977] ABSTRACT Ammonites referred to the family Lytoceratidae Neumayr, 1875, occur in rocks of Barremian, Aptian and Albian age in northern Zululand, South Africa, and represent the most southerly Cretaceous records of all the species present. The following are described: Lytoceras vogdti Karakasch, which is common in the Aptian, Lytoceras aff. sauclum (Drushchitza), represented by a single specimen from the Aptian, and Lytoceras hennigi Zwierzycki, which is frequent in the Upper Barremian and was previously known only from Tanzania. Eulytoceras is represented in the Upper Barremian by the widely occurring E. phestum (Matheron); A. (Ammonoceratites) mahadeva (Stoliczka) occurs in the Middle and Upper Albian, and A. (Ammonoceratites) crenulatum (Crick), A. (A.) ezoense (Yabe) and A. (A.) crenocostatum (Whiteaves) are shown to be synonyms of Stoliczka’s species. A. (Argonauticeras) depereti (Kilian), of which A. (A.) argonautarum (Anderson) and A. (A.) belliseptatum (Anthula) are considered synonyms, occurs in the Upper Aptian; a form referred to as A. (A.) aff. depereti is present in the Albian, accompanying A. (A.) besairiei Collignon. Protetragonites is represented in the Upper Albian by the widely occurring P. aeolus aeolus (d’Orbigny). The genus Pictetia is regarded as only a doubtful lytoceratid, perhaps better classed with the Ancyloceratina; it is represented by a specimen referred to as Pictetia aff. depressa (Pictet & Campiche). A number of important specimens, including the types of A. (Ammonoceratites) ezoense, A. (Am.) crenulatum and A. (Argonauticeras) belliseptatum, are figured photographi- cally for the first time. CONTENTS PAGE MOG HOME ye cn gee eee ah a Cian LOD WoOcatiOnuOlspecIMeNSie ees eae ees 258 ic Gulocalitiesi 4 ram mebers oat eek Cs sa OO ID MBG OCs O Gees 94958 9s 2c =e oe Zab SutunentenminOlOeyseucny! auto ap, anes ak hee LOD Systematic palaeontology SELLY etna ty nes SEMEL A fy) Ncknowledgementstaer, see we. eee! Sew | = 9330 INGECKeNCeSentek >, eee tet OR an ere hee bane e. Y S3O INTRODUCTION The Lytoceratidae are represented by nine species in the South African Cretaceous, all referrable to the subfamily Lytoceratinae Neumayr, 1875, of AST Ann. S. Afr. Mus. 74 (12), 1978: 257-333, 53 figs. 258 ANNALS OF THE SOUTH AFRICAN MUSEUM which the Hemilytoceratinae Spath, 1927, and Protetragonitidae Spath, 1927, are regarded as inseparable parts. The Lytoceratinae are the rootstock of the Lytoceratina, ranging from Triassic to mid-Cretaceous. They are a morphologically conservative group, with long and widely ranging genera and species. During the late Jurassic and Cretaceous the group were commonest in the Mesogean Realm, although known from as far north as Greenland and as far south as the sub-Antarctic Islands, and typically rare in the Boreal Realm of western Europe and the United States Western Interior. The following species are described below: Lytoceras vogdti Karakasch, 1907 Lytoceras aff. sauclum (Drushchitza, 1956) Lytoceras hennigi Zwierzycki, 1914 Eulytoceras phestum (Matheron, 1878) Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1865) Ammonoceratites (Argonauticeras) depereti (Kilian, 1892) Ammonoceratites (Argonauticeras) aft. depereti (Kilian, 1892) Ammonoceratites (Argonauticeras) besairiei Collignon, 1949 Protetragonites aeolus aeolus d’Orbigny, 1850 Pictetia aff. depressa (Pictet & Campiche, 1861) LOCATION OF SPECIMENS The following abbreviations are used to indicate the repositories of the materials studied: BMNH British Museum (Natural History), London IGS Geological Museum, London MNHP Muséum d’Histoire Naturelle, Paris SAM South African Museum, Cape Town SAS South African Geological Survey, Pretoria GSC Geological Survey of Canada, Ottawa UPG University of Pretoria, Geology department FIELD LOCALITIES Details of field localities referred to in the paper are given in Kennedy & Klinger (1975). DIMENSIONS OF SPECIMENS Dimensions of specimens are given in millimetres; abbreviations are as follows: D = diameter, Wb = whorl breadth, Wh = whorl height, U = umbilical diameter. Figures in parentheses are dimensions as a percentage of total diameter. CRETACEOUS FAUNAS FROM SOUTH AFRICA 259 SUTURE TERMINOLOGY The suture terminology of Wedekind (1916; see Kullman & Wiedmann 1970 for a recent review) is followed in the present work: I; = Internal lobe with septal lobe, U = Umbilical lobe, L = Lateral lobe, E = External lobe. SYSTEMATIC PALAEONTOLOGY Phylum MOLLUSCA Class CEPHALOPODA Cuvier, 1797 Subclass AMMONOIDEA Zittel, 1884 Order LYTOCERATIDA Hyatt, 1899 Superfamily LYTOCERATACEAE Neumayr, 1875 Family LYTOCERATIDAE Neumayr, 1875 Subfamily Lytoceratinae Neumayr, 1875 Genus Lytoceras Suess, 1865 Types species Ammonites fimbriatus J. Sowerby, 1817 (1.C.Z.N. Opinion 130). Diagnosis Evolute, serpenticone, whorls slowly expanding, with a rounded to quadrate whorl section. The surface of the shell is ornamented by fine crenulate ribs or growth lines, and there are periodic flares, each associated with a constriction on the internal mould. The shell surface may be finely striate. Suture formula ELU, (Uyy : Uya) Is; highly subdivided, with a short external lobe (E) and a triangular, highly subdivided lateral (L) lobe. There is a large septal lobe. Discussion Lytoceras is a conservative genus, ranging through the Jurassic and Lower Cretaceous with very little change. There have been repeated attempts to sub- divide the genus, but the authors follow Arkell & Wright (1957), Wiedmann & Dieni (1968) and others in regarding Ophiceras Suess, 1865, Fimbrilytoceras Buckman, 1918, Thysanoceras Hyatt, 1867, Thysanolytoceras Buckman, 1905, Kallilytoceras Buckman, 1921, Crenilytoceras Buckman, 1926, Orcholytoceras Buckman, 1926, Biasaloceras Drushchitza, 1953, Valentolytoceras Beznosov, 1958, and Dinolytoceras Beznosov, 1958, as synonyms. Hemilytoceras Spath, 1927 (type species Ammonites immanis Oppel, 1865), has rounded and smooth inner whorls, and on the outer whorl develops high, concave, closely spaced lamellar flares. Metalytoceras Spath, 1927 (type species Lytoceras triboleti Hohenegger in Uhlig 1883), is a compressed, finely ribbed genus, with very weak constrictions, and is readily separable from Lytoceras by virtue of the consistent splitting of the ribs into fine riblets over the venter. Pterolytoceras Spath, 1927 (type species Ammonites exoticus Oppel, 1865), is very loosely coiled, with slowly expanding subcircular whorls; the ornament is 260 ANNALS OF THE SOUTH AFRICAN MUSEUM much finer than in Lytoceras, and rather irregular. Eulytoceras Spath, 1927 (type species Ammonites inaequalicostatus d’Orbigny, 1840), differs from Lytoceras in having regular, distant ribs and periodic flares, with fine striae or riblets between. In the type species, this ornament survives in adults, as it does in E. phestum (Matheron). There are, however, species such as ‘Biasaloceras’ sauclum Drushchitza which have Eulytoceras-like inner whorls, but develop a Lytoceras-like adult ornament of fine, crinkled ribs and flares with associated constrictions. These forms are both classed with Lytoceras sensu stricto here, but they point to the homogeneity of the Lytoceratinae and the problem of placing many ‘intermediate’ species within the group. The remaining Cretaceous Lytoceratinae are readily distinguished from Lytoceras; in Pictetia Uhlig, 1883 (type species Crioceras astierianum d’Orbigny, 1842), the whorls are not in contact, whilst there are no flares or constrictions. These are also generally absent in the rapidly expanding, finely ribbed Ammono- ceratites (Ammonoceratites) and A. (Argonauticeras), although, as is noted later, the dividing line between many lytoceratid genera discussed here is sometimes exceedingly thin. Occurrence Lytoceras ranges from the Lower Jurassic (Pliensbachian) to the Cenomanian, with a world-wide distribution extending as far north as Japan, Greenland and Alaska, and as far south as Zululand and the sub-Antarctic Islands. In the Cretaceous, the group is unknown in the Boreal region of northern Europe and the Western Interior of North America. Lytoceras vogdti Karakasch, 1907 Figs 1-8, 9A, 12A, F-G Lytoceras vogdti Karakasch, 1907: 51, pl. 5 (fig. 1), pl. 24 (fig. 31), pl 26 (fig. 7). Forster, 1975: 143, pl. 1 (fig. 3). Pictetia vogdti Drushchitza, 1956: 82, pl. 5 (fig. 19), text-fig. 36a-d. Drushchitza & Kudryavtseva, 1960: 257, pl. 6 (fig. la—b), text-fig. 65. ‘Pictetia’ vogdti Schindewolf, 1961: 679. Type The larger of the two specimens figured by Karakasch (1907, pl. 5 (fig. 1)) is herein designated lectotype of this species; it is from the Barremian of the Crimea. Material Large lytoceratids referred to this species are common in the Upper Aptian of Zululand, especially along Mlambongwenya Spruit and in the Mkuze Game Reserve; SAS L/Li, Z1113-4, Z805, 805a and A1158, are all from the Makatini Formation, Aptian IV, at locality 151 in the Mkuze Game Reserve. SAS Z7 (Haughton Collection) is from the Makatini Formation at Haughton’s (1936) CRETACEOUS FAUNAS FROM SOUTH AFRICA 261 Fig. 1. Lytoceras vogdti Karakasch, 1907. SAS Z1731. x 0,38. locality Z7. UPG B391-393 and BMNH C78716 are from the Makatini Formation at locality 171; BMNH C78712, C78715, and C78717 are from the same formation, Aptian IV, at locality 172, Mlambongwenya Spruit. SAS LJE71, BMNH C78702, and possibly C78710 are from the Makatini Formation, Aptian III-IV, at locality 168, Mfongozi Spruit; and BMNH C78703 is from the same horizon at locality 34 on a tributary of the Mzinene River near Hluhluwe, Zululand. 262 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 2. Lytoceras vogdti Karakasch, 1907. SAS Z1731. x 0,38. CRETACEOUS FAUNAS FROM SOUTH AFRICA 263 Fig. 3. Lytoceras vogdti Karakasch, 1907. SAS LJE 71. x 0,38. 264 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 4. Lytoceras vogdti Karakasch, 1907. SAS LJE 71. x 0,38. CRETACEOUS FAUNAS FROM SOUTH AFRICA 265 Fig. 5. Lytoceras vogdti Karakasch, 1907. A. SAS Z1731, showing septal lobe. xX 0,45. B. SAS 2805, showing relationship between septal lobe and cruciform internal lobe. x 1,3. 266 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 6. External suture of Lytoceras vogdti Karakasch, 1907. x 4,5. 267 CRETACEOUS FAUNAS FROM SOUTH AFRICA T x “€0L8L0 HNWG L061 ¢ YOseyese yy Updo sv DIOJAT 268 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 8. Lytoceras vogdti Karakasch, 1907. Copy of the original figures of the lectotype. x 1. Dimensions Lectotype (after Karakasch) SAS L/Li SAS Z805 SAS Z805a UPG B393 LJE 71 SAS Z113 UPG B391 SAS Z1114 SAS A1158 Wb 34,0(40) 67,5(44) 82,0(—) 16,0(—) 95,0(45) 130,0(46) 149,0(45) 165,0(47) 198,0(49) 20) Wh 32,0(38) 67,0(43) TOS 16,0(—) 94,0(45) 135,0(48) 142,0(43) 161,0(46) 170,0(42) 200,0(—) Wb: Wh 1,06 1,00 1,01 1,00 1,00 0,96 1,05 1,02 1,16 1,05 U 30,0(35) 55,0(35) =) = (G3) 80,0(28) GS) 115,0(33) 120,0(30) —O CRETACEOUS FAUNAS FROM SOUTH AFRICA 269 Description This is a very large lytoceratid; the largest fragment is still septate at a whorl breadth of 211 mm, corresponding to an estimated total diameter of close on 450 mm for the phragmocone alone. Almost, all specimens have very poorly preserved inner whorls, commonly encrusted with oysters, and it has proved impossible to dissect out the nuclei of the giant specimens to confirm that they indeed correspond to the few associated juveniles, the best preserved of which is illustrated as Figures 7A-C, 12A. This individual is partially exfoliated and does not preserve the outer shell surface; it shows, however, very evolute coiling with a shallow dorsal impressed zone, a wide umbilicus and rapidly expanding depressed whorls (Fig. 7A). The exfoliated ornament (Fig. 7B-C) consists of prominent, narrow, fairly distant ribs separated by from three to four fine growth striae. Both ribs and growth striae arise at the umbilical seam; they are concave on the umbilical wall, sweep forwards across the shoulder and are markedly prorsiradiate on the flank, crossing the venter with a broad shallow convexity (Fig. 7B). The specimen bears a single broad constriction, parallel to ribs and striae. BMNH C78710 (Fig. 12F—G), a fragment tentatively referred to this species, shows perfectly preserved ornament at a whorl height of 42,5 mm. The ribs and striae are markedly prorsiradiate and pass across the venter with but a shallow convexity. Ribs are distant, and minutely crenulated, whilst between them are from three to five growth striae of variable strength and development. In addition, there is a delicate spiral ornament connecting the crinkles on successive ribs. Beyond: 150 mm diameter, it is convenient to describe the specimens in terms of two morphological extremes, with passage forms between. In variety « the dorsum is flattened and the venter quite narrowly rounded. The ribs flex slightly backwards over the umbilical wall and are radial to feebly prorsiradiate on the flanks, crossing the venter with only slight curvature. There are sparse, broad, shallow constrictions bounded by a prominent flared adoral rib. As diameter increases, the ribs become widely spaced and in most specimens growth striae are not preserved although a few specimens (e.g. Fig. 9A) show that both they and the delicate spiral ornament survive. On moulds the ribs appear as low, rounded undulations. In variety 8, the venter and dorsum are equally rounded, and the greatest breadth is at mid-flank—a square cross- section in which the corners are rounded. Ribs are distinctly prorsiradiate and there are occasional flared ribs, not, apparently, accompanied by constrictions. The suture line of this species is deeply and intricately subdivided (Fig. 6); L is large and asymmetrically bifid, U, larger than Uj, and I cruciform (Fig. 5B). The septal lobe is massive (Fig. 5A). Discussion The lectotype of Lytoceras vogdti is a juvenile, but shows an expansion rate and differentiation of ornament into ribs and striae like that of some of the 270 ANNALS OF THE SOUTH AFRICAN MUSEUM present similarly sized fragments, sufficient to suggest them to be conspecific. Some of the specimens resemble Lytoceras mikadyense Krenkel (1910: 233, pl. 22 (fig. 5)), but this species has consistently wider spaced ribs with striae between, indicating it to be closer to Eulytoceras. Of other species described by Krenkel, the unique holotype of ‘Crioceras’ schlosseri Krenkel (1910: 227, pl. 22(3) (fig. 15)), although more compressed, has ornament like that of some of the present specimens. Of species described from approximately contemporaneous strata in Madagascar, Lytoceras aff. vicinum Douvillé of Collignon (19625: 5, pl. 217 (fig. 952)) has a compressed whorl section with delicate concave, prorsiradiate flank ribs, whilst Lytoceras belliseptatiforme Collignon (19626: 6, pl. 217 (fig. 953), pl. 218 (fig. 953)) is densely and evenly ornamented by fine ribs, has a lower expansion rate and circular whorl section. Lytoceras mahafalense Collignon (1963: 14, pl. 246 (fig. 1503)) has a distinctly compressed oval whorl section and is of Albian age. The most similar species appears to be Lytoceras (Thysanolytoceras) strambergense mut. albensis Collignon (1963: 9, pl. 244 (fig. 1047)), a Lower Albian form with rather finer ribbing and a distinctively compressed whorl section and lower expansion rate. The species may be a descendant of the present form. Occurrence Lytoceras vogdti was first described from the Barremian of the Crimea; it has been subsequently recorded from the Upper Aptian of Mozambique, and now Zululand. Lytoceras aff. sauclum (Drushchitza, 1956) Figs 9B, 1OA—D, 11B Compare: Biasaloceras sauclum Drushchitza, 1956: 71, pl. 4 (fig. 13); Drushchitza & Kudryavtseva 1960: 256, pl. 5 (fig. 1). Lytoceras sauclum Wiedmann and Dieni, 1968: 29. Type The holotype is Drushchitza’s (1956, pl. 4 (fig. 13)) original specimen from the Hauterivian of the Crimea. Material BMNH C78704 from the Makatini Formation, Albian IV, locality 171, Mlambongwenya Spruit, Zululand. Dimensions D Wb Wh Wb: Wh U BMNH C78704. ._—‘122,0 52,5(43) 48,0(39) 1,1 49,0(40) Jjpsk CRETACEOUS FAUNAS FROM SOUTH AFRICA -ysniq) wnjonvs “jye sp4a20jAT ° a GX” TX JUSWIeUIO a]TUAANS Jo sjre}op ZuIMOYs ‘snefonu ‘POL8LD HNWA “(OSé6r “ez11Y9 ‘JUDWUIO JO Sleep SUIMOYS JUSLUSEIJ [IOYM ‘7OL8LD HNWE “LOGI “Yosexesey 4psoa spsa00jAT “VW "6 “SIF Die ANNALS OF THE SOUTH AFRICAN MUSEUM Description The coiling is very evolute, serpenticone, with a narrow, shallow, impressed dorsal area. The whorl section is depressed (whorl breadth to height ratio may be up to 1,1), expanding quite rapidly, with the greatest breadth some way below mid-flank. The umbilical seam is deeply incised, the umbilicus of moderate width (about 40% of diameter) and depth. The umbilical wall is high and rounded, the umbilical shoulder broadly rounded, the flanks flattened, ventro- lateral shoulders broadly rounded, and the venter somewhat flattened. The shell is ornamented by growth striae, and widely spaced crinkled ribs. Ribs and striae arise at the umbilical seam, curve backwards across the umbilical wall and forwards across the umbilical shoulder and are straight and slightly prorsiradiate across the flanks. The inner whorls have a distinctive Eulytoceras- like appearance, with between seven and ten striae between each pair of ribs, the ribs themselves being distinctly—if minutely—crenulate, with a gently inclined apical slope and an abrupt apertural face which give the surface of the shell a scale-like appearance (Fig. 9B). As size increases (Fig. 10) the ribs become more closely spaced, striae lose their prominence, crenulations become more distinct, and on some specimens, fine strigations appear on some areas of shell. All specimens bear occasional broad, deep constrictions on both shell and mould, preceded by a high, flared rib. Both flares and constrictions become increasingly prominent as diameter increases. The suture line (Fig. 11B) includes a deeply incised elongate E with a long, narrow lanceolate median saddle, a large, triangular, asymmetrically bifid L, and a similar smaller U, lobe, separated by a large, splayed bifid L/U, saddle. Discussion Drushchitza figured the type specimen of Biasaloceras sauclum in side view only, but the illustration shows clearly the differentiation of ornament on the Eulytoceras-like early whorls into ribs with groups of striae between, an inter- mediate stage with rather irregular ornament, and middle to late growth stages with straight, variably spaced crenulate ribs and occasional flares. The whorl section is slightly depressed, with the greatest breadth some way below mid-flank. Although of Hauterivian age, the striking similarity between the illustration and the present material suggests that they belong to the same group, in spite of age difference. Biasaloceras striatum Drushchitza (1956: 78, pl. 4 (figs 14a—b); Drushchitza & Kudryavtseva 1960, pl. 4 (figs 5a—b)) has a similar juvenile ornament, but a rather different whorl section; based on a juvenile, the authors are inclined to regard it as a possible variant of L. sauclum. The markedly different early and late ornament of Lytoceras sauclum helps to differentiate it from most Cretaceous Lytoceras species. Thus, of the long-ranging Stramberg species, Lytoceras liebigi Oppel (Zittel, 1868: 74, pl. 9 (figs Sa—b, 7a—b), pl. 10) has a depressed oval whorl with concave crenulate ribs. Lytoceras strambergense Zittel (1868: ATE: CRETACEOUS FAUNAS FROM SOUTH AFRICA ‘TX “pOL8LO HN ‘(S6r “eziyoysniqd) wnjonvs SX \ ‘ RRS ~ So SS Ye svsgd0JAT “OT “S14 te S aS YaR ss 274 ANNALS OF THE SOUTH AFRICAN MUSEUM 74, pl. 11 (figs 1-3)) has flexuous crenulate ribs, and Lytoceras sutile Oppel (1865: 551; Zittel 1868: 76, pl. 12 (figs 1-5)) has a circular whorl section and flexuous ribbing throughout ontogeny. Lytoceras aff. strambergense of Collignon (1949a: 71, pl. 12 (fig. 1), text-fig. 5) from the Hauterivian of Madagascar has distinctly concave ribs and flares. Lytoceras mahafalense Collignon (1963: 14, pl. 246 (fig. 1053)) from the Albian of Madagascar has a distinctly compressed whorl section and flattened flanks, whilst Lytoceras vicinum Douvillé (1916: 93, pl. 11 (fig. 6)) is also a compressed species with an elliptical to oval whorl section. A final species to mention, Lytoceras hennigi Zwierzycki, described below, has a depressed oval whorl section, and a very even, regular ornament. Occurrence Lytoceras sauclum was originally described from the Hauterivian of the Crimea and is also recorded from the Valanginian of Sardinia. Lytoceras hennigi Zwierzycki, 1914 Figs 11A, 12B—E, 13A-E Lytoceras sp. Krenkel, 1910: 224, pl. 22 (fig. 10). Lytoceras hennigi Zwierzycki, 1914: 40, pl. 4 (figs 6-7). Spath, 1939: 7. Type Lectotype herein designated is Zwierzycki’s (1914, pl. 4 (figs 6-7)) original figured specimen from the Trigonia schwartzi Bed (Barremian—Aptian) of Mikadi, Tanzania. Material Fragments of five individuals and one fairly complete specimen, BMNH C78705-9, C78711, from the Makatini Formation, Barremian 1, locality 171, Mlambongwenya Spruit, northern Zululand. Dimensions D Wb Wh Wb:Wh U Lectotype (from IE MATOS) 5 5 ‘ISS 58,0(37) 57,0(37) 1,02 61,0(39) BMNH C78708 — 37,0(—) 31,0(—) 1,19 — BMNH C78705___.. — 35,0(_ ) SUSE) 1,10 -- Description The coiling is very. evolute, with a narrow, shallow impressed area, the whorls expanding quite rapidly. The whorl section is a depressed oval, the greatest breadth being at mid-flank. Ornament consists of fine ribs, growth striae, and occasional flares. These arise at the umbilical seam, sweep sharply backwards over the inner umbilical wall where they are concave, then forwards CRETACEOUS FAUNAS FROM SOUTH AFRICA 275 Fig. 11. A. Partial suture of Lytoceras hennigi Zwierzycki, 1914. BMNH C78705. x 5. B. Partial suture of Lytoceras aff. sauclum (Drushchitza, 1956). BMNH C78704. x 4,5. 276 ANNALS OF THE SOUTH AFRICAN MUSEUM to pass straight across the umbilical shoulder and flanks where they are markedly prorsiradiate, to cross the venter with a faint forwards projection. The best preserved specimens show the ribs to have been even, sharp-topped and minutely crinkled, and separated by wider interspaces ornamented by irregular fine striae. Delicate spiral ridges connecting crinkles are present over the whole of the shell surface when well preserved. There are periodic high, sharp flares (Fig. 12D-E), although these are only rarely preserved. When shells are partially exfoliated, growth striae, crinkles and strigations disappear, leaving only evenly spaced, rounded ribs and occasional constrictions (corres- ponding to the site of flares). Moulds are smooth save for constrictions (Fig. 12D). The sutures are poorly exposed, but include a large E/L and smaller L/U,, both deeply incised, asymmetric and triangular, separated by a deep splayed L (Fig. 11A). A distinct siphonal band is present in one specimen (Fig. 12D). Discussion The present specimens are much smaller than Zwierzycki’s lectotype, but a comparison with topotype specimens in the British Museum (Natural History) suggests that they are indeed referrable to Lytoceras hennigi. The whorl section of topotype specimens is depressed at small diameters, whilst the ornament consists of even, crinkled ribs with smooth, somewhat wider interspaces between, and occasional strong periodic flares, marked by broad constrictions on the internal mould. The regular, fine ribs and periodic flares of L. hennigi readily distinguish it from Eulytoceras species discussed below, whilst Lytoceras liebigi, L. stramber- gense and L. sutile differ in proportions and ornament, as noted above. Lytoceras subsequens Karakash (1907: 49, pl. 5 (fig. 9, 9a), pl. 24 (fig. 32)) lacks prominent constrictions and flares; L. aff. sauclum has completely different, Eulytoceras- like juvenile ornament as discussed above, whilst Lytoceras puezanus Haug (1889: 197, pl. 8 (fig. 1) pl. 10 (fig. 1)) has less marked flares and very strong strigations. Lytoceras densefimbriatum Uhlig (1883: 191, pl. 6 (figs la—c, 2)) is a very finely ornamented, compressed form with weaker flares and stronger strigations, Lytoceras mahafalense Collignon is compressed, whilst Lytoceras subfimbriatum d’Orbigny (1840: 121, pl. 35) has a compressed whorl section, very fine fimbriate ornament, and distant flares, four to five per whorl. Occurrence Lytoceras hennigi is known from the Lower Cretaceous of Tanzania and the Upper Barremian of Zululand only. Genus Eulytoceras Spath, 1927 Type species Ammonites inaequalicostatus d’Orbigny 1840. CRETACEOUS FAUNAS FROM SOUTH AFRICA 277 MMM, yy to) y Ye TY, Yih: G Fig. 12, A, F, G. Lytoceras vogdti Karakasch, 1907. A. BMNH C78703. F-G. BMNH C78710. x 1. B-E, Lytoceras hennigi Zwierzycki, 1914. B—D. BMNH C78708. E. BMNH C78705. ae 278 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 13. Lytoceras hennigi Zwierzycki, 1914. A-B. BMNH C78705. C-E. BMNH C78706. Phe Os CRETACEOUS FAUNAS FROM SOUTH AFRICA 279 Diagnosis Compressed to round-whorled serpenticone lytoceratids with slowly expanding whorls ornamented by fine, regular, distant rectiradiate to prorsi- radiate ribs and periodic flares, all of which extend across the venter without interruption. Discussion The regular, fine, distant ribs and occasional simple flares of typical Eulytoceras clearly separate them from Lytoceras itself (with typically crinkled ribs and flares and periodic constrictions), Hemilytoceras (with smooth inner whorls and high, closely spaced concave flares), Pterolytoceras (with minutely crinkled growth lines and irregular fine ribs), Ammonoceratites (Ammonoceratites) and A. (Argonauticeras), both of which have fine, dense, subdued ornament. As the authors have noted, however, some species such as Lytoceras sauclum, described above, have Eulytoceras-like inner whorls, whilst ‘Lytoceras’ mikadyense Krenkel has Eulytoceras inner whorls and develops very closely spaced ornament when adult, being transitional to Lytoceras sensu stricto. Occurrence Eulytoceras ranges from the Hauterivian to Lower Aptian, and species are known from southern Europe, the Crimea, Caucasus, Bulgaria, east Africa (Tanzania), South Africa (Zululand), Madagascar, and California. Eulytoceras phestum (Matheron, 1878) Figs 14A-E, 15 Ammonites phestus Matheron, 1878, pl. c—20 (fig. 5). Lytoceras phestus Uhlig, 1883: 187, pl. 5 (figs 1-4, 20). Trautschold, 1886: 137. Haug, 1889: 196, pl. 8 (fig. 2). Simionescu, 1898: 59, pl. 2 (figs 4, 10), pl. 20 (fig. 17). Sarasin & Schondelmayer, 1901: 19. Karakasch, 1907: 46, pl. 4 (fig. 10), pl. 20 (figs 5—6). Pervinquieére, 1907: 64. Kilian & Reboul, 1915: 21, pl. 1 (figs 1-2). Petkovic, 1921: 48. Kulzhinskaya- Voronets, 1933: 5, fig. 6. Rouchadzé, 1933: 174. Ksiazkiewicz, 1938: 230, pl. 1 ee 1). Eristavi, 1955: 53; 1957: 59. Lytoceras gresslyi Somogyi (non Hantken), 1914, pl. 11 (fig. 12). ? Lytoceras lorentheyi Somogyi, 1914: 300, pl. 11 (figs 10-11). ? Protetragonites phestus Nagy, 1967: 62, pl. 2 (fig. 1). Eulytoceras phestum Spath, 1927: 64. Drushchitza, 1956: 87, pl. 5 (fig. 18); Drushchitza & Kudryavtseva, 1960: 258, pl. 6 (fig. 3). Collignon, 1962a: 95, pl. 214 (fig. 935). Dimitrova, 1967: 27, pl. 9 (fig. 3). Kotetichvili, 1970: 60, pl. 4 (fig. 3). Vasicek, 1972: 37, pl. 2 (fig. 6). Murphy, 1975: 18, pl. 2 (fig. 3). Type The ‘holotype’ is Matheron’s original specimen, in the collections of the Muséum d’Histoire Naturelle de Marseille according to Murphy (1975: 18), but Matheron in fact gives three figures, and it is not clear if these represent individual specimens or composite drawings. Lectotype designation (if necessary) must await fuller redescription of the type material. 280 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 14. Eulytoceras phestum (Matheron, 1878). A-C. BMNH C78698. D-E SAS L7(5). ; A=Di xX ke Bx xe CRETACEOUS FAUNAS FROM SOUTH AFRICA 281 Fig. 15. Partial suture of Eulytoceras phestum (Matheron, 1878) based on SAS L7(5). x 8. Material Two specimens, SAS. L7(5) and BMNH C78698, both from the Makatini Formation at locality 170, Mlambongwenya Spruit, Zululand (Barremian 1). Description The material consists of a wholly septate juvenile 27,5 mm in diameter, and a body chamber fragment of a larger individual. The coiling is very evolute, the whorls only just in contact, with, as a consequence, a very narrow impressed area. The coiling is evolute, the umbilicus being approximately 30 per cent of the diameter, and shallow with a rounded wall. The whorl section is as broad as high, rounded trapezoidal, with a rounded ventrolateral shoulder, convergent, somewhat flattened sides, the greatest breadth being some way beyond mid-flank, and a broadly rounded venter. Inner whorls up to a diameter of 15 mm appear to be devoid of all ornament. Beyond this ornament consists of fine, narrow ribs separated by wide, smooth interspaces. The ribs arise at the umbilical seam, pass forwards across the umbilical wall and shoulder and are prorsiradiate and weakly flexed or convex on the flanks. Interspaces are ornamented by fine growth striae. The external suture is partially exposed, as shown in Figure 15. Discussion The present specimens match closely with Matheron’s original figures in terms of general ornament and proportions, his illustrations being based upon crushed specimens. Eulytoceras phestum is readily distinguished from Eulytoceras inaequalicostatus (d’Orbigny) (1840: 18, pl. 29 (figs 3-4)), for that species develops strong flares, nine per whorl in D’Orbigny’s figure. Eulytoceras raricinctum (Uhlig) (1883: 88, pl. 5 (figs 5—7)) is a distantly ribbed form with only fifteen ribs per whorl rather than the forty to forty-two typical of E. phestum. Eulytoceras rotundatum Drushchitza (1956: 86, pl. 5 (fig. 15)) has a depressed 282 ANNALS OF THE SOUTH AFRICAN MUSEUM whorl section with fewer, more convex ribs. Eulytoceras intemperans (Coquand in Matheron) (1878, pl. c—20 (fig. 4-4c)) has only twenty-one ribs per whorl, with distinctive striae between, whilst Eulytoceras electra (Coquand in Matheron) (1878, pl. c-20 (fig 2a—2b)) has fifteen flexuous ribs per whorl, also with striae between. The east African “Lytoceras’ mikadyense (Krenkel) (1910: 223, pl. 22 (fig. 5)) is a depressed species, has periodic flares, rather irregular ribs, and prominent striae between. Eulytoceras belchasifakaense Collignon (1949a: 70, pl. 11 (fig. 2-2a), text-fig. 4), from the Hauterivian of Madagascar, has straight, quite distantly spaced ribs, rather than the slightly flexuous ornament of E. phestum, but may subsequently prove to be a synonym. E.? komihevitraense Collignon (1963: 15, pl. 247 (fig. 1055)), from the Middle Albian of Madagascar, is said to possess constrictions, and has dense, distinctly concave ribs with a ventral sinus. Finally, Eulytoceras lepidum (d’Orbigny) (1840: 149, pl. 48 (figs 3—4)) bears strong flares, and is more distantly ribbed. Occurrence Eulytoceras phestum is a typically Barremian species, known from southern France, the Tyrol, Czechoslovakia, Hungary, Bulgaria, the Crimea, Georgia and the Carpathians, Tunisia, Madagascar, and South Africa. Kilian & Reboul (1915: 21) also cite the species from the Lower Aptian of southern France, whilst Murphy (1975: 18) records it from the Upper Barremian of California. Genus Ammonoceratites Bowdich, 1822 Subgenus Ammonoceratites Bowdich, 1822 Type species Ammonoceratites lamarcki Bowdich, 1822. Diagnosis Large, evolute, serpenticone lytoceratids with a circular whorl section, ornamented by dense, crenulate fine ribs and growth striae, sometimes combined with faint spiral striae. Occasional broad, shallow constrictions may appear. Suture highly subdivided, with a massive cruciform septal lobe (Is). Discussion A general lack of flares and associated constrictions, fine ornament and rounded whorls separates Ammonoceratites (Ammonoceratites) from Lytoceras Suess, 1865 sensu stricto, whilst lack of regular, distant flares differentiates it from Eulytoceras Spath, 1927, Hemilytoceras Spath, 1927, and Pterolytoceras Spath, 1927. Ammonoceratites (Argonauticeras) is readily separated by virtue of its rapidly expanding whorls and an ornament of very fine, even, generally non-crenulate ribs. Considerable confusion surrounds the introduction of this genus, as much CRETACEOUS FAUNAS FROM SOUTH AFRICA 283 as that surrounding the type species, as discussed below. Following Casey (1960: 2), Ammonoceratites Rafinesque, 1815, is regarded as a nomen nudum and Ammonoceras Lamarck, 1822, a nomen dubium (as introduced) over which Ammonoceratites Bowdich, 1822 (both of which have type species based on the same specimen), has priority. The following species have been referred to the restricted subgenus: A. (Ammonoceratites) lamarcki Bowdich (1822: 21, pl. 3 (fig. 14)). A. (Ammonoceratites) glossoidea (Lamarck) (1822: 144 in Chenu 1859: 90, text-figs 391-392). | A. (Ammonoceratites) mahadeva (Stoliczka) (1865: 16, pl. 80 (fig. 1-1b)), Middle Albian of southern India. A. (Ammonoceratites) crenulatum (Crick) (1907: 236), Upper Albian of Zululand. A. (Ammonoceratites) betiokyense Collignon (19625: 8, pl. 218 (figs 954-955)), Upper Aptian of Madagascar. A. (Ammonoceratites) crenocostatum (Whiteaves) (1884: 45, pl. 9 (fig. 2)), Albian of British Columbia. . (Ammonoceratites) ezoense (Yabe) (1903: 9, pl. | (fig. 1), pl. 5 (fig. 1)), Upper Albian of Japan. aN Occurrence Ammonoceratites ranges from Upper Aptian to Upper Albian. There are records from British Columbia, southern France, southern India, Japan, Madagascar, and South Africa. Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1865) Figs 16-33 ? Ammonoceratites lamarcki Bowdich, 1822: 21, pl. 3 (fig. 14). ? Ammonoceras glossoidea Lamarck, 1822: 644. ? Ammonoceratites glossoidea Chenu, 1859: 90, text-figs 391-392. Ammonites mahadeva Stoliczka, 1865: 165, pl. 80 (fig. 1). Ammonites crenocostatus Whiteaves, 1876: 45, pl. 9 (fig. 2). Lytoceras batesi Whiteaves (non Trask), 1884: 202, pl. 27 (fig. 1); 1900: 270. Lytoceras mahadeva Kossmat, 1895: 17. ? Lytoceras ezoense Yabe, 1903: 9, pl. 1 (fig. 1), pl. 5 (fig. 1). ? Lytoceras crenulatum Crick, 1907: 236. ? Lytoceras sp. aff. mahadeva: Jeannet, 1908: 105-119, pls 3-6. Lytoceras (Ammonoceratites) crenulatum: Breistroffer, 1936: 169, text-fig. 10h. Ammonoceratites cf. mahadeva Collignon, 19495: 45, text-fig. 6. Ammonoceratites mahadeva: Collignon, 1963: 12, pl. 245 (fig. 1051). Ammonoceratites crenocostatus McLearn, 1972: 22, pl. 1 (fig. 5), pl. 2 (figs 1-3). Type The holotype is Stoliczka’s (1865, pl. 80 (fig. 1)) original specimen from the Lower Utatur Group of the environs of Moraviatoor, southern India, and presumably of Albian age. The original illustrations are reproduced here as Figures 27-28. 284 ANNALS OF THE SOUTH AFRICAN MUSEUM Material Eight specimens: BMNH C18265, the holotype of Lytoceras crenulatum Crick, from the Albian of the Mzinene River area; BMNH C78890, from Bed 2 of the Mzinene Formation at locality 35 on the Mzinene River (Albian III); BMNH C78700-1, from the Mzinene Formation at locality 54 on the Mzinene River (Albian V), SAS A1195 and 2004 come from the same horizon at locality 53. SAS Z418 and Z428 in the Van Hoepen Collection are from Impala, Ndumu, Mzinene Formation (Albian II or III). Dimensions D Wb Wh Wb:Wh U Holotype (after Stoliczka . 218,0 82,0(38) 78,0(36) 1,05 95,0(44) A. ezoense (after Yabe) . 200,0 75,0(37) 80,0(40) 0,94 70,0(35) A. crenocostatum holotype, GSC 4987 (after McLearn) 44,0 14,7(33,5) 15,4(35,0) 0,95 20,7(47) A. crenocostatum, GSC 4976 = 47,0(—) 40,0(—) 1,08 — (—) BMNH C78700 ae 44,8(—) 41,5(—) 1,08 — (—) — 29,0(—) 28,0(—) 1,04 — (—) SAS A1195, at . 115,0 45,0(39) 44,0(38) 1,03 48,0(42) SAS Z428, at 137,0 49,0(36) 48,0(36) 1,02 58,0(42) ate" 194,0 75,09) 74,0(38) 1,03 83,5(43) BMNH C78890 220,0 93,0(42) 88,0(40) 1,05 85,0(38) SAS Z418 — 78,0(—) 76,0(—) 1,03 — (—) SAS A2008 . . . 253,0 98,0(39) 96,0(38) 1,01 101,0(40) Description Very large (the Zululand specimens are still septate at diameters of over 300 mm), very evolute, the whorls only just in contact with a narrow, shallow dorsal impressed zone. The whorl section is almost circular, typically just slightly broader than high, the ratio varying from 1,0 to 1,05. The whorls expand quite rapidly and the umbilicus is broad, varying from 38 to 42 per cent of the diameter and increasing slightly during ontogeny, with a deeply incised umbilical seam. Ornament changes markedly throughout growth. At the smallest diameters visible (Figs 16A, 31C) the shell surface appears virtually smooth, bearing only fine prorsiradiate growth striae, distant, minutely crenulated ribs, and occasional flares which correspond to distinct constrictions on internal moulds. Ribs, flares and striae arise at the umbilical seam, are concave on the inner part of the umbilical wall, then sweep forwards and are prorsiradiate and feebly 285 CRETACEOUS FAUNAS FROM SOUTH AFRICA ~ we oe . ee SRE PR. B we x a RAT “aS SS © ASSES ERS ens . SSS Fig. 16. Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1865). A. SAS A2004. B. BMNH C78890. x 1. 286 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 17. Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1865). SAS A2004. x 0,45. convex on inner and mid-flank (Figs 17, 20). Beyond this, the crenulated ribs become increasingly frequent, and there are sparse prominent flares and associated constrictions. In middle and later growth ribs are strong and closely spaced (Fig. 20). During these later growth stages, ribs and striae arise at the umbilical seam, sweep strongly backwards across the inner part of the umbilical wall and forwards over the umbilical shoulder. They are distinctly prorsiradiate and variably, although commonly, weakly concave across the CRETACEOUS FAUNAS FROM SOUTH AFRICA 287 We Has highs y ws oe (a Fig. 18. Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1865). SAS A2004. x 0,45. flanks and sweep forwards across the ventrolateral shoulder to produce a broad ventral peak of variable depth. Growth striae are scarcely visible to the naked eye, but the ribs are very distinct and strongly crenulate with a vertical apertural face and a gently inclined apical slope. On the ventral and ventrolateral region of some of the Zululand specimens there are distinct, curved longitudinal striae between and connecting the crinkled ribs, arising from individual crinkles (Fig. 20). . Partially exfoliated shells show low, broad, irregular, fold-like constricted 288 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 19. Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1865). SAS 2418. x 1. CRETACEOUS FAUNAS FROM SOUTH AFRICA 289 Fig. 20. Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1865). BMNH C78890. x 0,6. areas during middle and later growth (Figs 22—23), and in some places, transverse rows of large pits corresponding to the site of crinkled ribs. Weathered shell surfaces are also covered in minute pits. Internal moulds are essentially smooth, save for broad, shallow, fold-like constrictions (Fig. 19). One specimen shows a dorsal callus, through which traces of-the under- lying ventral ornament are still visible. ANNALS OF THE SOUTH AFRICAN MUSEUM 290 BMNH C78890. text-fig. 391. ° 9 1865) ka, icz Copy of Chenu 1859 h. i Ammonoceratites (Ammonoceratites) mahadeva (Stol A. ° Dal x 0,6. B—D. Ammonoceratites lamarcki Bowd 1g F CRETACEOUS FAUNAS FROM SOUTH AFRICA 291 * < : - , 4 - i, i ‘ dic Ae oe Gee ‘ Gy 7. ae Fig. 22. The holotype of Ammonoceratites (Ammonoceratites) crenulatum (Crick, 1907). BMNH C18265. Slightly reduced. The suture line is highly subdivided with a long spatulate ventral saddle, a large bifid asymmetric E/L saddle and a smaller also asymmetrically bifid L/U, saddle, separated by a very large lateral lobe (L), deeper than the external lobe (E) and bifid, with a large subdivided median element. U, is highly sub- divided, dorsal internal lobe (1) cruciform, septal lobe large (Fig. 18). Discussion The Zululand specimens show some variation in strength and direction of ornament and relative proportions when compared with Stoliczka’s magnificent 292 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 23. The holotype of Ammonoceratites (Ammonoceratites) crenulatum (Crick, 1907). BMNH C18265S. Slightly reduced. specimen. In particular, the ventral peak is more prominent than shown in his figure and the ribs are prorsiradiate rather than radial. The material varies in these respects, and rather than trying to separate it into Ammonoceratites mahadeva with straight ribs and a shallow ventral peak and Ammonoceratites crenulatum, with prorsiradiate ribs and a deeper peak, the authors regard their material as one variable species. It is, however, far from clear whether Ammonoceratites mahadeva is the earliest name used for this species. Lamarck (1822, vol. 7: 644) introduced the name Ammonoceras glossoidea, CRETACEOUS FAUNAS FROM SOUTH AFRICA 293 A B C Fig. 24. Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1865). BMNH C78700. x 1. without figuring the material. His description is brief: ‘Cette coquille rompue en trois morceaux, qui s’appartiennent successivement, et dont l’un offre Pextrémité supérieure de cette méme coquille, est d’une assez grande taille, fort épaisse en sa partie inférieure, arcuée presque en demi-cercle, et se termine supérieurement en forme de langue. Ses loges sont remplies de matiére pierreuse, et leurs cloisons ne se distinguent que dans les parois ot! leurs concours forment des sutures lobées, laciniées, rameuses, tout a fait analogues a celles des ammo- nites. Mais la coquille dont il s’agit en est trés-distincte par sa forme générale; car malgré son arcuation, elle n’efit point forme de tours contigus, si la nature Petit agrandie davantage. Sa longueur est de cinquante centimetres.’ In the same year, Bowdich (1822: 21, pl. 3 (fig. 14)) introduced the name Ammonoceratites lamarcki, based upon the same fragments. His description is ANNALS OF THE SOUTH AFRICAN MUSEUM 294 T X “00L8L0 HNWA “(6987 “8 4Z91101¢) Aapvynur (sazwsaoouoUMU) SaqyvsaoUoUMMUp *SZ “BIZ | oe SS x CRETACEOUS FAUNAS FROM SOUTH AFRICA 295 : | a | Fig. 26. External suture of Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1865). BMNH C78890. x 1,6. also brief: -“Septa numerous, undulated at the margins, pl. 3, fig. 14. Siphon marginal, interior. Rolled on itself in the same plane? The locality is unknown, M. Lamarck purchased it by accident: he kindly allowed me to take it home in order that the figure, the first that has been made, might be as accurate as possible.’ Subsequent authors have varied in their views of the nature of Ammonoceras glossoidea— Ammonoceratites lamarcki (the latter has priority; fide Casey 1960), regarding it as of both Jurassic and Cretaceous age and as a species of Hamites, Toxoceras, and Pictetia. The age is, however, in all probability Cretaceous, for as Chenu (1859: 90) notes, the specimen was not purchased by Lamarck, for he (Chenu) quotes Valenciennes as telling him that ‘Lamarck lui a souvent répété que ce fossile, dont il faisait grand cas, avait été rapporté par le secrétaire de la colonie de Pondichéry, et donné au fils de Buffon, qui lui en avait luiméme fait cadeau. M. Valenciennes ajoute que parmi les ammonites rapportées de l’Inde par Jacquemont, il se trouve aussi un fragment de I’A. fimbriatus.’ Chenu’s figure is much better than that of Bowdich, and is reproduced here as Figure 21B—D. The specimen was said to be 500 mm in diameter, and corresponds to the generally accepted interpretation of Ammonoceratites; its occurrence in southern India indicates that it may well be a specimen of what has subsequently been termed Ammonoceratites mahadeva. The figures are, 296 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 27. Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1861). Copy of Stoliczka’s figures of the holotype, reduced here by approximately 0,75. CRETACEOUS FAUNAS FROM SOUTH AFRICA 297 Fig. 28. Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1861). Copy of Stoliczka’s figure of the holotype, reduced here by approximately 0,75. however, so poor that, until the specimen is redescribed and refigured, the authors would prefer to use Stoliczka’s name, having sought without success for the Lamarck/Bowdich type in the Paris museums. Of other species, Ammonoceratites (Am.) betiokyense, from the Aptian of Madagascar, is based on two small fragments only, but appears to have distinctly flattened sides and a markedly asymmetric or even subtrifid U, lobe 298 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 29. Ammonoceratites (Ammonoceratites) ezoense (Yabe, 1903). Lectotype. x 0,75. according to Collignon (19625: 8). Ammonoceratites (Am.) ezoense (Yabe), lectotype, herein designated, Yabe’s (1903: 9, pl. 1 (fig. 1), pl. 5 (fig. 1)) original specimen is also an Albian species (see range charts in Matsumoto 1954), the original material being preserved in the collections of the Geology Department of Tokyo University, and refigured here as Figures 29-30, 31A—B. Yabe separated it from Ammo- CRETACEOUS FAUNAS FROM SOUTH AFRICA 299 Fig. 30. Ammonoceratites (Ammonoceratites) ezoense (Yabe, 1903). Lectotype. x 0,75. noceratites (Am.) mahadeva on the basis of more rapidly expanding whorls and a smaller umbilicus (Yabe’s measurements are reproduced above), but as can be seen from the present photographs, the preservation is poor and the specimen probably within an acceptable range of variation for A. (Am.) mahadeva as shown by the Zululand specimens (e.g. compare Figs 29-30, 31A—B with Figs 17-19); the authors therefore place it as a questionable synonym. 300 ANNALS OF THE SOUTH AFRICAN MUSEUM » iy oe se if RY re , Fig. 31 A-B. Ammonoceratites (Ammonoceratites) ezoense (Yabe, 1903). Part of lectotype, showing suture. C. Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1861); inner whorls of SAS A2004. x 1,5. CRETACEOUS FAUNAS FROM SOUTH AFRICA 301 Ammonoceratites (Am.) crenocostatus (Whiteaves) is a further Middle? to Upper Albian species known only from the Queen Charlotte Islands off the coast of British Columbia. The authors have been able to examine casts of the holotype and other specimens from the area, illustrated here as Figures 32-33. The holotype is only 43 mm in diameter, and it retains only part of the shell. The inner whorls are strongly constricted on the mould up to a diameter of 20 mm, with traces of delicate striae where shell is preserved; from 26 to 40 mm there are distant fine ribs with delicate growth striae between, both ribs and striae being prorsiradiate and feebly convex on the venter. Over the last preserved part of the specimen the ribs crowd markedly. GSC 4976 is a larger specimen, with an original diameter of approximately 120-130 mm. The outer whorl shows distinctly concave prorsiradiate flank ribs, with a marked convex ventral peak, rib density, form and strength matching closely to the similarly sized Zululand specimen BMNH C78700 (Figs 24-25). So far as can be judged there are no criteria that can be used to separate A.(Am.) crenocostatum from the Zululand material, and it is therefore suggested that it is a further synonym of Ammonoceratites (Am.) mahadeva. Occurrence As defined above, Ammonoceratites (Am.) mahadeva ranges from Middle to Upper Albian, and is known from southern India, the Queen Charlotte Islands, Madagascar, Japan, Zululand, and perhaps the Jura (Jeannet 1908: 105-119, pls 3-6). Subgenus Argonauticeras Anderson, 1938 Type species Lytoceras argonautarum Anderson, 1902. Diagnosis A subgenus of Ammonoceratites with a high expansion rate, subrectangular to trapezoidal whorl section, fine, equal, dense, straight to gently flexed ribs or striae without obvious crinkles. Weak, broad constrictions may be present on early whorls. Discussion Argonauticeras can be separated from other Cretaceous lytoceratid genera by its fine, even, typically non-crenulate ornament and high expansion rate. The trapezoidal whorl section, emphasized by Wright (in Arkell & Wright 1957), is a less distinctive feature, as discussed below. Differences from Ammonoceratites (Ammonoceratites) are outlined above. Four species have been referred to the subgenus: Argonauticeras argonautarum (Anderson) (1902: 85, pl. 7 (figs 154-155); 1938: 149, pl. 17 (fig. 3), pl. 19 (figs 1-2)). Upper Aptian of California. ANNALS OF THE SOUTH AFRICAN MUSEUM 302 ‘L860 OSD ‘odA}0[OH{ “(QL8I ‘SoAvoUY AA) SJDJSODOUAAI (SAJNDAQIOUOWIU) SaJ1JDAIIOUOUP*Z | “BLT J dq | V 303 CRETACEOUS FAUNAS FROM SOUTH AFRICA TX ‘OL6b OSD ‘adModAH “(OL8T ‘soAvoY AMA) SNIDJSOIOUAID (SaJIIDAZIOUOUILUP) SafjjosQIOUOUUMp “EE “SLA pS 304 ANNALS OF THE SOUTH AFRICAN MUSEUM Argonauticeras depereti (Kilian) (1892: 8, pl. 1 (fig. 2). Thomel, 1968: 684-687, pl. 35 (figs 1-4)). Upper Aptian of southern France. Argonauticeras besairiei Collignon (19495: 46, pl. 8 (fig. 1)). Upper Aptian to Middle Albian of Madagascar. Argonauticeras belliseptatum (Anthula) (1899: 97, pl. 6 (fig. 1), pl. 7 (fig. 1)). Aptian of the Caucasus and Madagascar. Of these species, Dimitrova (1967: 28, pl. 10 (fig. 6)) figured a fragment referred to as Pictetia belliseptata (Anthula) which appears actually to uncoil. The authors therefore take the opportunity to figure photographically the holotype of A. (Ar.) belliseptatum (Figs 40-43) which shows it to be an Argonauticeras. Pseudotetragonites Drushchitza, 1956 (type species P. kudrjavzevi Drushchitza), is a subjective synonym of Argonauticeras, based on juveniles. Occurrence Argonauticeras is known from the Upper Aptian of southern France, the Caucasus, California, the Lower and Middle Albian of Madagascar, and the Middle Albian of Zululand. Ammonoceratites (Argonauticeras) depereti (Kilian, 1892) Figs 34-37, 38B, 39-43 Lytoceras depereti Kilian, 1892: 8, pl. 1 (fig. 2a—b). Simionescu, 1900: 656. Lytoceras belliseptatum Anthula, 1899: 97, pl. 6 (fig. 1), pl. 7 (fig. 1). Collignon, 1963: 12, pl. 245 (fig. 1052). Lytoceras argonautarum Anderson, 1902: 85, pl. 7 (figs 154-155). Tetragonites depereti Jacob, 1907: 12. Non Fallot, 1920: 243, pl. 2 (fig. 5), text-fig. 13. Lytoceras (Argonauticeras) argonautarum: Anderson, 1938: 140, pl. 17 (fig. 3), pl. 19 (figs 1-2). Pseudotetragonites kudrjavzevi Drushchitza, 1956: 80, pl. 7 (fig. 25), text-fig. 35. Orlov, 1958: 58, pl. 20 (fig. 4), Drushchitza & Kudryavtseva, 1960: 257, pl. 5 (fig. 2a—b), text-fig. 64. Ammonoceratites (Argonauticeras) depereti Collignon, 1956: 100. Thomel, 1968: 684, pl. 35 (figs 1-4), text-figs. ? Pictetia belliseptata Dimitrova, 1967: 28, pl. 10 (fig. 6). Argonauticeras belliseptatum Forster, 1975: 144, pl. 1 (fig. 6). Type The lectotype, herein designated, is Kilian’s original specimen from the Upper Aptian of the environs of Barréme (Basses Alpes), south-eastern France, in the collections of Muséum d’Histoire Naturelle, Lyon. Material Two specimens, UPG B4 and UPG B5 from the Makatini Formation, Upper Aptian, Manyola Drift on the Usutu River, at 26°49’15” S 30°13’30" E. CRETACEOUS FAUNAS FROM SOUTH AFRICA 305 Fig. 34. Ammonoceratites (Argonauticeras) depereti (Kilian, 1892). UPG B4. x 0,78. 306 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 35. External suture of Ammonoceratites (Argonauticeras) depereti (Kilian, 1892). x 0,8. Dimensions Californian specimen (after Anderson, 1933) ere ree Holotype of A. (Ar.) belliseptatum (after Anthula, 1899) Lectotype of A. (Ar.) depereti (after Kilian, 1892) . French specimens (after Thomel, 1968) LSI. i527. 158) UPG B4. UPG B5. Description D 250,0 32550 53,0 38,3 55,0 102,0 179,0 Wb 125,0(50) 153,0(47) 25,0(47) 19,3(50) 27,0(49) 43,0(42) 70,0(39) 40,0(—) Wh 113,0(45) 130,0(40) 27,0(39) 16,0(41) 23,0(41) 44,5(43) 67,0(27) 7.0) Wb: Wh iol 1,20 7 0,96 1,04 1,08 U 87,0(35) 123,0(38) 21,0(39) 14,4(37) 19,2(34) 38,0(36) 73,0(40) aa ee) The largest specimen in the collection consists of one and a half whorls, parts of which are damaged. Coiling is very evolute, and whorls increase rapidly in height and width. The whorl section is slightly wider than high with rounded umbilical and ventrolateral edges; the dorsal impressed zone is small. Ornament consists of narrow, dense unbranched ribs which pass forwards over the flanks with a slight curvature and straight across the venter. Broad, CRETACEOUS FAUNAS FROM SOUTH AFRICA 307 A B Fig. 36. Ammonoceratites (Argonauticeras) depereti (Kilian, 1892). UPG BS. x 1. low constrictions are also present, paralleling striae; the number of constrictions per whorl cannot be determined, but on the smaller specimen, UPG BS, there are four equidistant constrictions in one-third of a whorl. The suture is only partially exposed; E is narrow and bifid, L is very wide and deeper than both E and U;. The saddles E/L and I/U, are bifid and have slender stems. Discussion Thomel recently (1968) reviewed this species and pointed to the changes in whorl section, relative dimensions and density of ornamentation during ontogeny. The Zululand specimens fall within the range of relative proportions given by Thomel (1968: 685) and are virtually identical to the figured specimens as far as the whorl section, fine ornamentation and constrictions are concerned. The largest known French specimen of A. (Ar.) depereti is only 102 mm in 308 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 37. Ammonoceratites (Ammonoceratites) depereti (Kilian, 1892). A-—B. Copies of Kilian’s figures of the holotype. C. Copy of Anderson’s (1938: pl. 17 (fig. 3)) figure of a juvenile Lytoceras (Argonauticeras) argonautarum (Anderson, 1902). All figures x 1. 309 CRETACEOUS FAUNAS FROM SOUTH AFRICA ¢ UIE) Madadap (spda1jnvUosAP) Sajj{DADIOUOUU d I Xx ‘I X ‘ph DdN “(681 SOP DdN (Z68T “UPI ) Nadsadap “jye (SDAZINNDUOSAP) SAUJVADIOUOUUp “WY “BE ‘SI 310 ANNALS OF THE SOUTH AFRICAN MUSEUM Midge EME th papappps aidlai ond We isi isi Lipton ue a ewe jp lus gu Sj” SE MMLZ io SE Me Bos WE RAED << giana ee Le. ea ies. X 0,75. 1€S ] figures of the spec S origina Fig. 39. Ammonoceratites (Argonauticeras) argonautarum (Anderson, 1902). Copies of Anderson’ CRETACEOUS FAUNAS FROM SOUTH AFRICA 311 Fig. 40. Ammonoceratites (Argonauticeras) belliseptatum (Anthula, 1899). Holotype. x 0,45. (Photograph supplied by R. A. Reyment.) 312 ANNALS OF THE SOUTH AFRICAN MUSEUM ZL WOW Fig. 41. Ammonoceratites (Argonauticeras) belliseptatum (Anthula, 1899). Holotype. x 0,45. (Photograph supplied by R. A. Reyment.) CRETACEOUS FAUNAS FROM SOUTH AFRICA 313 Fig. 42. Ammonoceratites (Argonauticeras) belliseptatum (Anthula, 1899). Holotype. x 0,45. (Photograph supplied by R. A. Reyment.) 314 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 43. Ammonoceratites (Argonauticeras) belliseptatum (Anthula, 1899). Holotype. x 0,45. (Photograph supplied by R. A. Reyment.) CRETACEOUS FAUNAS FROM SOUTH AFRICA B15 diameter, and thus difficult to compare with the holotype of Argonauticeras argonautarum (Anderson), the original figures of which are reproduced here as Figure 39. A smaller specimen figured subsequently by Anderson (1938: pl. 17 (fig. 3)) is reproduced here as Figure 37C; it appears identical to Kilian’s lectotype (reproduced here as Fig. 37A—B), and in consequence the authors regard A. (Ar.) argonautarum as a junior subjective synonym of A. (Ar.) depereti. A. (Ar.) belliseptatum Anthula is based on a giant disc 325 mm in diameter. When compared with the present material, the overall proportions, expansion rate, style and direction of ornament are so similar that the authors believe this to be a synonym of A. (Ar.) depereti. The remaining species referred to the subgenus, A. (Ar.) besairiei Collignon (19495: 46, pl. 8 (fig. 1)), is also very similar, and, indeed, Forster (1975: 144) regarded it as a synonym of A. (Ar.) belliseptatum. The juvenile holotype has, however, a distinctly trapezoidal whorl section with a broad, flattened venter, flattened flanks and, according to Collignon, has straight and rather coarse ribs for the genus. A larger Mada- gascan specimen figured subsequently (Collignon 19626: 10, pls 219-220 (fig. 956)) has a very high expansion rate, the proportions (after Collignon) being: D Wb Wh Wb:Wh U 270,0 160,0(58) 136,0(48). 1,18 82,0(30) Given a larger sample it might prove possible to demonstrate that A. (Ar.) besairiei is also within the range of variation of A. (Ar.) depereti, but the authors maintain the species separate at this time whilst acknowledging the fine division between them. Occurrence As defined above, Ammonoceratites (Argonauticeras) depereti is known from the Upper Aptian of the Caucasus, Bulgaria, southern France, California, Mozambique and Zululand, and the Albian of Madagascar. Ammonoceratites (Argonauticeras) aff. depereti (Kilian, 1892) Figs 38A, 44-46 Material One specimen only, UPG B465, from the Mzinene Formation, Albian III-IV, Mlambongwenya Spruit, northern Zululand. Dimensions D Wb Wh Wb:Wh U UPRG B465*.5 052% 47,5 23,5(49) 21,0(44) 12 — (—) 84,0 40,0(48) 38,5(46) 1,04 — (—) 149,0 68,5(46) 67,5(45) 1,02 — (—) 248,0 99,5(40) 98,5(40) 1,01 — (—) 352,0 127,5(36) 134,0(38) 0,97 ~—«135,0(38) 316 ANNALS OF THE SOUTH AFRICAN MUSEUM OL IOT yy Yl Fig. 44. Ammonoceratites (Argonauticeras) aff. depereti (Kilian, 1892). UPG B465. x OLS: CRETACEOUS FAUNAS FROM SOUTH AFRICA B17 Yj YY Fig. 45. Ammonoceratites (Argonauticeras) aff. depereti (Kilian, 1892). UPG B465. x 0,5. Description The specimen is beautifully preserved, retaining part of the original shell. All but the last quarter whorl are septate. Coiling is very evolute with the whorls increasing gradually in height and width. The dorsal area of impression is very shallow. The whorl section varies with increasing diameter as shown in the table of dimensions. It is initially rect- angular, but with increasing diameter the venter becomes narrower and the whorl height becomes greater than the whorl breadth. Maximum whorl breadth is at the dorsal third of the flanks. The surface is ornamented by numerous thin, 318 ANNALS OF THE SOUTH AFRICAN MUSEUM (NEage \ Fig. 46. Suture of Ammonoceratites (Argonauticeras) aff. depereti (Kilian, 1892). UPG B465. x 3,6. fine ribs. The ribs pass radially over the umbilical wall and then forwards across the flanks with a slight curvature, and straight over the venter. On the venter of one of the inner whorls, where the specimen had been broken, the ribs are minutely crinkled and are connected longitudinally by faint striae. At a whorl height of 63 mm there are 45 ribs in a distance equal to the whorl height; at 85 mm the figure is 42. On the inner whorls the flanks bear low folds, so that the ribs appear to bunch. On the outer whorls this effect is less noticeable, but two wide and shallow, barely noticeable constrictions occur. The external suture (Fig. 46) is highly divided with narrow-stemmed saddles. The lobes are asymmetrically bifid. Discussion As can be seen from the dimensions, the ratio of Wb/Wh decreases with increase in diameter, a situation somewhat comparable to that encountered in A. (Ar.) depereti, from which it differs in the lack of constrictions on inner whorls (Fig. 38A), as a consequence of which the authors refer to it as A. (Ar.) aff. depereti. It is readily separated from A. (Ar.) besairiei which is markedly depressed at large diameters. Occurrence Middle Albian of northern Zululand. Ammonoceratites (Argonauticeras) besairiei Collignon, 1949 Figs 47-48A Argonauticeras besairiei Collignon, 1949a: 46, pl. 8 (fig. 1); 1950: 36, pl. 5 (fig. 6); 19625: 10, pl. 219-220 (fig. 956). Holotype The original of Collignon (1949a: pl. 8 (fig. 1)), a juvenile specimen from the Lower/Middle Albian transition beds of Ambarimaninga, Madagascar. CRETACEOUS FAUNAS FROM SOUTH AFRICA 319 Material A single fragment, BMNH C78697, from the Mzinene Formation, Albian III, of locality 171, Mlambongwenya Spruit, northern Zululand. Dimensions D Wb Wh Wb:Wh U Holotype (after Collignon, 1949a) 43,0 21,0(49) 18,0(42) LLY 16,0(37) Specimen 1 of Collignon, 1950 . 70,0 35,0(50) 33,0(47) 1,06 26,0(35) Specimen 2 of Collignon, 1950 . 130,0 51,0(50) 48,0(47) 1,06 28,0(27) Original of Collig- non, 19625: pls 219-220 (fig. 956) 200,0 106,0(53) 94,0(47) 1,13 63,0(32) BMNH C78697 __.. — 121,0(—) 107,0(—) (133 — (—) Description The specimen is a wholly septate fragment of a large, massive ammonite whose original diameter must have been well over 200 mm. The coiling is very evolute, with a narrow, shallow impressed area. The umbilicus is of moderate size, and deep. The whorls expand rapidly and are depressed, rounded- trapezoidal, the greatest breadth being well below mid-flank. The umbilical wall is high, rounded, merging with broadly rounded umbilical shoulders and lower flanks. The upper flanks are flattened and converge to a broadly rounded venter. Ornament consists of fine, non-crenulate, narrow, flattened ribs separated by wider interspaces. These arise at the umbilical seam, sweep abruptly back across the lower part of the umbilical wall and curve forwards over the shoulder to pass across the flanks in a rectiradiate or slightly prorsiradiate direction, with a faint convexity. They pass straight across the venter. The external suture is not visible, but the fractured sections indicate the presence of large L and U, lobes, together with a large septal lobe. The dorsum shows the impression of the venter of the penultimate whorl, which was distinctly flattened. Discussion At the diameter represented by the Zululand specimen, the trapezoidal whorl section of Argonauticeras is lost, but when compared with the series of specimens of A. (Argonauticeras) besairiei figured by Collignon (1949a—19625), the present specimen is clearly an adult of that species. Occurrence Upper Aptian to Middle Albian of Madagascar, Middle Albian of Zululand. 320 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 47. Ammonoceratites (Argonauticeras) besairiei Collignon, 1949. BMNH C78697. x 0,7. CRETACEOUS FAUNAS FROM SOUTH AFRICA SAI Genus Protetragonites Hyatt, 1900 Type species Ammonites quadrisulcatus d’Orbigny, 1841. Diagnosis | Very evolute lytoceratids with circular to oval whorl sections, sparse to frequent radial or curved constrictions and a shell ornamented by growth striae only. Discussion Repeated attempts have been made to justify a separation of a family Protetragonitidae Spath, 1927, from the Lytoceratidae. This has been based on the tendency for species to develop one or more auxiliary elements in the suture line. Schindewolf (1960: 681) and Wiedmann (1962: 17) have both demonstrated that Protetragonites follow the same sutural development as the Lytoceratidae, and are derived from Lytoceras sensu stricto. The authors further follow Schindewolf and Wiedmann in regarding Hemitetragonites Spath, 1927, and Leptotetragonites Spath, 1927, as subjective synonyms. Occurrence Protetragonites is a typically Mesogean genus, known chiefly from the Mediterranean region, but also known from central Europe (Austria), the Crimea, Caucasus, north Africa, Madagascar, and now Zululand. Its strati- graphic range is from Tithonian to Upper Albian. Protetragonites aeolus aeolus (d’Orbigny, 1850) Fig. 48B—F Ammonites aeolus @’Orbigny, 1850: 125. Lytoceras (Gaudryceras) aeolus Jacob, 1908: 14 (pars), pl. 1 (fig. 17), non pl. 1 (figs 14-16). Gaudryceras aeolus: Fallot, 1920: 12. Collignon, 19495: 50. Hemitetragonites aeolus Breistroffer, 1936: 175. Breistroffer & Villoutreys, 1953: 71 (Pars): Protetragonites aeolus aeolus Wiedmann, 1962: 24, pl. 10 (fig. 3), text-fig. 6. Protetragonites aeolus: Wiedmann & Dieni, 1968: 31. Type The lectotype, designated by Wiedmann (1962: 25), is MNHP 5770 in the D’Orbigny Collection, Muséum d’Histoire Naturelle, Paris, and is from the Middle? Albian of Escragnolles, Var, France. Material Two specimens, SAS PJR/1—2, from the Mzinene Formation, Albian V, at locality 65 along the Munywana Creek, east of Hluhluwe, Zululand, are definitely referred to the subspecies, whilst a third specimen, BMNH C78699, from the same horizon at locality 56 on the Mzinene River, may also belong here. 322 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 48. A. Ammonoceratites( Argonauticeras) besairiei Collignon, 1949. BMNH C78697. x 0,7. B-F. Protetragonites aeolus aeolus (d’Orbigny). B—C. PJR/1. D-F. BMNH C78699. B-C, x 1; Ds <2: 5E-F alk CRETACEOUS FAUNAS FROM SOUTH AFRICA 323 Dimensions D Wb Wh Wb:Wh U Lectotype, MNHP SHOW : . % 1 31,0 12,5(40) 10,5(34) 1,19 14,5(47) RMB ee 30,7 12,5(41) 8,7(28) 1,43 14,3(47) BMNH C78699 (approximate) . 30,1 — (—) 9,3(31) — 15,0(50) Description . The coiling is very evolute, serpenticone, with a very wide umbilicus (47-50% of diameter). The whorl section is depressed, reniform, with a gently rounded venter and swollen flanks (the greatest breadth is below mid-flank). The dorsal impressed zone is narrow and shallow. All specimens are corroded and no external shell surface survives except in a few areas which retain a suggestion of growth striae. There are four straight, prorsiradiate constrictions per whorl, bounded aperturally by a thickened rib. The sutures are undecipherable. Discussion Relative proportions, expansion rate, position and form of constrictions indicate these specimens to be Protetragonites, most closely resembling the well-known species Protetragonites aeolus (d’Orbigny). Wiedmann (1962: 23) recognized three subspecies, P. aeolus aeolus, P. aeolus neptuni Wiedmann (1962: 29, pl. 1 (fig. 2), text-fig. 7b), and P. aeolus aeoliformis (P. Fallot) (Wiedmann 1962: 26, pl. 1 (fig. 1), pl. 2 (fig. 4), text-fig. 4a). P. aeolus aeoliformis has a more or less rounded whorl section rather than reniform as in the nominate subspecies, whilst P. aeolus neptuni has strongly ornamented inner whorls and much more prominent constrictions. Of other species, Protetragonites crebrisulcatus (Uhlig) (Wiedmann 1962: 19, pl. 1 (fig. 3), pl. 3 (figs 2, 4)) has rather similar overall proportions, but bears more constrictions which efface at maturity. P. obliquestrangulatus (Kilian) and its subspecies (Wiedmann 1926: 21 et seg.) bear many more prorsi- radiate constrictions, whilst Protetragonites laevis Wiedmann (1962: 31, pl. 10 (fig. 4) text-figs 9-10) has a subquadrate depressed whorl section and is smooth. Occurrence Protetragonites aeolus and its subspecies are best known from the western Mediterranean, i.e. the Balearics, Sardinia and south-eastern France, but there are also records from Madagascar, and now Zululand. The species is restricted to the Albian. SYSTEMATIC POSITION UNCERTAIN Genus Pictetia Uhlig, 1883 Type species Crioceras astierianus d’Orbigny, 1842. 324 ANNALS OF THE SOUTH AFRICAN MUSEUM Diagnosis Loosely coiled with whorls separated throughout; body chamber may straighten. Whorl section rounded, compressed to depressed; ornamented by weak, irregular, feebly crinkled ribs. Discussion Pictetia is an enigmatic genus, known chiefly from septate fragments only which are either curved or straight. Few good illustrations of the type species are available and in consequence the authors reillustrated (Fig. 49) the original of D’Orbigny (1842: pl. 115 (figs 3-5)) which is herein designated lectotype of P. astieriana (the specimen has been commonly referred to as the holotype, but D’Orbigny clearly states that he was aware of specimens in the Astier, Duval and Puzos collections, although that which he figured was ‘a lévé tous mes doutes a été découverte par M. Astier’. The authors also take the opportunity of illustrating two of the straight fragments, doubtful paralectotypes, also from Escragnolles, Var (Figs 50-51). In his generic diagnosis of Pictetia, Casey (1960: 3) indicated that it had a ‘suture line with simple lobe formula IU LE, but highly complex’. Such a formula is not typical of the Lytoceratina, which are characterized by a quin- quelobate suture. Unfortunately there is no detailed work on the sutural ontogeny of Pictetia; Schindewolf (1961: 678) examined specimens of the type species, and concluded that the small incision in saddle I/U should be regarded as an incision, and not a lobe. If this be true, then Pictetia is either to be trans- ferred to the Ancyloceratina (some members of which have fimbriate ornament), as did Schindewolf and Hyatt (1900: 588) before him, or retained within the Lytoceratida as an uncoiled Lytoceras (Casey 1960: 3) in which the quadrilobate suture is secondarily derived from quinquelobate ancestors. In Spiroceras bifurcati (Quenstedt) Schindewolf (1961: 762, text-fig. 45) demonstrated that two of the original five lobes became so reduced that a pseudo-trilobate condition prevailed in adults; might not reduction in Pictetia be a similar reflection of acquisition of the heteromorph habit? Unfortunately the Zululand material does not help in the resolution of this interesting problem. Occurrence Pictetia is known from the Lower and Middle Albian of western Europe, Transcaspia, India, Madagascar and Zululand; Collignon (19626: 13) has recorded two species from the Upper Aptian of Madagascar. Pictetia aff. depressa (Pictet & Campiche, 1861) | Figs 52A-C, 53 Compare: Crioceras depressus Pictet & Campiche, 1861: 28, pl. 45 (figs 3-4). Pictetia depressa Casey, 1960: 5, pl. 1 (fig. la—c), text-fig. la—d (with synonomy). Collignon, 19626: 13, pl. 221 (fig. 457). 35) CRETACEOUS FAUNAS FROM SOUTH AFRICA "TX °€669p HNING ‘2d410399T “(78 T “AUSIQIO.P peste ye ) DUDLAAIISD DIAN “6p “314 326 ANNALS OF THE SOUTH AFRICAN MUSEUM Ge B Fig. 50. Pictetia astieriana (d’Orbigny, 1842). BMNH C5440, wholly septate ancyloceratid- like shaft. x 1. CRETACEOUS FAUNAS FROM SOUTH AFRICA a0) A B Fig. 51. Pictetia astieriana (d’Orbigny, 1842). BMNH C5439, septate straight shaft. x 1. 328 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 52. A-C. Pictetia aff. depressa (Pictet & Campiche, 1861). SAS LJE 173. x 1. D-F. Pictetia depressa (Pictet & Campiche, 1861). Copies of Pictet & Campiche’s original figures of the species. CRETACEOUS FAUNAS FROM SOUTH AFRICA 329 Fig. 53. Pictetia aff. depressa (Pictet & Campiche, 1861). Suture of SAS LJE 173. x 3,6. Material One specimen only, SAS LJE 173 from the Mzinene Formation, Albian II-III, locality 171, Mlambongwenya Spruit, northern Zululand. Description The fragment is wholly septate, consisting of three-quarters of a whorl only of a juvenile specimen. Coiling is very open, the whorl section depressed, reniform, with a very small dorsal impressed zone. Ornament is subdued, and consists of single, low prorsiradiate folds on the shell surface. The suture line is partially exposed, showing a long E/L saddle with a narrow stem, a large L and low L/U saddle. I is trifid (Fig. 53). Discussion Depressed whorl section, together with the presence of a narrow dorsal impressed area place this specimen closest to Pictet & Campiche’s species, the original illustrations of which are reproduced here as Figure 52D-—F; the whorl section is more depressed in that species, whilst internal moulds are smooth, in consequence of which the authors refer to their specimen as P. aff. depressa. In P. astieriana (Fig. 49) the whorl section is more rounded; in P. crassecostata Collignon (1963: 8, pl. 243 (fig. 1046)) the whorl section is oval with broad, well-differentiated convex ribs; in P. ovalis Collignon (1963: 8, pl. 243 (fig. 1045)) the whorl section is also oval, with dense, relatively prominent flexuous ribs. P. arcuata Collignon (19626: 13, pl. 221 (fig. 958)) is an Aptian species based on gently curved fragments with a circular whorl section, smooth surface and folds corresponding to the site of individual sutures. 330 ANNALS OF THE SOUTH AFRICAN MUSEUM Occurrence Pictetia depressa is best known from the Lower Albian of western Europe and Transcaspia, although also recorded from the Upper Aptian of Madagascar. ACKNOWLEDGEMENTS We thank Dr H. W. Ball, Dr M. K. Howarth and Mr D. Phillips of the British Museum (Natural History), Dr R. Casey and Mr C. J. Wood (Institute of Geological Sciences), Dr J. Sornay (Muséum d’Histoire Naturelle, Paris), Professor T. Matsumoto (Kyushu, Japan), Dr I. Hayami (Tokyo), Professor R. A. Reyment (Uppsala) and Professor D. Visser (Pretoria) for allowing us to study material in their care and for assistance in many other ways. We also thank General M. Collignon (Moirans), Professor Dr J. Wiedmann (Tubingen), Mr C. W. Wright and Dr M. R. Cooper (Oxford) for their advice and useful discussions. Dr J. A. Jeletzky of the Canadian Geological Survey kindly supplied casts of Ammonoceratites crenocostatus and Professor R. A. Reyment (Uppsala) provided photographs of the holotype of Argonauticeras belliseptatum for this publication. The help of the staff of the Geological Collections, Oxford University Museum; Geological Survey of South Africa, Pretoria; and South African Museum, Cape Town, is gratefully acknowledged, as is the financial support of the Trustees of the Sir Henry Strakosch Bequest, the Royal Society, and the Natural Environment Research Council. Thanks are due to the Director, Geological Survey of South Africa, Pretoria, for the loan of part of the material described. REFERENCES ANDERSON, F. M. 1902. Cretaceous deposits of the Pacific Coast. Proc. Calif. Acad. Sci. 2: 1-132. ANDERSON, F. M. 1938. Lower Cretaceous deposits in California and Oregon. Spec. Pap. geol. Soc. Am. 16: 1-339. ANTHULA, D. J. 1899. 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Palaeontogr. Soc. (Monogr.): 1-787. SPATH, L. F. 1927. Revision of the Jurassic Cephalopod fauna of Kachh (Cutch). Mem. geol. Surv. India Palaeont. indica n.s. 11: 1-71. SPATH, L. F. 1939. The Cephalopoda of the Neocomian Belemnite Beds of the Salt Range. Mem. geol. Surv. India Palaeont. indica n.s. 25: 1-154. STOLICZKA, F. 1863-1866. The fossil Cephalopoda of the Cretaceous rocks of southern India. Mem. geol. Surv. India Palaeont. indica 1: 41-56 (1863); 2-5: 47-106 (1864); 6-9: 107-154 (1865); 10-13: 155-216 (1866). Suess, E. 1865. Uber Ammoniten. Sitz.-Ber. k.k. Akad. Wiss. 52: 71-89; 305-322. THOMEL, G. 1968. A propos de l’Ammonoceratites (Argonauticeras) depereti (Kilian) du Gargasien des Basses-Alpes orientales. Bull. Soc. géol. Fr. (7) 10: 684-687. UHLIG, V. 1883. Die Cephalopoden der Wernsdorfer Schichten. Denkschr. Akad. Wiss., Wien 46 (2): 127-290. VASICEK, Z. 1972. Ammonoidea of the Tésin-Hradisté Formation (Lower Cretaceous) in the Moravskoslzské Beskydy Mountains. Ustredniho ustavu geologického 38: 1-103. WEDEKIND, R. 1916. Uber Lobus, Suturallobus und Inzision. Zentbl. Miner. Geol. Paldont. 1916: 185-195. WHITEAVES, J. F. 1876. On some invertebrates from the coal-bearing rocks of the Queen Charlotte Islands. Mesozoic Fossils 1 (1): 1-92. Ottawa: Geological Survey of Canada. WHITEAVES, J. F. 1884. On the fossils of the coal-bearing deposits of the Queen Charlotte Islands collected by Dr G. M. Dawson in 1878. Mesozoic Fossils 1 (3): 191-262. Ottawa: Geological Survey of Canada. WHITEAVES, J. F. 1900. On some additional or imperfectly understood fossils from the Cretaceous Rocks of the Queen Charlotte Islands, with a revised list of the species from these rocks. Mesozoic Fossils 1 (4): 263-308. WIEDMANN, J. 1962. Unterkreide-ammoniten von Mallorca J. Liefr. Lytoceratina, Aptychi. Abh. math.-naturw. Kl. Akad. Wiss. Mainz 1962 (1): 1-148. WIEDMANN, J. & DIENI, I. 1968. Die Kreide Sardiniens und ihre Cephalopoden. Palaeontogr. ital. 64: 1-171. YABE, H. 1903. Cretaceous Cephalopoda from Hokkaido. Part 1. J. Coll. Sci. imp. Univ. Tokyo 18: 1-55. ZITTEL, K. A. VON. 1868. Die Cephalopoden der Stramberger Schichten. Palaeont. Mitt. Mus. k. bayer. St. 2: 1-118. ZITTEL, K. A. VON. 1884. Handbuch der Palaeontologie. (Palaeozoologie). i Abt., ii Band, Lief. ili. Cephalopoda. Munich & Leipzig: Oldenbourg. ZWIERZYCKI, J. 1914. Die Cephalopoden-fauna der Tendaguru-schichten in Deutsch-Ostafrika. Arch. Biontol. 3: 7-96. ’ “bts bie I P ‘ 1 he eh a ane ) end 4 6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature (particularly Articles 22 and 51). 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Spawning behaviour, ee ta and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): THIELE, J. 1910. Mollusca: B. aha Re Ee Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. (continued inside back cover) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 74 ~ Band March 1978 Maart Part 13 #Deel RE-EXAMINATION OF ARCHAEICHNIUM, A FOSSIL FROM THE NAMA GROUP By MARTIN F. GLAESSNER Cape Town Kaapstad The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular intervals as material becomes available Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000 Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM word uitgegee in dele op ongereelde tye na beskikbaarheid van stof Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 8000 OUT OF PRINT/UIT DRUK 1, 201-3, 5-8), 3(1-2, 4-5, 8, t.-p.i.), 51-3, 5, 7-9), 6(1, t.—p.i.), 711-4), 8, 911-2, 7), 10(1-3), 11(1-2, 5, 7, t--p.i.), 15(4-5), 24(2), 27, 31(1-3), 32(5), 33 Price of this part/Prys van hierdie deel R1,80 Copyright enquiries to the South African Museum Kopieregnavrae na die Suid-Afrikaanse Museum ISBN 0 908407 42 4 Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms:, Bpk., Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap RE-EXAMINATION OF ARCHAEICHNIUM, A FOSSIL FROM THE NAMA GROUP By MARTIN F. GLAESSNER University of Adelaide, South Australia (With 3 figures) - [MS. accepted 6 September 1977] ABSTRACT The fossils described by Haughton in 1960 as Archaeocyatha from the Nama Group were later, after the study of plaster casts, considered as worm tubes or worm burrows by the present author and named Archaeichnium haughtoni. The type specimens have now been re-examined. They consist of agglutinated sand grains and are similar to tubes made actively by living polychaete worms. They are not closely similar to the partly calcareous tubes of Cloudina or to the distinctively structured Volborthella. In the absence of any taxonomically significant characters of tube construction the systematic position of the originator of these fossils remains uncertain. CONTENTS PAGE Introduction=." 2" cas see 335 New observations . © 2... 338 INntenpretaliOniass ewan tee ee SS Conclusions = «bh ees 341 Acknowledgements. . . . . 342 References: 2) Ge) haere 342 INTRODUCTION Haughton (1960) described what he considered “An archaeocyathid from the Nama System’, from two specimens (housed in the South African Museum) of a quartzitic sandstone on Griindoorn near the Ham River, ‘a typical member of the Kuibis beds’. Germs later re-examined the stratigraphy of the Nama Group. From his brief description of the eastern area it appears that Haughton’s fossils came from the Upper Clastic Member of the Nababis Formation, probably corresponding to the Upper Clastic Member of the Kuibis Formation in the west (Germs 1972a: 19, figs 14, 22). The interpretation of these fossils hinges on their mode of fossilization. It is therefore necessary to summarize Haughton’s views on this matter. He considered them as moulds on bedding planes and noted correctly that some of the supposed moulds are ‘marked off’ by a bordering depression and that some are lying parallel to each other, as if deposited by current action. They are described as elongated tubular cones with circular cross- sections. “The more complete individuals have a pointed lower end.’ In two 335 Ann. S. Afr. Mus. 74 (13), 1977: 335-342, 3 figs. 336 ANNALS OF THE SOUTH AFRICAN MUSEUM instances, ‘two individuals appear to have been joined at this pointed end’. Haughton noted also the presence of a number of elongate (longitudinal) ‘pustulate ridges’ and of an individual ‘mould’ appearing as an approximately circular ridge which ‘can be interpreted as an infilling of the intervallum or space between the internal and external walls of a cup, or as a replacement of a single thick exoskeleton’. The second alternative was abandoned without further com- ment and the first one adopted. Thus the longitudinal section (as seen below the centre of Fig. 1) was interpreted as showing an infilling of a central cavity sur- rounded by concentric inner and outer walls which are now indicated by depressions. The solid specimens being considered as internal moulds, the present longitudinal ridges on their outer surfaces were thought to separate what would have been broad, rounded ridges on the inner surfaces of the vanished outer walls. If pustules are now present, they would have originated as rows ‘of evenly spaced pores’ in longitudinal channels. This reconstruction of the fossils from their supposed ‘moulds’ led Haughton without much difficulty to his interpreta- tion as Archaeocyatha. The obvious significance of Haughton’s classification of his fossils for the dating of the Nama Group led to their re-examination (Glaessner 1963). Those studies were based on plaster casts of the originals. It appeared that the fossils represented infillings of cemented tubes made, probably, by worm-like animals which incompletely agglutinated the sediment surrounding their burrows. The tubes were considered as generally cylindrical but they appeared to be conical where they entered the rock matrix. The specimen (below the centre in Fig. 1, marked M in Glaessner 1963, pl. 3 (fig. 1)) which Haughton saw as two indivi- duals joined at their pointed ends, represents one tube dipping from the bedding plane into the rock matrix and re-emerging with a 90° change in direction. The other specimen, marked with an arrow on Haughton’s Plate 4 (L in Glaessner 1963, pl. 3 (fig. 1)), shows a similar change of direction. The number of longitu- dinal ridges was correctly described by Haughton as ten to twelve on each individual. Five to six ridges can be seen on one side but the supposed pustules are sand grains. The ridges are also visible on several fragmentary specimens but appear only faintly at the ends of the largest specimen. The longitudinal ridges on the outer surfaces of the agglutinated tubes and the transverse layering of the 1 mm thick walls appear to be diagnostic for these fossils. None of the distinctive characters of the Archaeocyatha can be seen in them. There are no pointed cones, no traces of a former calcareous double wall, no septa or other intervallum structures. There is no evidence of an intervallum. The description of the partly agglutinated tubes of Cloudina by Germs (1972a, 1972b) and recent studies of this genus and of the fully agglutinated Volborthella (Glaessner 1976) suggested that a re-examination of Archaeichnium would be desirable, not because they could still be considered as Archaeocyatha or as identical with Cloudina, as subsequently suggested by Kaever & Richter (1976: 30), but because more is now known about early tube-building worms. The observations of the morphology of these fossils made previously on plaster RE-EXAMINATION OF ARCHAEICHNIUM, A FOSSIL FROM THE NAMA GROUP 331 Fig. 1. Archaeichnium haughtoni Glaessner, 1963, on a slab of quartzite from the Nama Group (Late Precambrian). Slab B, SAM-K4812. The holotype is the specimen below the centre of the figure. Natural size. 338 ANNALS OF THE SOUTH AFRICAN MUSEUM casts have been supplemented by examination of the original specimens. One of them is marked Slab A, SAM-K4813, referred to correctly as Slab A by Haugh- ton (1960) on pages 57 and 58, but as Slab B on plate 3 (true magnification x 0,9, not ‘above natural size’). The other is marked Slab B, SAM—K4812, which is correctly referred to by Haughton on pages 57 and 58 and on plate 5, but as Slab A on plate 4. The true magnification of this plate is about x 1,5, not x 5/8. Part of a plaster cast of Slab A was previously figured (Glaessner 1963) as plate 3, figure 2, and part of Slab B as plate 3, figure 1 and they are now shown as Figures | (Slab B) and 2 (Slab A). NEW OBSERVATIONS The fossils consist of firmly cemented sand grains, mostly quartz. Although the slabs could not be cut and without sections no measurements could be made, the grains forming the tubes appear to be slightly coarser and more uniform than the matrix. They must have been agglutinated prior to fossilization as they are mostly sharply delimited from the matrix and do not grade into it on the bedding planes. There is no indication of any previous presence of thin ‘walls’ between the matrix and the fossils. The rock slabs consist of a slightly glassy quartzite. Diagenetic alteration of a sandy layer has fused the tubes with the sediment represented by the slabs, leaving them projecting in sharp semirelief from one bedding plane. On the analogy of similarly preserved Australian fossils collected in situ, this bedding plane was probably underlain by a silt or clay layer. The preservation is likely to be in hyporelief. On Slab A (Fig. 2) and possibly also on Slab B (Fig. 1) where numbers are smaller, the tubes show (as Haughton had recognized) a preferred orientation, indicating some current sorting. This is demonstrated by Figure 3. Sizes of the tubular fossils range from | mm to almost 5 mm in diameter. There is some flattening and some apparent twisting of tubes. Some are cylindrical, some appear to be barrel-shaped, some were probably conical and a few, including one near the bottom of Figure | (L in pl. 3 (fig. 1) in Glaessner 1963) thin abruptly at one end to form a long, thin, twisted tail. Irregular forms are more common than geometrically regular ones. Some ends are sharp, others fade out in the matrix. Few if any clean breaks which would indicate brittleness are visible and it is concluded that the tubes were Cea sible and flexible, particularly where their walls were thin. INTERPRETATION In 1963 it was demonstrated that the fossils could not be considered as Archaeocyatha. What are these fossils? Specimen M (Fig. 1, below centre) was selected as the holotype of Archaeichnium haughtoni because it shows the most distinctive characters of the new monotypic genus which are seen less clearly among the other fossils on the two slabs. The holotype specimen can be followed in appropriate lighting conditions over a length of 70 mm, almost to the edge of RE-EXAMINATION OF ARCHAEICHNIUM, A FOSSIL FROM THE NAMA GROUP 339 Fig. 2. Archaeichnium haughtoni Glaessner, 1963. Slab A, SAM-—K4813. (The lobe projecting from the lower part of the bedding plane may be a bioturbated load cast.) Natural size. 340 ANNALS OF THE SOUTH AFRICAN MUSEUM fo) ° ° Clo 0. 0 20° or 20 AN 60 B80 oO" Fig. 3. Histogram of positions of 73 individual fossils on Slab A showing deviation of long axis from modal direction. the slab. Its maximum width is almost 5 mm. Its wall is about 1 mm thick and shows pronounced transverse layering of sand grains. The centre of the tube is filled with sand grains which show a much fainter and less regular transverse fabric. The outer surface of the tube is marked with fine longitudinal ribs, but they are visible in this specimen only faintly near the ends of the tube, which is either smoothed by weathering or split open for most of its length. At the end nearer the edge of the slab the ribs are preserved as impressions in the matrix, hence they are an external sculpture, not part of an internal mould.-In the other large specimen (bottom of Fig. 1, L in pl. 3 (fig. 1) in Glaessner 1963) the external ribs are well preserved and straight but its internal structure is not revealed. This tube is 35 mm long and up to 4,5 mm wide. At one end it is narrowed and appears twisted. Several other specimens show some of these characters such as external longitudinal ribbing or transverse wall fabric. Their size ranges fit those of the larger specimens and the indications of original flexibility are similar. On these grounds the author sees no reason to distinguis] any specimens taxonomically from the typical ones. It is difficult to decide whether these fossils are burrows in the strict sensi of the word. There is no doubt that they were built by a worm-shaped animal from sand grains selected and agglutinated by some organic cement which allowed them some flexibility. The clear separation between the tube walls and the matrix brought out by weathering of the bedding plane indicates that the composition of the walls was different from that of the matrix. The wall material appears to have been actively selected by the animal. There is no evidence for calcareous walls having been dissolved during fossilization. The selection of grains and their arrangement so as to form external and possibly strengthening ridges shows active building from transported grains, rather than passive cemen- RE-EXAMINATION OF ARCHAEICHNIUM, A FOSSIL FROM THE NAMA GROUP 341 tation of a burrow wall from grains of matrix in situ. Faint transverse texture of the infilling of the tube in the holotype specimen may indicate active filling, but there is no confirmation from other specimens. It is likely that the tubes were made by organisms burrowing in or living on the sediment and constructing agglutinated walls from sand grains. It is also probable that subsequent current action as indicated particularly by the orientation of burrow fragments on Slab A washed the weakly cemented burrows or tubes out of a layer of sediment and distributed them over a bedding plane. If this were the case, it raises the semantic point whether we are dealing with trace fossils in the strict sense of the word or whether worm tubes agglutinated from sand grains with organic cement are properly considered as body fossils, but this is not a matter of substantial importance. The Slabs A and B are so similar that they may well represent not very distant parts of the same bedding plane, yet one (A) indicates current action while the other (B) may be thought to have at least the two curved, long tubes preserved in situ. These questions cannot be solved without further field studies and discovery of material in situ. CONCLUSIONS Archaeichnium haughtoni is a name given to tubular fossils with walls agglutinated from small detrital quartz grains. The best preserved of these tubes show external, longitudinal, straight, fine ribs and a transverse layering of the grains forming the wall. They may have been actively infilled. Most of the tubes seem to be fragments washed out of the original sediment within or from which they were built. They appear to have been redeposited on bedding planes where they were subsequently buried by layers of sand and diagenetically fused with it. There is no significant resemblance between this fossil and others from the Nama Group or indeed from anywhere else. After this study had commenced, Kaever & Richter (1976) described Buschmannia roeringi from the Buschmannsklippe Formation of the Nama Group as Archaeocyatha. The author was able to re-examine this fossil. Its resemblance to Praesolenopora Tsao & Zhao, 1972, from the Late Precambrian (Upper Sinian) of China (see Tsao & Liang 1974) was noted and it was concluded that it is probably a calcareous alga. Archaeichnium differs from Cloudina Germs from the Nama Group (Germs 1972a, 19726). This genus builds partly calcareous, secreted tubes with only minor, external agglutination of sand grains and transverse sculpture. ‘The tube walls seem to have contained organic material. Their distortions suggest some flexibility ... reminiscent of some agglutinating foraminifera with walls of quartz or carbonate grains and much organic cement’ (Glaessner 1976: 266). Cloudina is generally placed in the Order Cribricyathida, most of which are of early Cambrian age. Their morphological characters make probable their ‘possible relation to the serpulids’ (Germs 1972b: 753) and suggest that they are ‘best classified as an extinct Order of the Class Polychaeta’ (Glaessner 1976: 268). Archaeichnium differs from Volborthella which is built from invaginated cones 342 ANNALS OF THE SOUTH AFRICAN MUSEUM of agglutinated sandy material round a central tube in distinctly sloping layers, and from the similar but predominantly calcareous Sal/terella. This mode of tube construction is similar in some detail to that used by living Sabellariidae. Volborthella can be considered as the product of polychaete annelid worms. While the tubes and the inferred tube-building activities of Archaeichnium are comparable and compatible with those of polychaete worms of the Order Tere- bellida (and probably others), there is as yet nothing sufficiently specific in the known morphological characters of this fossil to support its definite systematic assignment to the Annelida Polychaeta. It can only be tentatively attached to them and must remain for the present incertae ‘sedis. No occurrences of this fossil other than that at its type locality, in strata considered to be of Late Precambrian age, are known. ACKNOWLEDGEMENTS I wish to thank the authorities of the South African Museum for the loan of the type material for this study; the support of this loan by Mr P. J. Rossouw, South African Geological Survey, Pretoria, is gratefully acknowledged. Through the kind co-operation of Prof. M. Kaever I was able to re-examine Buschmannia roeringi. Mr Richard Barrett, Department of Geology, University of Adelaide, took the photographs of the rock specimens. REFERENCES GERMS, G. J. B. 1972a. The stratigraphy and paleontology of the lower Nama Group, South West Africa. Bull. Chamber Mines Precambr. Res. Unit 12. GERMS, G. J. B. 1972b. New shelly fossils from the Nama Group, South West Africa. Am. J. Sci. 272: 752-761. GLAESSNER, M. F. 1963. Zur Kenntnis der Nama-Fossilien Stidwest-Afrikas. Annln naturh. Mus. Wien 66: 133-120. GLAESSNER, M. F. 1976. Early Phanerozoic annelid worms and their geological and biological significance. J. geol. Soc. Lond. 132: 259-275. HAUGHTON, S. H. 1960. An archaeocyathid from the Nama System. Trans. R. Soc. S. Afr. 36: 57-59. KAEVER, M. & RICHTER, P. 1976. Buschmannia roeringi n. gen. n. sp. (Archaeocyatha) aus dei Nama-Gruppe Siidwestafrikas. Paldont. Z. 50: 27-33. Tsao, RuI-CuHi & LIANG, YU-ZHOU. 1974. On the classification and correlation of the Sinian System in China, based on a study of algae and stromatolites. Mem. Nanking Inst. Geol. Pal. 5: 1-26. —1 \ Boegicny,” the ° = Vee ed f ae ir 7 6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature (particularly Articles 22 and 51). Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov., etc. An author’s name when cited must follow the name of the taxon without intervening punctuation and not be abbreviated; if the year is added, a comma must separate author’s name and year. The author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is transferred from its original genus. The name of a subsequent user of a scientific name must be separated from the scientific name by a colon. Synonymy arrangement should be according to chronology of names, i.e. all published scientific names by which the species previously has been designated are listed in chronological order, with all references to that name following in-chronological order, e.g.: Family Nuculanidae Nuculana (Lembulus) bicuspidata (Gould, 1845) Figs 14-15A Nucula (Leda) bicuspidata Gould, 1845: 37. Leda plicifera A. Adams, 1856: 50. Laeda bicuspidata Hanley, 1859: he pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b). Nucula largillierti Philippi, 1861: Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. Note punctuation in the above example: comma separates author’s name and year semicolon separates more than one reference by the same author full stop separates references by different authors figures of plates are enclosed in parentheses to distinguish them from text-figures dash, not comma, separates consecutive numbers Synonymy arrangement according to chronology of bibliographic references, whereby the year is placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable. In describing new Species, One specimen must be designated as the holotype; other speci- mens mentioned in the original description are to be designated paratypes; additional material not regarded as paratypes should be listed separately. The complete data (registration number, depository, description of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.: Relaivea: SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973. Note standard form of writing South African Museum registration numbers and date. 7. SPECIAL HOUSE RULES Capital initial letters (a) The Figures, Maps and Tables of the paper when referred to in the text e.g. *... the Figure depicting C. namacolus ...’; °. . . in C. namacolus (Fig. 10) . (b) The prenines of prefixed surnames in all dames: hen used in the text, if not precicd by initials or full names e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene (c) Scientific names, but not their vernacular derivatives e.g. Therocephalia, but therocephalian Punctuation should be loose, omitting all not strictly necessary Reference to the author should be expressed in the third person Roman numerals should be converted to arabic, except when LONE part of the title of a book or article, such as ‘Revision of the Crustacea. Part VIII. The Amphipoda.’ Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial capital letter, provided the same generic name is used consecutively. Name of new genus or species is not to be included in the title: it should be included in the abstract, counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts. MARTIN F. GLAESSNER RE-EXAMINATION OF ARCHAEICHNIUM, A FOSSIL FROM THE NAMA GROUP r iy f i. VCR a Ne Ave PAY Ce Lary MCZ ERNST MAYR LIBRARY (ONO 8 444 064 wh Ee a Oe " My sh = 2 ute Bike RQe var b Es s : ‘ crys oe 4 ; : ? ¢ en alia " . ae tg : ws 7 Biles 4B ps oe Sacome te ih Se Seah pasar tee a eer is oo bo bee aol ee. we BAG Te ke Ce : ae ; . - a = : eng™ . as ‘ ; ; : 7 sa i . a ‘ j : eer : : f eee 3 a te i :- Ned > - mane aoe - dais ls . oe : - : hd Jie 6 ¢ Sic ie x é a ; : aoe - : a A « a : — as ~ . - é : 7 - a . » + < meee “ . ame * ‘i > 7 iy ‘- ¥ i is of Bs o - ee, = . 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