HARVARD UNIVERSITY e Library of the Museum of Comparative Zoology ae «wir ae’ A vi | vat. i" | Me ut nea A MUSEUM © RIK AAS ge: UMiy 4 ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM VOLUME 77 BAND THE TRUSTEES OF THE DIE TRUSTEES VAN DIE SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM CAPE TOWN KAAPSTAD 1979 ® SET, PRINTED AND BOUND IN THE REPUBLIC OF SOUTH AFRICA BY THE RUSTICA PRESS (PTY.) LTD., WYNBERG, CAPE 932 4 A $ a oS ANNALS OF THE ANNALE VAN DIE SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM VOLUME 77 BAND 77 LIST OF CONTENTS APPLETON, C. C. The Unionacea (Mollusca, Lamellibranchiata) of south-central Africa. (Published May 1979.) : a: BARNARD, J. L. Redescription of Plioplateia K. H. Barnard, a genus of amphipod (Crustacea) from South Africa. (Published December 1978.) in = we is Cooper, M. R. & KENNEDY, W. J. Uppermost Albian (Stoliczkaia dispar Zone) ammonites from the Angolan littoral. (Published June 1979.) : mac a oe é De Freitas, A. J. A new genus and species of the penaeoid family Solenoceridae (Crustacea, Deca- poda) from south-east African waters. (Published February 1979.) KENNEDY, W. J. see Cooper, M. R. KENNEDY, W. J. & KLINGER, H. C. Cretaceous faunas from Zululand and Natal, South Africa. A new genus and species of Gastroplitinae from the Mzinene Formation (Albian). (Published December 1978.) oh ae + KENNEDY, W. J. & KLINGER, H. C. Cretaceous faunas from Zululand and Natal, South Africa. The ammonite super- family Haplocerataceae Zittel, 1884. (Published February 1979.) KENSLEY, B. The South African Museum’s Meiring Naude cruises. Part 8. Isopoda Anthuridea. (Published October 1978.) 42 oe Shs er KLINGER, H. C. see KENNEDY, W. J. KLINGER, H. C. & Lock, B. E. Upper Cretaceous sediments from the Igoda River Mouth, East London, South Africa. (Published December 1978.) Bes ap oe ae MILLARD, N. A. H. Type specimens of Hydroida aun uraae in the South African Museum. oe lished February 1979.) 2 . Rau, R. E. Additions to the revised list of preserved material of the extinct Cape Colony quagga and notes on the relationship and distribution of southern plains zebras. (Published December 1978.) Page 151 47 175 123 57 85 71 133 27 NEW GENERIC NAMES PROPOSED IN THIS VOLUME Alopecoceras Kennedy & Klinger, 1978 Cryptopenaeus de Freitas, 1979 ‘OLUME 77 sail 1 OCTOBER 1978 ISSN 0303-2515 ae RY DEC 27 1978 UNIVERSES, ANNALS OF THE SOUTH AFRICAN ae MUSEUM CAPE TOWN INSTRUCTIONS TO AUTHORS 1. MATERIAL should be original and not published elsewhere, in whole or in part. 2. 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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 a/. 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. Fiscuer, 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 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. TurELe, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: 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 77 Band October 1978 Oktober Part 1 Deel THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES PART 8 ISOPODA ANTHURIDEA 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 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, 21-3, 5-8), 3(1-2, 4-5, 8, t.-p.i.), 51-3, 5, 7-9), 6(1, t—p.i.), 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), 32(5), 33 Copyright enquiries to the South African Museum Kopieregnavrae na die Suid-Afrikaanse Museum ISBN 0 908407 56 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 THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES PART 8 ISOPODA ANTHURIDEA By Brian Kensley South African Museum, Cape Town (With 13 figures) LMS. accepted 21 March 1978] ABSTRACT Ten species of anthurid and paranthurid isopods are recorded from deep water off Natal. One is described as a new genus, viz. Natalanthura foveolata. A further five species are described as new, viz. Apanthura insignifica, Neohyssura skolops, Colanthura uncinata, Leptanthura minuta, and Leptanthura natalensis. CONTENTS PAGE Introduction . . 3 ; : ; ; 1 Species list . : ; : : : : 1 Systematic discussion . ‘ ; ; : 2 Acknowledgements. : P : ae >, References. : ‘ : 4 , mete ee INTRODUCTION The anthuridean isopods were excluded from the report on the isopods of the 1975 and 1976 Meiring Naude cruises (Kensley 1977) either for lack of specimens of some species or because of taxonomic uncertainty. With the 1977 cruise more anthurids were collected and some of the taxonomic problems have been resolved. The anthurids of all three cruises are presented here. Station data from the first two cruises may be obtained from Louw (1977), and the South African Museum’s unpublished reports and station lists for the 1977 cruise. SPECIES LIST SM Station 3¢ 22 ~—so ovig. 92.—s jus Family Anthuridae Anthelura remipes Barnard ; ; : 123 os l — — 129 — — — Apanthura insignifica sp. nov. . ; : 123 —- 2 — — 129 | 3 oe -- Mesanthura catenula (Stimpson) . 7 86 — 1 — l Natalanthura foveolata gen. et sp. nov. . 86 = 6 os -— 123 — 1 — - 129 —_ _— l _ Neohyssura skolops sp. nov. : : : 129 — 1 — -- Ann. S. Afr. Mus. T7 (1), 1978: 1-25, 13 figs. 2 ANNALS OF THE SOUTH AFRICAN MUSEUM SM Station oo cole) ovig. 22 juvs Family Paranthuridae Colanthura uncinata sp.nov. . ; : 86 103 Leptanthura minuta sp.nov. . : e 78 86 129 Leptanthura natalensis sp. nov. . : ‘ 53 Ww 9 12 — 1 | Paranthura punctata (Stimpson) , 2 103 Pseudanthura tenuis Kensley ‘ é 53 a oO | |] | eerenne | i Oe ok fe ie RF a eI cee pasealel | lent | | SYSTEMATIC DISCUSSION Family Anthuridae Apanthura insignifica sp. nov. Figs 1-2 Description Female Integument thin. Body with following proportions: C<1=2=3<4=5>6>7. Pereonite 7 half length of 6. Pereonites 4 to 7 with faint anterior constriction, pereonite 4 with single, shallow, circular dorsal pit, pereonite 5 with transversely elongate shallow pit, pereonite 6 with two small, shallow pits. Dorsolateral grooves present. Pleonites | to 5 completely separate, pleonite 5 slightly longer than preceding pleonites, pleonite 6 with middorsal incision in posterior margin. Telson distally broadly rounded with several setae, widest at about midlength, with low weakly developed middorsal ridge; two large statocysts present at telsonic base. Cephalon with rostrum projecting beyond anterolateral corners; eyes lacking. Antennular peduncle 4-segmented, basal segment equal to second and third segments together; fourth segment very short, flagellum of two articles bearing three terminal aesthetascs. Antennal peduncle 5-segmented, two basal segments fairly broad; flagellum of four articles. Mandibular palp 3-segmented, first and second segments subequal, third segment shorter, with four distal serrate spines; incisor of three teeth, lacinia plate with five teeth; molar process well developed, distally narrowly rounded. Maxilla elongate, with one strong tooth and five smaller spines distally; inner ramus tipped with single seta. Lower lip complex tipped with single terminal hook-like lobe on each side. THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 3 B G Fig. 1. Apanthura insignifica. A. Holotype in dorsal view. B. Antennule. C. Antenna. D. Mandible. E. Maxilla. F. Lower lip complex. G. Maxilliped. 4 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 2. Apanthura insignifica. A. Pereonite 7 and pleon in lateral view. B. Pereopod 1. C. Pereopod 2. D. Pereopod 7. E. Pleopod 1. F. Telson. G. Uropod. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 5 Maxilliped 5-segmented, terminal segment rectangular with several setae on mediodistal corner. Pereopod | unguis half length of dactylus, with small spine at base; pro- podus broad, palm concave with strong, rounded lobe at about midpoint; carpus triangular, distally narrowly rounded. Pereopods 2 to 7 similar, with carpi barely underriding propodi, and becoming slightly more elongate posteriorly. Pleopod | exopod operculate, about three times wider and slightly longer than the endopod, bearing numerous distal plumose setae; endopod with three distal plumose setae; basis with three retinaculae. Uropodal exopod folding over telson, oval in outline, reaching to end of basis, bearing several sparsely plumose setae; endopod oval, fringed with simple setae. Submale Antennular flagellum of about twelve articles extending posteriorly to second pereonite, lacking whorls of aesthetascs. Pereopod | as in female. Stylet on pleopod 2 not yet detached from endopod. Material Holotype SAM-A15646 92° TL 5,9 mm SM 123 30°33’S 30°48’E 690 m Paratype SAM-A15646 12 TL 5,2 mm SM 123 Paratypes SAM-A15647 1 sub. ¢ TL 5,6 mm 12 TL 4,5 mm SM 129 30°54’S 30°31’E 850 m Paratypes USNM 170542 2 9° TL 5,6 mm 5,4 mm SM 129 Remarks Of the twelve species of Apanthura described, only A. coppingeri Barnard from the Dundas Straits (northern Australia) and A. africana Barnard from South Africa lack eyes. The rhomboid-shaped telson, and the slender uropodal endopod of A. coppingeri easily distinguish this species from A. insignifica. A. africana, known from Saldanha Bay to Still Bay, is a much larger species (up to 20 mm) recorded from shallow water. A. africana can further be separated from A. insignifica by the maxilliped, which possesses a small terminal segment, and an apically acute telson. Etymology The specific name refers to the small size of this species. Natalanthura gen. nov. Diagnosis Integument heavily indurate and very obviously pitted. Mandible with spike-shaped molar on right side, molar reduced to a spine on left side. 6 ANNALS OF THE SOUTH AFRICAN MUSEUM Maxilliped 5-segmented with well developed endite. Carpus of pereopods 4 to 7 slightly underriding propodus. Pleopod | operculate. Pleonites 1 to 3 subequal, separate; pleonites 4 and 5 fused; pleonite 6 indistinguishably fused with telson. Paired statocysts difficult to detect but probably present beneath two pits at telsonic base. Type species Natalanthura foveolata Etymology The generic name Natalanthura is derived from the South African province of Natal, in whose waters the species was caught, plus the usual ‘anthura’ suffix. Gender: feminine. Remarks The unusual fusion of the pleonites is not encountered in any of the described genera of anthurids. The mandibular structure would also seem to be unique, although reminiscent of Panathura. These two features demand the erection of a new genus. Natalanthura foveolata gen. et sp. nov. Figs 3-4 Description Female Integument indurate and obviously pitted. Anterolateral corners of cephalon extending slightly beyond rostrum. Eyes feebly pigmented, of three or four ocelli. Pereonites with strong dorsolateral ridges and two large dorsal pits per segment in addition to numerous smaller pits. Lateral walls of pereonites pitted. Pereonites 1 to 4 increasing in width and length, pereonite 4 widest and longest; posterior three pereonites decreasing in length, seventh half length of sixth. Pereonites 3 to 5 with oblique dorsal grooves converging anteriorly. Pleon with anterior three pleonites free and distinct; pleonite 4 and 5 fused, line of fusion marked by row of pits; pleonite 6 completely fused with telson. Latter longer than pleonites 1 to 5 combined, distal margin evenly convex, dentate, with strong mediodorsal rounded ridge. Antennule with broadly curved basal segment plus six additional segments; fourth segment short, terminal segment bearing setae and three or four aesthetascs. Antenna with 4-segmented peduncle; broad and relatively elongate basal segment with distal triangular part folded over antennule; flagellum of six articles. Mandible with distal incisor tridentate, quite strongly sclerotized; lacinia plate with row of twelve short teeth; right mandible with digitiform molar THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 7 process bearing row of small denticles and striations; molar process reduced to small spine in left mandible; palp 3-segmented, middle segment longest and widest, terminal segment bearing three stout setae. Maxilla slender and elongate, with four terminal teeth. Maxilliped 5-segmented, terminal segment rounded, bearing four setae; endite elongate-lanceolate, apically acute, with single seta on median margin, reaching just beond proximal margin of third segment. See F Fig. 3. Natalanthura foveolata. A. Holotype in dorsal view. B. Antennule. C. Antenna. D. Maxilla. E. Right mandible. F. Maxilliped. 8 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 4. Natalanthura foveolata. A. Pereopod 1. B. Pereopod 7. C. Pereonites 4 and 5 in lateral view. D. Pleopod 1. E. Uropod. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 9 Pereopods | and 2 similar, unguis one-third length of dactylus, with single spine at base; propodus with single distal spine and numerous setules on palmar margin. Posterior pereopods with unguis one-third length of dactylus, with short spine at base, propodus with strong distal spine and four or five short, slender spines on ventral margin, and single serrate seta on upper distal margin; carpus about half length of, and slightly underriding, propodus. Brood pouch formed by four pairs of oostegites. Pleopod | operculate, rami of equal length, endopod half width of exopod, with four distal plumose setae; exopod with numerous plumose setae distally, and with outer surface bearing scales; basis bearing two coupling hooks. Uropodal endopod subequal to basis in length, entire margin strongly dentate; exopod crescentic and curved, margin strongly dentate; basis triangular in cross-section. Material Holotype SAM-A15648 2 TL 5,8 mm SM 86 27°59’S 32°40’E 550 m Paratypes SAM-A15648 2 292 TL 3,5 mm 5,4 mm SM 86 Paratypes USNM-170543 3 92 TL 4,3 mm 4,3 mm 4,6 mm SM 86 Paratype SAM-A15649 1 2 TL3,4mm SM 123 30°33’S 30°48’E 690 m Paratype SAM-A15650 | ovig. 2 TL 6,9 mm SM 129 30°53’S 30°31’E 850 m Etymology The specific name refers to the pitted nature of the integument. Neohyssura skolops sp. nov. Figs 5-6 Description Female Integument thin, only uropods and telson slightly indurate. Body pro- portions: C<1<2=3<4=5=6>7. Dorsolateral grooves present on pereonites but very difficult to see; dorsal pits absent. Anterior five pleonites separate, equal in length and breadth; pleonite 6 fused with telson; latter with broad, rounded base, tapering rapidly into cylindrical spike-like structure tipped with setae; no statocysts visible. Eyes absent. Anterolateral corner of cephalon extending beyond low rostrum. Antennular peduncle 4-segmented, two basal segments broad, fourth segment set obliquely into third, short; flagellum of five articles with few setae and single apical aesthetasc. Antennal peduncle 5-segmented, second and fifth segments subequal, longer than third and fourth segments; second segment grooved to receive antennule; flagellum of seven articles. Mandibular palp 3-segmented, first and third segments subequal, about one-third length of second segment; incisor of three teeth, lacinia plate bearing five blunt teeth; molar process blunt, tapering. 10 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 5. Neohyssura skolops. A. Pereonite 7 and pleon in lateral view. B. Telson and uropods in dorsal view. C. Antenna. D. Antennule. E. Maxilla. F. Maxilliped. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 11 Fig. 6. Neohyssura skolops. A. Pereopod 1. B. Mandible. C. Pleopod 1. D. Lower lip complex. E. Pereopod 7. 12 ANNALS OF THE SOUTH AFRICAN MUSEUM Maxilla with one strong, outer spine and five smaller spines distally. Lobes of lower lip distally bearing two short acute processes. Maxilliped 7-segmented, terminal segment short, bearing four setae; thin-walled endite tapering, with single terminal seta. Pereopods | to 3 similar, subchelate, pereopod 2 more robust than | or 3; unguis one-third length of dactylus, propodus proximally broad, palm convex between three short sensory spines, with few scattered setae on inner face; carpus triangular, produced distally well beyond base of propodus to meet dactylus. Pereopods 4 to 7 slender, unguis one-sixth length of dactylus; propodus with strong posterodistal spine; carpus short, underriding propodus, with strong sensory spine on posterior margin. Uropods somewhat indurate, both rami reaching telsonic apex; exopod situated dorsal to telson, with four strong teeth on medial margin, apically rounded; basis with strong mediodistal spine visible in ventral view. Material Holotype SAM-A15651 2 TL 5,8 mm SM 129 30°53’S 30°31’E 850m Remarks Barnard (1925) noted that ‘Hyssura’ spinicauda Walker, 1901, did not fit the generic diagnosis of Hyssura but he did not decide on the generic status.. Amar (1952), on examining a specimen from Corsica, erected the new genus Neohyssura which he defined as follows: close to Hyssura but the maxillipeds with seven segments, lacking endite, seventh pereopod present, pereopods 4 to 7 with triangular carpus underriding propodus. The present specimen agrees with all except one of these characters. The maxilliped does have a thin-walled endite (easily overlooked), but for the rest is very similar to that of N. spinicauda. The shape of the telson also differs, being narrowly triangular and bearing lateral spines in N. spinicauda but narrowly spike-like in N. skolops. (See Walker 1901, pl. 27, figs 7-11; Monod 1925, pl. 44.) Etymology The specific name ‘skolops’ is taken from the Greek word meaning a sharp object such as spike, and refers to the shape of the telson. Family Paranthuridae Colanthura uncinata sp. nov. Figs 7-8 Description Male Entire integument covered with small imbricate scales. Rostrum extending slightly beyond anterolateral corners of cephalon. Eyes ovate, lateral, consisting of fourteen ocelli. Cephalon two-thirds length of pereonite 1. Pereonites | to 5 THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 13 Ty W/ wf f; Y Tf Fig. 7. Colanthura uncinata. A. Holotype in dorsal view. B. Maxilla. C. Maxilliped. D. Antenna. E. Pereopod 1 with spine further enlarged. F. Antennule 2. G. Antennule 3. 14 ANNALS OF THE SOUTH A8RICAN MUSEUM F Fig. 8. Colanthura uncinata. A. Pleopod 1. B. Pleopod 2 ¢ with apex of stylet further enlarged. C. Pereopod 2. D. Pereopod 6. E. Telson. F. Uropodal exopod. G. Uropodal endopod j and basis. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 15 increasing in width, but subequal in length; posterior margins of pereonites 1 and 2 bilobed. Pereonite 6 two-thirds length of pereonite 5; pereonite 7 very short, subequal to anterior pleonites in length and width, lacking pereopods. Ventral surface of pereonites formed by two longitudinal, low, rounded ridges, conspicuously scaled. Pleon narrower than pereon; pleonites 1 to 5 subequal, pleonite 6 posteriorly bilobed, almost twice length of preceding pleonite. Telson equal to pleonites | to 6 in length, distally evenly convex; dorsally with low, rounded median area widest proximally. Antennular peduncle 4-segmented, basal segment largest; flagellum of six articles, five proximal articles broad, bearing dense aesthetascs, terminal article narrow, bearing setae. Antennal peduncle 5-segmented, second segment largest and broadest; flagellum reduced to setiferous terminal article about one-third width and half length of penultimate segment. Mandible reduced to blunt lobe lacking palp. Maxilla very slender, acute, with about ten faint distal serrations. Maxilliped elongate, consisting of a single segment, distally narrowed, with about seven terminal setae. Pereopod 1 dactylus strongly curved, unguis short; propodus broadly oval, palm gently sinuous, with triangular proximal process and row of fourteen fringed spines set back from the triangular process. Pereopod 2 dactylus gently curved, unguis short; propodus proximally broad, distally narrowed, palm with short spines. Pereopods 3 to 7 similar, unguis of dactylus relatively longer than in pereopods | and 2; propodus with three spines and numerous fine setules on ventral margin, plus two distal fringed spines; carpus not underriding propodus, bearing two ventral spines. Pleopod 1 exopod broad and operculate, with scales on external face; endopod as long as, but one-third width of, exopod. Pleopod 2 exopod and endopod subequal in length and width, each with few distal plumose setae; stylet of endopod extending well beyond apices of rami, distally rounded with strong subapical hook. Uropodal endopod just extending to telsonic apex, oval; exopod narrowly ovate, apically narrowly rounded. Ovigerous female Antennule consisting of five segments, penultimate segment very short. Pereopod | similar to that of male, but with spine row of six (rather than fourteen) spines. Brood pouch formed by four pairs of oostegites, containing four eggs or four larvae. Material Holotype SAM-A15652 3 TL 3,9 mm SM 103 28°31’S 32°34’E 680m Allotype SAM-A15652 ovig. 2 TL 4,5 mm SM 103 16 ANNALS OF THE SOUTH AFRICAN MUSEUM Paratypes USNM 170544 2 3g 1 ovig. 2 SM 103 10 gd 2 ovig. 92 12 immature SM 103 Paratypes SAM-A15653 2 gg 1 ovig. 29 SM 86 27°59’S 32°40’E 550 m 1g lovig.2 9 immature SM 86 Remarks Four species of Colanthura have been described. These are C. tenuis Richardson, 1902, from Bermuda; C. squamosissima Menzies, 1951, from California; C. nigra Nunomura, 1975, from Japan, and C. caeca Mezhov, 1976, from Russia. The present material most closely resembles the Californian species, this similarity being most marked in the character of the integument, both species being almost covered with imbricate scales. Several differences between these two species are apparent. These include pleonites | to 5, which in C. uncinata are subequal, while in C. squamosissima pleonite 5 is four times the length of the preceding segment; the telson, which is more obviously tapered in C. uncinata; and the apex of the stylet of pleopod 2 3, which has a swollen apical lobe in Menzies’s species, but is hooked in C. uncinata. The stylet of pleopod 2 3 in C. caeca is also apically hooked, but possesses three smaller supplementary apical spines (Mezhov 1976, fig. 3). The shape of the uropodal exopod and telson, and the propodal spination of the pereopods of the Russian species differs from C. uncinata. Etymology The specific name is taken from the hooked stylet of pleopod 2 2. Leptanthura minuta sp. nov. Figs 9-10 Description Male Body slender, elongate, not indurate. No dorsal pits or dorsolateral grooves present. Cephalon subequal to pereonite | in length; eyes lacking. Body proportions: C=1<2=3=4<5>6>7. Pleon equal to pereonites 6 and 7 together in length; all pleonites distinct, pleonites | and 5 slightly longer than 2 to 4; pleonite 6 with convex distal margin. Telson elongate-oval in outline, distally evenly rounded, with four median setae; ventrally concave; dorsally with gently raised central area; prominent statocyst present with single medio- dorsal pore. Antennule with 4-segmented peduncle, basal segment largest, fourth segment very narrow; flagellum of eight articles each with dense whorl of aesthetascs. Antenna with 5-segmented peduncle, second segment longest, with disto- dorsal triangular part folding over basal antennular segment; flagellum of four articles. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 17 Mandible narrowly triangular, apically acute; 3-segmented palp with middle segment two and a half times length of basal segment, terminal segment short and curved, with single terminal seta. Maxilla lanceolate, apically acute, with about twelve faint distal serrations. Maxilliped 3-segmented, with few distal setae; second segment six times longer than wide. Pereopod | dactylus fairly stout, unguis one-third length of rest of segment; propodus proximally broad, palm straight with low triangular proximal D Fig. 9. Leptanthura minuta. A. Holotype in dorsal view. B. Mandible. C. Maxilla. D. Maxilliped. E. Antennule ¢. FF. Antennule 2. G. Antenna. H. Pereopod | 9. I. Pereopod 1 ¢. 18 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 10. Leptanthura minuta. A. Pleopod 1. B. Pleopod 2g. C. Pereopod 2. D. Pereopod 7. E. Telson and uropod. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 19 tooth, armed with six short serrate spines and numerous simple setae; carpus triangular, small, with two spines and four setae on ventral margin. Pereopod 2 slightly less robust than pereopod 1, propodus with six spines on palm, lacking dense setae found on pereopod 1; carpus with single ventral spine. Pereopods 4 to 7 similar, slender; propodi with three serrate spines on ventral margin; carpus short, triangular, underriding propodus, with single ventral spine. Pleopod | basis with four retinaculae on median margin, exopod operculate, two and a half times longer than broad with about ten to twelve distal elongate plumose setae; endopod one-third width and two-thirds length of exopod, with eight distal plumose setae. Pleopod 2 with three or four retinaculae on basis; rami subequal in length; exopod with transverse articulation at midlength and about eight distal plumose setae; endopod with four distal plumose setae, stylet on median margin extend- ing well beyond apex of ramus, apically blunt. Uropodal exopod extending to base of endopod, triangular-oval, median margin crenulate and setose; basis with single elongate plumose seta at medio- distal angle; endopod with several distal setae, apically rounded. Female Antennule with 4-segmented peduncle, basal segment longest and broadest, fourth segment very narrow; flagellum reduced, consisting of three articles. Pereopod | very similar to pereopod 1 3, but lacking numerous simple setae on palm. Brood pouch of four pairs of oostegites, anterior pair smallest. Material Holotype SAM-A15654 3 TL 4,6 mm SM 86 27°59’S 32°40’E 550 m Allotype SAM-A15654 ovig. 9 TL 4,5 mm SM 86 Paratypes SAM-A15654 1 gl ovig.2 12 SM 86 Paratypes USNM 170545 1g 1 ovig.2 12 SM 86 SAM-A15655_ 1 ovig. 2 SM 7S. 2731S. 32°S0'E 750 m SAM-A15656 2g3¢ 12 SM 129 30°53’S 30°31’E 850 m Remarks The present small blind species with its characteristic elongate-oval telson and crenulate uropodal exopod bears no close resemblance to any of the approximately eighteen described species of Leptanthura. Certainly it is very different from the described South African species, viz. L. agulhasensis, L. laevigata, and L. urospinosa. The shape of the telson of L. minuta to some extent resembles that of L. antarctica Kussakin, 1967, but the shape of the uropodal endopod and exopod, the antennular peduncle, and the first pereopod easily separate these species. 20 ANNALS OF THE SOUTH AFRICAN MUSEUM Etymology The specific name derives from the small size of the adult animals. Leptanthura natalensis sp. nov. Figs 11-12 Description Male Body very slender and elongate. No dorsal pits. No dorsolateral grooves present, but dorsolateral ridge developed on cephalon and pereonites | and 2. Cephalon lacking eyes. Body proportions: 1<2<3<4>5>6>7. Pereonite 5 in small males (6-7 mm) with two rounded posteroventral lobes; holotype male (20,6 mm) with similar lobes on pereonite 7. Pleonites free. Telson distally broadly rounded, not indurated, dorsally slightly concave, with single, large proximal statocyst. Antennule with 4-segmented peduncle, basal segment largest, fourth seg- ment short; flagellum of thirteen articles. Antenna 8- or 9-segmented, of which three distal segments are probably flagellum; second segment longest. Mandible acutely triangular, palp 3-segmented, basal segment two-thirds length of middle segment, latter with single elongate distal seta, terminal seg- ment short and curved. Maxilla slender, with twelve serrations and three barbs distally. Maxilliped 4-segmented, second segment four and a half times longer than wide, with short distal lobe bearing two setae at mediodistal angle; two distal segments narrowing, bearing few setae. Pereopod | dactylus with very short unguis; palm of propodus almost straight, with low proximal triangular lobe, and row of fourteen short sensory spines, several elongate setae, and dense border of short simple setae. Pereopod 2 propodus narrower than that of pereopod 1, with nine or ten sensory spines on palm, lacking dense border of simple setae; carpus small, triangular, with two spines on ventral margin. Pereopod 7 with short carpus bearing two spines and underriding propodus; latter with three spines on ventral margin. Pleopod | operculate, not indurated; endopod considerably shorter than exopod, both rami bearing distal plumose setae; basis bearing about six retinaculae on medial margin. Pleopod 2 endopod shorter than exopod, with apically blunt stylet on medial margin; exopod with distinct transverse articulation at about midlength, both rami with distal plumose setae; basis armed with five or six retinaculae on medial margin. Uropodal basis triangular in cross-section, longer than endopod; latter narrower than basis, with tuft of apical setae; exopod broadly oval, with few marginal setae, exopods overlapping telson dorsally. ae THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 21 Fig. 11. Leptanthura natalensis. A. Holotype in lateral view. B. Pereonite 5 of 6,8 mm ¢. C. Antennule g. D. Antenna. E. Mandible. F. Maxilla. G. Maxilliped. 22 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 12. Leptanthura natalensis. A. Pleopod 1. B. Pleopod 2 3. C. Pereopod 1 3, with palm enlarged (short, simple setae omitted). D. Pereopod 2. E. Pereopod 7. F. Telson. G. Uropodal exopod. H. Uropodal basis and endopod. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 23 Material Holotype SAM-A15657_ 3 TL 20,6 mm SM 60 27°09’S 32°58’E 800 m Paratypes SAM-A15658 2 gg TL6,8 mm 6,7 mm 1 juv. 4,8 mm SM 53 26°51’S 33°12’E 720m Paratypes SAM-A15659 2 gg TL 6,7 mm 7,5 mm SM 61 27°10’S 32°58’E 820 m Paratypes USNM 170546 1 ¢ TL 8,0mm 1 2 5,7mm SM 117 30°17’S 31°10’E 820 m SAM-A15660 1 ¢ 9,9 mm 1 juv. SM 123 30°33’S 30°48’E 690 m SAM-A15661 7 922 complete but damaged SM 129 30°53’S 30°31’E 850m Remarks The general structure of the pereon and pleon, as well as the mandible, maxilliped, antennae, and pereopods of the present material, agrees with Leptanthura, and quite closely resembles L. lJaevigata and L. glacialis. The broadly rounded telson is similar to that of L. /aevigata (Stimpson) (Barnard 1925), but L. natalensis lacks the characteristic notch in the uropodal exopod, and is a more attenuated species both in body shape and pereopodal propor- tions. The terminal maxillipedal segment is longer than that of L. laevigata. L. glacialis Hodgson (Kussakin 1967) has a uropodal exopod similar to L. natalensis but possesses an apically pointed telson, and shorter and broader antennal segments. L. glacialis appears to have a 3-segmented maxilliped, rather than the 4-segmented structure of L. natalensis. Etymology The specific name is taken from the province, Natal, from whose waters the species was taken. Pseudanthura tenuis Kensley Fig. 13 Pseudanthura tenuis Kensley, 1978: 222, figs 1-2. Description Male Antennular peduncle 4-segmented; segments broad, basal segment equal in length to three distal segments combined; basal flagellar article broad and almost as long as remaining seven flagellar articles together; six distal articles each with pair of aesthetascs. Pleopod 2 stylet of endopod extending well beyond apex of ramus, straight, apically rounded. 24 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 13. Pseudanthura tenuis. A. Pleopod 2 3. B. Antennule ¢. THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 25 Material Le TE9 9mm SM53 26°5S1'S 33°1E 720m 692 TL6,0-25,33mm SM 103 28°31’S 32°34’E 680m be TE 20,7 mm SM 129 30°53’S 30°31’E 850m Remarks P. tenuis Kensley was described from seven females from localities very close to that of the present material. Now that a single male is available, the two main sex-linked characters are described and figured. ACKNOWLEDGEMENTS In addition to the people and institutions thanked in the introductory paper in this series, I should like to thank Mr R. Downes of the South African Museum for sorting the samples of the 1977 cruise. My sincere thanks are also due to Dr T. E. Bowman of the Smithsonian Institution for useful dis- cussions and for critically reading the manuscript. REFERENCES Amar, R. 1952. Isopodes marins du littoral Corse. Bull. Soc. zool. Fr. 77: 349-355. BARNARD, K. H. 1925. A revision of the family Anthuridae (Crustacea Isopoda) with remarks on certain morphological peculiarities. J. Linn. Soc. 36: 109-160. KENSLEY, B. 1977. The South African Museum’s Meiring Naude cruises. Part 7. Marine Isopoda. Ann. S. Afr. Mus. 74: 125-157. KENSLEY, B. F. 1978. Two new species of the genus Pseudanthura Richardson (Crustacea, Isopoda, Anthuridea). Proc. biol. Soc. Wash. 91: 222-233. KussAKIN, O. G. 1967. Fauna of Isopoda and Tanaidacea in the coast zones of the Antarctic and Subantarctic waters. Biol. Rep. Sov. Antarct. Exped. (1955-58) 3: 220-380. Louw, E. 1977. The South African Museum’s Meiring Naude cruises. Part 1. Station data 1975, 1976. Ann. S. Afr. Mus. 72: 147-159. Menzies, R. J. 1951. New marine isopods, chiefly from northern California, with notes on related forms. Proc. U.S. natn. Mus. 101: 105-156. MEzHov, B. V. 1976. New species of Anthuroidea from the upper part of the sublittoral zone of the Middle Kurile Islands. Biol. Sea 5: 19-27. Monon, T. 1925. Tanaidacés et Isopodes aquatiques de l’Afrique Occidentale et Septentrionale. Bull. Soc. Sci. nat. Maroc 5: 233-247. NunomurA, N. 1975. Marine Isopoda from the rocky shore of Osaka Bay, Middle Japan. Bull Osaka Mus. nat. Hist. 29: 15-35. RICHARDSON, H. 1902. The marine and terrestrial isopods of the Bermudas, with descriptions of new genera and species. Trans. Conn. Acad. Arts Sci. 11: 277-310. WALKER, A. O. 1901. Contributions to the Malacostracan fauna of the Mediterranean. J. Linn. Soc. 28: 290-307. 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: Nickles, 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 shou!d 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 8 ISOPODA ANTHURIDEA a AL. Caplou— OLUME 77 PART 2 DECEMBER 1978 MUS. COMP. ZOOS LIBRARY SN 0303-2515 FEB 15 07) HARV far UNIVERSITET? ANNALS OF THE SOUTH AFRICAN MUSEUM CAPE TOWN 1. MATERIAL should be original and not published elsewhere, in whole or in part. 2. LAYOUT should be as follows: (a) INSTRUCTIONS TO AUTHORS 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) Abstract of not more than 200 words, intelligible to the reader without reference to the text Table of contents giving hierarchy of headings and subheadings Introduction Subject-matter of the paper, divided into sections to correspond with those given in table of contents Summary, if paper is lengthy Acknowledgements References 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.: (b) ‘Smith (1969) describes...’ ‘Smith (1969: 36, fig. 16) describes .. .” “As described (Smith 1969a, 19696; Jones 1971)’ ‘As described (Haughton & Broom 1927)...’ “As described (Haughton ef 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. Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, 5, 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. FiscHErR, 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. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51. TurELe, 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 77 Band December 1978 Desember Part 2 Deel BODITIONS TO THE REVISED LIST OF PRESERVED maALERIAL OF THE EXTINCT CAPE COLONY QUAGGA AND NOTES ON THE RELATIONSHIP AND DISTRIBUTION OF SOUTHERN PLAINS ZEBRAS By cE; EY RAY 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, 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, 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 Copyright enquiries to the South African Museum Kopieregnavrae na die Suid-Afrikaanse Museum ISBN 0 908407 57 2 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 ADDITIONS TO THE REVISED LIST OF PRESERVED MATERIAL OF THE EXTINCT CAPE COLONY QUAGGA AND NOTES ON THE RELATIONSHIP AND DISTRIBUTION OF SOUTHERN PLAINS ZEBRAS By R. E. RAu South African Museum, Cape Town (With 9 figures) LMS. accepted 7 September 1978] ABSTRACT Previously unpublished photographs and data of two mounted skins of the extinct quagga (Equus quagga quagga) and a further relic are added to the list of preserved material. Photographs of the destroyed KGnigsberg quagga are reproduced. Examination of plains zebra (Equus ‘burchelli’) skins from southern Africa and comparison with the true Burchell’s zebra (Equus ‘burchelli burchelli’) and the quagga (Equus quagga quagga) suggest that all belong to one species (Equus quagga) and are forms which are dominant in certain areas within the continuous distribution, rather than subspecies. In addition, the true Burchell’s zebra is considered to be still extant in Zululand and possibly Swaziland, if the generally accepted but vague range of that ‘subspecies’ is ignored. CONTENTS PAGE Introduction . ; : : : 2 : eT Additional material : ; : : : eaves Notes on Burchell’s zebra and quagga : Ay si Reintroductions of zebras. . : : Ae: | Acknowledgements . : d ‘ : : . 44 References . : : , : : : = . 44 INTRODUCTION Since the publication of the revised list of preserved material of the quagga (Equus quagga quagga) a photograph of and additional data on the quagga at the Zoological Museum, State University at Kazan, Soviet Union, have now been obtained. In addition, a mounted quagga has been discovered at the Muséum d’Histoire Naturelle Lyon, France, and a mounted foot at the Royal Albert Memorial Museum at Exeter, England. These new data are presented in accordance with the method followed by Rau (1974). The photographs included in the description of the Kénigsberg quagga (Hilzheimer 1930), which was destroyed at the end of the Second World War, are reproduced to complete the illustrations of mounted skins which existed into this century. 27 Ann. S. Afr. Mus. 77 (2), 1978: 27-45, 9 figs. 28 ANNALS OF THE SOUTH AFRICAN MUSEUM In a monograph Antonius (1951), a world authority on zebras, included the quagga and the Burchell’s zebras in the single species Equus quagga, as had been done earlier (Pocock 1904; St Leger 1932). This view is supported by Rzasnicki (1951), Mohr (1964) and Klingel (1969, 1972) amongst others. There is, however, a second school of thought which divides them into two species, Equus quagga and Equus burchelli (Ansell 1971). As the present study has revealed that there is no definite division between the quagga and Burchell’s zebras, the former view is supported. ADDITIONAL MATERIAL KAZAN— MOUNTED SKIN Fig. 1 Catalogue number: 109 Sex: female Locality: unknown Date of acquisition: 1843 Remarks on acquisition: bought in Hamburg from a Mr Brandt by Professor Eversman for the Zoological Museum of Kazan University Fig. 1. Kazan quagga. ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 29 History of mount: bought as mounted specimen in bad condition and remounted ; a second remounting (‘restoring’) was performed in 1969 by M. A. Zaslavski in Leningrad, U.S.S.R. Description of striking features: face clearly striped; shadow-stripes on neck; posterior part of feet (fetlock-hoof) dark; the photograph seems to show faint striping on buttock Measurements: head-body 2,36 m* (2,42 m)* tail 0,54m (0,53 m) ear 0,14m (0,17 m) hind foot 0,56 m (0,54 m) shoulder height 1,29 m_ (1,28 m) State of preservation: appears to be good Further material of same individual: none Remarks: exhibited open in gallery References: none known LYON— MOUNTED SKIN Fig. 2 Catalogue number: 6102 Sex: female Locality: Kaffraria, South Africa Date of acquisition: unknown Remarks on acquisition: none History of mount: appears to be the original mount Description of striking features: face not clearly striped Measurements: head-body 2,10 m tail 0,40 m ear 0,17 m hind foot 0,50 m shoulder height 1,15 m State of preservation: fair Further material of same individual: none Remarks: exhibited; specimen not mentioned in previous literature; V. Eisen- mann of the Muséum d’Histoire Naturelle, Paris, discovered this specimen early in 1975 and kindly informed the author References: M. Philippe, ‘La zoologie au Muséum d'Histoire Naturelle de Lyon’ (in preparation) * Measurements received through two intermediaries are not identical. 30 ANNALS OF THE SOUTH AFRICAN MUSEUM —_ Fig. 2. Lyon quagga. EXETER— FOOT TROPHY Fig. 3 Catalogue number: A758 Locality: Bontebok Flats, British Kaffraria, South Africa (between present Cathcart and Queenstown) Date of acquisition: after 1869 ) Remarks on acquisition: from the collection of W. S. M. D’Urban, first curator of Exeter Museum Measurements (hoof only): anterior height 6,0 cm width 6,7 cm \ length 8,4 cm . References: Howes & Bamber (1970) ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 3] Fig. 3. Exeter quagga foot; anterior and lateral views. KONIGSBERG QUAGGA Fig. 4 The K6nigsberg quagga was destroyed at the end of the Second World War when Schloss Waldhof, about 20 km from KGnigsberg, where museum specimens had been taken for safe-keeping, was shelled and completely destroyed by fire (pers. comm. Prof. Dr O. Koehler (1972), director of the K6nigsberg Museum until 1945). The preserved material of Equus quagga quagga, including the above specimens, now consists of: skins—23, plus 1 mounted head and neck complete skeletons—7 skulls—20, including 7 from complete skeletons; at least 3 of the mounted skins appear to contain skulls loose skeletal parts—2 sets of foot bones; upper and lower incisivae in portion of jaws other—dried connective tissue, flesh, etc., removed from skin; nasal cartilages and palate removed from skull (in alcohol) (Rau 1974: 58) NOTES ON BURCHELL’S ZEBRA AND QUAGGA During February to October 1977 approximately 400 privately owned skins of plains zebra Equus quagga (burchelli) and mountain zebra Equus zebra hartmannae were processed at a Cape Town tannery and examined by the author. These originated from many different localities in southern Africa. Skins of 113 plains zebras were photographed, the initial 88 non-selectively and 32 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 4. Konigsberg quagga (destroyed), adapted from Hilzheimer (1930). ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 35 the remaining 25 selectively. The proportion of the two species processed during a two-month period (February and March) was 88 plains zebra: 62 mountain zebra. A great number of ‘subspecies’ and even ‘species’ of plains zebra have been described. In bestowing names to the many variations, the stripe pattern has been the principal, and often the only, character considered (Ansell 1971). This is certainly true in the case of the ‘subspecies’ burchelli. The diagnostic characteristics of the true Burchell’s zebra Equus quagga burchelli are usually given as follows: whitish, unstriped legs except occasionally some traces of stripes on the hocks (Pocock (1897: 52) accepts some stripes on the ‘knees’ as well); basic colour of dorsal parts white to ochre and brownish; interspaces with shadow-stripes, which may be broad and intense, or less conspicuous; body-stripes not meeting the ventral midline. Equus quagga antiquorum is defined by the striping extending on to the legs to at least the ‘knees’ and hocks, but sometimes to the pasterns. Equus quagga burchelli (E. burchelli burchelli) is one of the few still recog- nized as a valid subspecies. The accepted distributional range of this subspecies included the Orange Free State, the north-eastern Cape Province, southern Botswana and the south-western Transvaal. As zebras no longer occur in these regions, other than reintroduced specimens, the true Burchell’s zebra is usually considered to have been extinct since about 1910. The locality of the type specimen on which Gray (1825) based the descrip- tion of the subspecies is between Campbell and the Vaal River. The specimen figured as Equus quagga burchelli by Antonius (1951, fig. 22) originated from British Bechuanaland (Botswana) and lived at Hagenbeck’s Tierpark, Germany, during 1907-9. However, most preserved specimens of this subspecies lack locality data, as do those individuals that lived at menageries or zoos and are known from illustrations and/or descriptions. It appears, therefore, that the accepted range is based on old reports rather than collected material, and that the true range is not known. Indeed, different opinions as to the range of Equus quagga burchelli have been expressed. Matschie (1894) has included Zululand while Shortridge (1934) and Eloff (1967) extend the range to Great Namaqualand (South West Africa). The concept of the range of a subspecies within the continuous distribution of a species is open to question, especially in the case of fast-moving and migrating animals such as the plains zebra. With regard to zebra distribution, Tegetmeier & Sutherland (1895) wrote: ‘Disputing, therefore, as to whether two closely allied animals are specifically or sub-specifically distinct is almost a waste of words. We know that a species spread over a wide area will change according to the conditions of life, until at last the two extremes are so diverse as to be regarded as distinct species, but no one can say where one species ends and the other begins, for they merge gradually into each other.’ The uncertainty about the former range of the subspecies Equus quagga burchelli has been aggravated by the indiscriminate use of the names ‘quagga’ 34 ANNALS OF THE SOUTH AFRICAN MUSEUM and ‘Burchell’s zebra’, and by the difficulty in distinguishing between those two forms as well as between the true Burchell’s zebra and its northern relatives. Accurate assessment of old documents is, therefore, practically impossible. Pocock (1897: 42) wrote 13 years before the alleged extinction of Equus quagga burchelli: *. .. and on account of the general application of the term Burchelli to any or all of its subspecies, we are in ignorance of the exact area that it now occupies in S. Africa.’ If all the mounted zebra specimens preserved in museums had precise locality data, there would be in all probability antiquorum-types from the burchelli ‘range’ and vice versa. Since most older mounted zebras do not, however, have conclusive locality data, it is accepted that apparent burchelli specimens originate from the accepted burchelli range while antiquorum patterned specimens are thought to be from further north. Since the plains zebra has a continuous distribution, recent interferences by man excepted, within which certain forms or variations predominate in certain areas, it follows that, in intergradational areas between two forms, the one form will gradually decrease in percentage of total numbers as the area of the next form is approached. Despite the fact that for most of the preserved or known plains zebra specimens, which conform basically to the description of Equus quagga burchelli, the place of origin is not known, they are accepted as belonging to that sub- species, as is shown in the figures by Cabrera (1936) (Fig. 5) and Antonius (1951). The subspecies Equus quagga antiquorum is presently considered to include all the extant southern forms of the plains zebra, ranging from southern Angola and (historical) Damaraland through Botswana and Transvaal to Zululand. Consequently, individual variation within antiquorum is considerable (Fig. 6). Eloff (1967) stated that the three subspecies antiquorum, chapmani and wahlbergi occur together in Zululand. Antonius (1951) had observed that in Damara zebra and especially in Wahlberg zebra burchelli-like individuals do occur. However, such individuals originate from outside the presumed range of the true Burchell’s zebra and are consequently not accepted as such. Neverthe- less, some authors are cautious. Cabrera (1936: 94), while saying that the subspecies ‘appears to be extinct’, also states, ‘in fact, I regard as typical burchelli all members of the species having all four legs white from the elbow and the stifle joint, except for the occasional presence of a few short markings across the hocks, and with the thighs free of complete and well-defined dark stripes behind the stifle stripe’. Dorst & Dandelot (1970) list the subspecies burchelli as ‘probably extinct’. Amongst the extremely variable skins (many with numerous small, light dots in stripes and shadow-stripes) examined at the tannery, were several specimens from a Zululand game farm with a surprising combination of light basic colour and few and/or weak shadow-stripes and considerable stripe reduction (Fig. 7A). It is generally believed that the change in coloration of plains zebras to the south of their distributional range is achieved by stripe- ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 35 Fig. 5. Examples of true Burchell’s zebra, Equus quagga burchelli, shown in standardized outline. A. Type of ‘Equus burchelli paucistriatus’ Hilzheimer, 1912, at Mainz Museum. B. Animal that lived at Jardin des Plantes, Paris, 1826. C. Type of subspecies from Little Klibbolikhonnifontein, British Museum (Natural History) (destroyed). D-E. Animals that lived at Knowsly Park, about 1845. F. Animal that lived at Zoological Garden, Dresden. (Adapted from Cabrera (1936).) 36 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 6. Examples of Equus quagga antiquorum, shown in standardized outline. A. From Rietfontein West, Munich Museum. B. Type of subspecies from Angola (Smith 1841). C. From eastern Transvaal. D. From west of Lake Ngami. E. From Zululand. F. From eastern Transvaal. (Adapted from Cabrera (1936).) ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 37 reduction and the darkening of basic colour and the occurrence of shadow- stripes. However, the above specimens indicate that these processes do not necessarily occur together. Indeed, there are specimens from other areas which, in spite of more extensive striping, display dark basic colour (Fig. 8A), some with numerous strong shadow-stripes (sometimes three in one interspace). From the Zululand skins (Fig. 7) and additional photographs it appears that stripe- reduction is the dominant feature in this eastern population, while to the west on the interior plateau the darkening in both basic colour and through shadow- stripes is more pronounced. In terms of stripe reduction these Zululand skins certainly do not fit into the concept of Equus quagga antiquorum (Fig. 6), but they do agree with the description of Equus quagga burchelli. Indeed, a few are less striped than some of the burchelli specimens figured by Cabrera (1936) (Fig. 5) and others. It appears, therefore, that Equus quagga burchelli is, in fact, not extinct but still survives in Zululand (and possibly Swaziland), and that its range does extend eastward into Zululand, as Matschie (1894) stated. (The ‘myriads of Quaggas’ which Chapman (1868, cited in Antonius 1951: 96) encountered in the extreme north-eastern Orange Free State could thus have been true burchelli, and not as Antonius (1951: 96) considered, Wahlberg zebra.) Otherwise the subspecies burchelli would have to be considered as part of the antiquorum complex, a thought similarly expressed by Rzasnicki (1951) and Roberts (1951), cited in Ansell (1971). The only other alternative would be not to recognize such individuals (Figs 7B—D, 9B, D) as members of the subspecies burchelli. However, had they been old mounted specimens, they would be regarded as true Bur- chell’s zebras. It must be noted that the farm where these specimens had been shot is a hunting farm which was originally stocked with 300 zebras from Hluhluwe Game Reserve, and that hunters select the most striped specimens (C. Tinley, pers. comm.). Selective breeding is thus exercised unintentionally and this might have ‘re-created’ burchelli, although similar specimens do occur in Hluhluwe and Umfolozi Game Reserves, the only area in Zululand where the original zebra population has not been interfered with through reintroductions (P. M. Brooks, pers. comm.). Zululand undoubtedly is an inter-gradational area between the two forms burchelli and antiquorum. Observations made during a recent visit to that area and Swaziland suggest that the percentage of burchelli types may be as high as 15 in Zululand, although P. M. Brooks, biologist at Hluhluwe, estimates it to be 5-10 per cent (pers. comm.). The north-eastern and eastern parts of Swaziland also retain an endemic population of plains zebra (T. E. Reilly, pers. comm.). There is no geographic barrier between Zululand and the Swaziland lowveld, and it is not surprising that some of the zebras observed there fit or approach the description of Equus quagga burchelli, though the percentage of such specimens must be expected to be lower than further south. 38 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 7. Skins of plains zebras from Zululand, 1977. A. Note stripe-reduction, few shadow- stripes. B-D. Note shadow-stripes on neck and stripe-reduction. (B now at South African Museum, ZM38819.) ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 39 ys _ es Petey. Td | - | ww 7g _- * “ mm << Fig. 8. Skins of plains zebras from southern Africa. A. Precise locality unknown; note dark basic colour of upper parts compared to lower leg. B. Precise locality unknown; note numerous shadow-stripes and extensive striping. C. From game farm in Zululand; note dark basic colour of upper parts. D. From southern Kaokoveld, South West Africa; note little contact between body-stripes and dorsal midline. 40 ANNALS OF THE SOUTH AFRICAN MUSEUM Despite the uncertainty whether the ‘quaggas’ of old reports were, indeed, the true quagga, Equus quagga quagga, the northern boundary of its range is usually considered to have been formed roughly by the Orange and Vaal rivers. The eastern limit might have been about the boundary between the Orange Free State and Natal (Grout, cited in Shortridge 1934: 402). It is also generally accepted that the ranges of quagga and true Burchell’s zebra overlapped in the Orange Free State (Antonius 1951; Liebenberg 1964; Ansell 1971). Antonius (1951) was of the opinion that the quagga, being essentially a form adapted to the Karoo and possibly the most distinct of all the plains zebra subspecies, expanded its range secondarily in a northerly direction and met and overlapped along a broad front with the southward expanding Burchell’s zebra. He obviously did not believe in reports about the social and reproductive separation of the two forms, since he considered one of the preserved quagga specimens (the more striped specimen of ‘Equus burchelli paucistriatus’ at Mainz (Fig. 9A and Rau 1974, fig. 13) as a possible cross between the two forms (Antonius 1951: 26). It is difficult to accept that separate herds of the two forms existed in the same locality. Even if they did represent separate species, as some are inclined to believe, herds formed by members of both must be expected, since this is known to occur in extant zebra species where their ranges overlap. For example, Equus zebra hartmannae overlaps with Equus quagga antiquorum in South West Africa (Antonius 1951: 82) and Equus grevyi forms mixed herds with Equus quagga boehmi in Kenya (Antonius 1951: 82; Bartlett 1963). Even true quaggas were seen together with the Cape mountain zebra at Geelbeckfontein, at the southern limit of the Karoo, by Barrow (1801, cited in Antonius 1951: 82). As shown previously (Rau 1974) the quagga and the true Burchell’s zebra approach one another in basic colour and in stripe reduction, so much so that some of the preserved specimens (e.g. Rau 1974, fig. 12 right) have variously been regarded as one or other of the two. Furthermore, some of the charac- teristics previously considered to be typical of the quagga (Rau 1974) have been observed in the skins at the Cape Town tannery and in a specimen from Kruger National Park. These characteristics include narrow interspaces (Smuts 1974, fig. 1.4.5), dorsal midline being flanked by a light band on either side, and body- Stripes not in contact with dorsal midline except sometimes via narrow bridges (Fig. 8D). Antonius (1951: 29) has already pointed out that the quagga had reddish dark-brown stripes which, especially on the body, merge into the reddish basic colour. It has previously been explained why the coloration of the quagga has been incorrectly interpreted as light stripes on a dark background (Rau 1974: 43). Careful examination reveals beyond all doubt that the quagga, like all the other plains zebras, is dark-striped and that the interspaces darken progressively from the shoulder in a posterior direction until they become equal in tone with stripes and shadow-stripes to produce the ‘uniform’ posterior portion. In other southern plains zebras the interspaces are also lightest on head and neck. Ansell (1971: 5) ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 4] and others are wrong in defining the striping in the quagga as ‘whitish on dark background’. Since identification is therefore complex, and since it is impossible to observe the stripe pattern of each individual when looking at a herd, the alleged recognition of separate herds of quaggas and Burchell’s zebras in the same locality must be treated with suspicion. The grouping of plains zebras into subspecies is thus somewhat vague, or, as it is expressed by Ansell (1971) ‘has never been satisfactory’. In an attempt to sort out this confusion, Pocock (1897) pointed out that throughout the distri- bution area of the plains zebra ‘intermediate types’ do exist between particular forms which prevail in particular localities. In addition, as mentioned earlier, typical representatives of the two forms occur in intergradational areas, the closer to the area of the one form the lower the percentage of the other form. The high degree of variability, both between geographically distant populations and within one herd, has been attributed by Antonius (1951) to the process of eliminating the contrasting stripe-pattern. It should be noted that Antonius’s ‘Riickbildung der Bein- und Rumpfstreifung’ has been misinterpreted by Eloff (1966, 1967) as being a sign of genetic pauperization. As shown above, the gradual change from one form to another in plains zebras appears to include, at least in exterior characteristics, the ‘separation’ into Equus quagga and Equus burchelli. Relatively few skulls of the true quagga are available and some of these lack sufficient data to establish their authenticity. The fact that some skulls may have been obtained by breaking up an old stuffed -quagga skin (as has happened at Berlin, where the skull was later identified as that of Equus caballus (Antonius 1951: 35)) is no guarantee of their identity as Equus quagga quagga. However, some zoologists believe that there are specific morphological differences between skulls of the quagga and the plains (Burchell’s) zebra (e.g. Cooke 1950; Lundholm 1951; Gentry 1975; V. Eisenmann, pers. comm.). The recent study by V. Eisenmann (pers. comm.) has shown that in some skull characters there is a cline from northern to southern plains zebras. In other words, in some respects at least skulls of southern plains zebras approach those of the quagga more closely than do northern forms. This conforms to the well-documented cline in colour and stripe-pattern. To conclude, it would appear that the Burchell’s zebras and the quagga should be regarded as one species, Equus quagga. Furthermore, since the grouping into subspecies, either by appearance or by distribution, is not clear-cut in the case of the plains zebras, it might be preferable to refer them to forms, types or variations prevailing in certain areas, rather than to subspecies. REINTRODUCTIONS OF ZEBRAS The reintroduction of animal species into areas where natural populations have been exterminated is increasingly common. There are, however, problems with this practice, some of which have been discussed by Greig (1977). 42 ANNALS OF THE SOUTH AFRICAN MUSEUM V BRANCO Dis A h - ( V BRANCO Fig. 9. Stripe patterns and tone of basic colours of various plains zebras, shown in standardized outline. A. Female quagga at Mainz Museum. B. Burchell’s zebra from Zululand (same as Fig. 7B). C. Male Burchell’s zebra at Leiden Museum. D. Burchell’s zebra from Zululand (same as Fig. 7D). E. Quagga at Tring Museum (near London). F. Type of true Burchell’s zebra, British Museum (Natural History) (destroyed). G. Quagga at Berlin Museum. H. Male quagga at Mainz Museum. I. Burchell’s zebra at Mainz Museum (type of ‘Equus burchelli paucistriatus’ Hilzheimer, 1912). = —--—- ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 43 44 ANNALS OF THE SOUTH AFRICAN MUSEUM Both plains and mountain zebras are amongst those South African mammals whose ranges have been much restricted during the past three centuries. Now, and in the recent past, zebras are being translocated from areas where populations survive to areas which they formerly inhabited. In the case of the plains zebras such reiniroductions have involved the movement into certain areas of forms dissimilar to those which previously occurred there, even though the original form, or one close to it, still exists. As indicated above, the ‘subspecies’, Equus quagga burchelli, apparently survives in Zululand and thus, when reintroductions of zebras into the former range of the true Burchell’s zebra are contemplated, it would be desirable that selected specimens from Zululand be used, rather than animals of the antiquorum type. It would even be desirable to eliminate former indiscriminate reintro- ductions and to replace them with individuals which are closer to the original population. With a minimum of initial selective breeding in order to eradicate recessive antiquorum characteristics, which are present also in burchelli-like individuals from Zululand (an intergradational area between the two forms), a ‘pure’ population of plains zebras resembling the former endemics could be achieved within a relatively short period. ACKNOWLEDGEMENTS My sincere thanks are due to Dr Vera Eisenmann of the Muséum National d’Histoire Naturelle, Paris, for kindly informing me of the mounted quagga which she discovered at the museum at Lyon, France. I also thank Dr C. A. Howes for drawing my attention to the quagga foot which is housed at the Royal Albert Memorial Museum, Exeter, England, Dr I. Sokolov of the Academy of Sciences, Leningrad, U.S.S.R., and Mr E. Grangqvist of the Zoological Museum, Helsinki, Finland, for being instrumental in obtaining photographs and data of the mounted quagga in the Museum at Kazan, U.S.S.R. I am indebted to Mr V. Branco for producing the drawings, Mr A. Byron for some of the photographic reproductions and to Prof. G. H. Findlay, and Drs Q. B. Hendey and P. A. Hulley for their co-operation. REFERENCES ANSELL, W. F. H. 1971. Perissodactyla. Jn: MEESTER, J. & SETZER, H. W. eds. The mammals of Africa. An identification manual 14: 1-14. Washington, D.C.: Smithsonian Institution Press. ANTONIUS, O. 1951. Die Tigerpferde: die Zebras. Monogrn Wildsdugetiere 11: 1-148. BARTLETT, D. 1963. Zeehorses and Zebronkeys. Animals 2: 394-396, CABRERA, A. 1936. Subspecific and individual variation in the Burchell zebras. J. Mammal. 17: 89-112. Cooke, H. B. S. 1950. A critical revision of the quaternary Perissodactyla of Southern Africa. Ann. S. Afr. Mus. 31: 393-479. Dorst, J. & DANDELOT, P. 1970. A field guide to the larger mammals of Africa. London: Collins. ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 45 Etorr, G. 1966. The passing of the true quagga and the little Klibbolikhonnifontein Burchell’s zebra: which is to be next the Cape mountain zebra or Wahlberg’s zebra of Zululand? Tydskr. Natuurwet. 6: 193-207. Exorr, G. 1967. The phenomenon of the obliteration of stripes of the Burchell zebra with a southerly migration. Zodn. Pretoria 8: 10-13. Gentry, A. W. 1975. A quagga, Equus quagga (Mammalia, Equidae), at University College, London, and a note on a supposed quagga in the city museum, Bristol. Bull. Br. Mus. nat. Hist. (Zool.) 28: 217-226. Gray, J. E. 1825. A revision of the family Equidae. Zool. J. Lond. 1: 241-248. GreiG, J. C. 1977. The principles and problems of genetic conservation in relation to wildlife in South Africa. Cape Dept. of Nature & Environmental Conservation, Research report, September 1977: 1-55. (Mimeographed.) HivzHemer, M. 1930. Das K6nigsberger Quagga. Z. Sdugetierk. 5: 86-95. Howes, C. A. & BAMBER, M. 1970. Rarities in a museum. Oryx 10: 326-328. KLINGEL, H. 1969. The social organisation and population ecology of the plains zebra, (Equus quagga). Zoologica afr. 4: 249-263. KuINGEL, H. 1972. Das Verhalten der Pferde. Handb. Zool., Berl. 8 (49): 1-68. LIEBENBERG, L. C. C. 1964. Die grotere soogdiere wat vroeér dae voorgekom het in die omgewing van die Golden Gate-Hooglandpark. Koedoe 7: 99-104. LUNDHOLM, B. 1951.-A skull of the true quagga (Equus quagga) in the collection of the Transvaal Museum. S. Afr. J. Sci. 47: 307-312. Martscule, P. 1894. Die afrikanischen Wildpferde als Vertreter zoogeographischer Subregionen. Zool. Gart. Lpz. 35: 33-39; 65-74. Monr, E. 1964. Von Rosshauten und Zebrafellen. Das Pelzgewerbe 15: 161-168. Pocock, R. I. 1897. The species and subspecies of zebras. Ann. Mag. nat. Hist. (6) 20: 33-52. Pocock, R. I. 1904. The Cape Colony quaggas. Ann. Mag. nat. Hist. (7) 14: 313-338. Rau, R. E. 1974. Revised list of the preserved material of the extinct Cape Colony quagga, Equus quagga quagga (Gmelin). Ann. S. Afr. Mus. 65: 41-87. Roserts, A. 1951. The mammals of South Africa. Johannesburg: Trustees of The mammals of South Africa Book Fund. RZASNICKI, A. 1951. Zebras and quaggas. Ann/s Mus. zool. pol. 14: 203-251. SHORTRIDGE, G. C. 1934. The mammals of South West Africa 1: 397-411. London: Heinemann. SmitH, C. H. 1841. The natural history of horses. Edinburgh: Lizars; London: Highley; Dublin: Curry. (The naturalist’s library [Vol. 31]. Mammalia 12.) Smuts, G. L. 1974. Growth, reproduction and population characteristics of Burchell’s zebra in the Kruger National Park. Unpublished D.Sc. thesis, University of Stellenbosch. St Lecer, J. 1932. On Equus quagga of south-western and eastern Africa. Ann. Mag. nat. Hist. (10) 10: 587-593. TEGETMEIER, W. B. & SUTHERLAND, C. L. 1895. Horses, asses, zebras, mules and mule breeding. London: Horace Cox. ~— , ae) erties a oe Ye baal £ hell 2» an re 7 > , Sf 7 1 : >» — “a 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 14-15SA 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 shou!d 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. 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. R=. RAV ADDITIONS TO THE REVISED LIST OF PRESERVED MATERIAL OF THE EXTINCT CAPE COLONY QUAGGA AND NOTES ON THE RELATIONSHIP AND DISTRIBUTION OF SOUTHERN PLAINS ZEBRAS PART 3 DECEMBER 1978 OLUME 77 id ee tea: ~ so. !SSN 0303-2515 LIBRARY FEB 15 1979 HARVARD 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. 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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; Jones 1971)’ ‘As described (Haughton & Broom 1927)...’ ‘As described (Haughton ef 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) BuLLouGu, 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. Konun, 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. TutELe, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: 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 77 Band December 1978 Desember Part 3 Deel REDESCRIPTION OF PLIOPLATEIA K. H. BARNARD, A GENUS OF AMPHIPOD (CRUSTACEA) FROM SOUTH AFRICA By J. LAURENS BARNARD 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, 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, 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 Copyright enquiries to the South African Museum Kopieregnavrae na die Suid-Afrikaanse Museum ISBN 0 908407 60 2 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 REDESCRIPTION OF PLIOPLATEIA K. H. BARNARD, A GENUS OF AMPHIPOD (CRUSTACEA) FROM SOUTH AFRICA By J. LAURENS BARNARD Department of Invertebrate Zoology, Smithsonian Institution, Washington (With 4 figures) [MS. accepted 12 September 1978] ABSTRACT Plioplateia K. H. Barnard (1916) is removed from Phliantidae to form the type genus of a new family demonstrating evolutionary outflow from the southern Pacific family Ceinidae towards the circumtropical family Phliantidae. At least nine major characters constrain Plioplateia from assignment to Phliantidae. Plioplateia appears to be the last living relict of what may have been a diverse group of taxa standing between ceinids and phliantids. It joins many other South African amphipods now considered to be relicts. CONTENTS PAGE Introduction . . ee ce ed ee A, Plioplateidae, fam. nov. . . . . . 48 Plioplateia K. H. Barnard . 49) Plioplateia triquetra K. H. Barnard . : OO Acknowledgements . .. . . . S55 Retercncest Paie® Fhe eres, Legett 55 INTRODUCTION Plioplateia, a gammaridean amphipod, was described by K. H. Barnard (1916) and placed in the family Phliantidae. Since that time many genera in this evolutionary vicinity have been described and then later realigned, mainly by J. L. Barnard (1972a, 19726) and Griffiths (1975), into families Phliantidae, Temnophliidae, Kuriidae and Ceinidae, with subfamily Chiltoniinae. Plioplateia appears to stand between Ceinidae and Phliantidae but differs in so many ways from either family group that a new family is established for Plioplateia. All of these families are included in the superfamily Talitroidea. The Plioplateidae join the Temnophliidae of South Africa as the narrowest relict families known in the Gammaridea except for the Kuriidae from Abd-el-Kuri near the Gulf of Aden. South Africa is noted for other isolated or relict amphipods, mainly the genera Calliopiella Schellenberg, Cypsiphimedia K. H. Barnard, Dikwa Griffiths, Triodos K. H. Barnard, Hoplopleon K. H. Barnard, Exampithoe K. H. Barnard, Macropisthopus K. H. Barnard, Unciolella Chevreux, Elasmopoides Stebbing, Cunicus Griffiths and Phoxostoma K. H. Barnard. 47 Ann. S. Afr. Mus. T7 (3), 1978: 47-55, 4 figs. 48 ANNALS OF THE SOUTH AFRICAN MUSEUM Plioplateidae fam. noy. Type genus Plioplateia K. H. Barnard, 1916. Diagnosis Talitroidea with laterally compressed tall, cuspidate bodies but coxae poorly splayed and excavate or cuspidate; coxa 4 small. Cuticle humped and rough. Head dorsoventrally compressed, complexly cuspidate. Antennae disparate, cuspidate, flagella elongate. Anterior part of body dorsally cuspidate. Mandibular molar huge, granular, not distinctly triturative. Inner lobes of lower lip well developed. Outer plate of maxilla | with eight spines. Inner plate of maxilla 2 much narrower and shorter than outer. Outer plate of maxilliped not larger than inner plate, palp articles | and 4 elongate, palp thin. Gnathopods subchelate, hands expanded, lacking giant facial setae. Article 2 of pereopods 5-7 narrow, small. Pleon small, deeply flexed below thorax, urosomite 3 distinct but vestigial. Pleopods elongate, peduncles thin, rami elongate, thin, uni-articulate. Uropod 3 formed of leaf-like peduncle lacking ramus. Telson forming erect cowl, deeply cleft. Relationship On first sight Plioplateia appears to belong to the Phliantidae as originally placed by K. H. Barnard, but Plioplateia differs from phliantids in the elongate flagella of the antennae, the elongate, thin pleopods with equal and uniarticulate rami, the giant mandibular molar, the presence of well-developed inner lobes on the lower lip, the small inner plate of maxilla 2, the small outer plate of the maxilliped, the thin maxillipedal palp with elongate article 1, the subchelate gnathopods lacking giant setae, and the cowl-like cleft telson. Plioplateia bears the bidentate pereonite | similar to that of many phliantids but would appear to be much more primitive and to share many features of the plesiomorphic Ceinidae (see J. L. Barnard, 1972a, 1972b). To a great extent Plioplateia fits many aspects of a model ancestor to phliantids with evolutionary outflow from ceinids, and, in addition, its isolation in South Africa fits the role of a relict. Plioplateia shares the tall body and subchelate gnathopods of ceinids, and in many other characters (to follow) appears to have a foundation in Taihape J. L. Barnard (1972a) or Waitomo J. L. Barnard (1972a): size and flagellar condition of antennae in Waitomo; posterior dorsal cuspidation of Waitomo; molar of Waitomo; outer plate of maxilla 1 in both genera; maxillipedal palp of Taihape; gnathopods (less giant setae) and telson of either genus. Plioplateia differs from Ceinidae in the dorsoventrally depressed head with complex cuspidation, the anterodorsal body cuspidation, the cuspidation and excavation of coxae and antennae, the special form of maxilla 2, the lack of REDESCRIPTION OF PLIOPLATEIA K. H. BARNARD 49 large setae on the gnathopods and the presence of inner lobes on the lower lip, wholly unique to Pliop/ateia in the Ceinid—Phliantid scheme. The reduction in segmentation on the pleopods suggests that Plioplateia has progressed far from a perfect intergrade between ceinids and phliantids and this is also supported by the extra cuspidation on the head and coxae, the development of inner lobes on the lower lip, the reduction in size of the inner lobe on maxilla | and the outer plate of the maxilliped, the enlargement of the molar, the loss of giant setae on the gnathopods. The brood plates appear to be much larger than in ceinids and phliantids whereas the gills are much smaller. The weakly developed multispination on the apices of uropods 1-2 would appear to be a remnant of ancestry in ceinids where these spines are well developed. The Kuriidae, composed only of Kuria Walker & Scott (1903), from Abd-el-Kuri, appear also to be in a level of evolution similar to Plioplateia. For example, the gnathopods of Kuria are subchelate, though weakly, the palms being almost transverse, and the outer plate of the maxilliped is small as in Plioplateia. Plioplateidae differ from Kuriidae in the ornamentation of the head, body and coxae, the small coxa 4, small article 2 of pereopods 5-7, expanded hands of the gnathopods, the disparity in sizes of antennae, and the extremely large mandibular molar. In minor ways, Plioplateidae differ from Kuriidae in the larger dactyls of the maxillipeds, and the absence of a ramus on uropod 3. Pleopods, maxillae and lower lip have not been reported for Kuria. Plioplateia K. H. Barnard Plioplateia K. H. Barnard, 1916: 155. Type species Plioplateia triquetra K. H. Barnard, 1916 (monotypy). Diagnosis With the characters of the family. Description Rostrum well developed, erect, thorn-like, bearing bilateral subsidiary tooth. Antenna | much longer than antenna 2, flagella of both pairs highly articulate, those of antenna | bearing 1-2 aesthetascs each. Epistome rounded anteriorly, upper lip deeply incised, asymmetrical. Mandibular incisor well toothed, right lacinia mobilis, if present, composed of three fused spines (or these actually rakers and lacinia mobilis absent), left lacinia mobilis large and well toothed, left mandible with three raker spines; molar very large, broad, blunt, tumid, poorly triturative, mainly granular apically; palp absent or possibly represented by small leaf. Lower lip with well-developed inner lobes. Outer plate of maxilla 1 with eight spines, palp uniarticulate, of medium size, bearing one medium apical seta. Inner plate of maxilla 2 much shorter and 50 ANNALS OF THE SOUTH AFRICAN MUSEUM narrower than outer, inner subconical, outer subrectangular, both poorly setose but inner with one medial and outer with two lateral setae. Inner plate of maxillipeds leaf-like, with tapered base, poorly armed, outer plates as large as inner; palp thin, article 1 elongate, article 4 unguiform, greatly elongate, with three apical setae. Pleurae of pereonites produced, rugose, humped or cuspidate. Plioplateia triquetra K. H. Barnard Figs 1-4 Plioplateia triquetra K. H. Barnard, 1916: 156, pl. 26 (figs 18-24); Griffiths, 1974: 328. Diagnosis With the characters of the family and genus. Description Head depressed but with erect thorn-like, rostrum with smaller basal tooth on each side, lateral lobes with weak dorsal and strong ventral tooth, middle of lobe bulging laterally and containing small but multifaceted ommati- dial eye, antenna | inserted by sleeve into pocket anterior to ocular lobe, anteroventral area of head extended forward through fusion of articles 1 and 2 of antenna 2, lateral surface of article 2 with large cusp, article 3 also with large lateral cusp, gland cones emerging ventrally from fused article 2. Mouthpart field from lateral view, apart from maxillipeds, dominated by outer surface of lower lip. Antenna | elongate, articles 1 and 2 cuspidate, article 3 shorter than article 1 of flagellum, latter 12-articulate, each article of flagellum with 1-2 aesthetascs and several curled setae. Accessory flagellum absent but marked by weak sinuate stripes inside of article 3. Antenna 2 small and slender, article 4 weakly cuspidate, article 5 slightly longer than article 4 of peduncle on article 1 of flagellum, latter 9-articulate, with short stiff curled setae. Upper lip scarcely distinct from epistome, together rounded anteriorly, upper lip deeply bilobed. Each mandibular molar with large setule, right incisor with 8-9 teeth, left with 3, left lacinia mobilis with 7 teeth, right either absent or formed of 3 fused spines, left mandible with 3 distinct rakers each independent and mostly fused to mandible. Mandibular lobes of lower lip well developed, inner lobes distinct, thin across faces, broad, widely separating outer lobes. Inner plate of maxilla | linguiform, of medium size. Wrist of gnathopod | longer than hand, shorter on gnathopod 2, neither lobate, palm well developed, oblique, defined by pair of spines, armed with pairs of wire-setules. Coxa 1 apically expanded, with deep posteroventral notch, coxae 2-4 somewhat tapered, each with weak or moderate notch, coxa 4 smaller than coxa 1, not excavate posteriorly; coxae 5-7 short, coxae 5 and 7 bilobed and acuminate, coxa 6 trilobed and acuminate. REDESCRIPTION OF PLIOPLATEIA K. H. BARNARD 51 Fig. 1. Plioplateia triquetra K. H. Barnard. h, holotype, female ‘h’ unmeasured; p, male ‘p’ 5,72 mm. A. Prebuccal outline, left lateral. B. left mandibular rakers. C. Ventral view of head, upper lip hatched; a, antenna 2, g, gland cone. D. Head, lateral; a, antenna 2 broken off. E. Prebuccal, left lateral. F. Body, scale = 1 mm. G. Right mandibular rakers. H. Head. I. Pleon, left lateral; u, uropod 3. J. Upper lip, anterior. 52 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 2. Plioplateia triquetra K. H. Barnard. male ‘p’ 5,72 mm. A. Antenna 2. B. Antenna 1. C. Left mandible. D. Mouthparts, left side; e, epistome, i, lower lip, 1, upper lip, m, mandible, x, maxilla. E. cuticle. F. Right mandible. G. Left mandible. H. Lower lip, oral side. I. Outer plate of maxilla 1. J. Maxilla 1. K. Palp of maxilliped, flattened. L. Maxilla 2. M. Lower lip, aboral side. N. Maxilliped. REDESCRIPTION OF PLIOPLATEIA K. H. BARNARD 53 Fig. 3. Plioplateia triquetra K. H. Barnard. male ‘p’ 5,72 mm, 0, outer ramus. A. Gnathopod 1. B. Gnathopod 2. C. Pleopod 1, rear. D. Pleopod 2, rear. E. Pleopod 3, rear. F. Pereo- pod 7. G. Pereopod 5. H. Pereopod 6. I. Pereopod 4. 54 ANNALS OF THE SOUTH AFRICAN MUSEUM Article 2 of pereopods 3-4 uncuspidate, that of pereopods 5—7 with bicuspi- date posteroventral process; defining armaments on article 6 composed of setae. Coxal gills very small, attached to segments 2-6, somewhat triquetral (see appendages attached to figures of coxae 2-4). Brood plates very large, lamellar, attached to coxae 2-5, densely furnished with coil-tipped setae. Male penes of pereonite 7 highly lateral, just basal to coxae, very elongate and sausage- shaped. Epimera tapering distally, poorly armed and unornamented. Pleopods tightly clumped, decreasing in size from front to rear slightly, inner rami scarcely shortened, each with 2 apical setae longer than ramus, except inner ramus of pleopod 3 with only one apical seta, peduncles elongate, only pleopod 3 with pair of apicomedial coupling spines. Urosomite 3 vestigial, represented only by ventral plate, telson on dorsal side appearing attached directly to urosomite 2, telson formed of bifid cowl lacking macroscopic armament. Uropods 1-2 short, stout, poorly armed, outer Fig. 4. Plioplateia triquetra K. H. Barnard. male ‘p’ 5,72 mm, g, gill, o, outer ramus. A. Uropod 3. B. Uropod 2. C. Coxa 2. D. Coxa 1. E. Coxa 3. F. Pleonite 6, ventral, telson hatched. G. Coxa 4. H. Uropod 1. I. Uropod 1, rami enlarged. J. Uropod 2. K. Pleonites 1-2 dorsal showing telson, t. REDESCRIPTION OF PLIOPLATEIA K. H. BARNARD yes ramus of uropod | shortened, each ramus of both uropods with apical jewel-nail plus accessory spine except on inner ramus of uropod 2, inner ramus of uropod 1 with third spine fixed subapically. Uropod 3 ovate, bearing weak apical spinule. Cuticle densely pebbled, each pebble usually complex, either with apparent pit or appearing ring-shaped (with central vacuole) bulbar setules sparse, pipes often abundant; no pits typical of ceinids have been observed. Pereonite 1 with large tooth dorsally, remaining pereonites each with single tooth or hump, pleonites 1-3 each with increasingly smaller dorsal hump, urosomite | with slightly larger hump, this segment covering urosomites 2-3 dorsally; pereonites 1-7 from front to rear with increasingly complex distolateral rugosities or cusps. Holotype South African Museum A174, female ‘h’ (not measured to prevent damage) lacking right antennae, uropods 2-3, telson, right uropod 1. Other material South African Museum, University of Cape Town SCD 310 F, male ‘p’ 5,72 mm (dissected and illustrated herein). Distribution South Africa, 50-91 m. ACKNOWLEDGEMENTS I thank Dr C. L. Griffiths of the C.S.I.R. Oceanographic Research Unit, University of Cape Town, for locating this material, and Dr T. H. Barry Director of the South African Museum, for his kind assistance. Carolyn L. Cox of Smithsonian Institution inked and prepared the illustrations for publication; she also created several of the original drawings. REFERENCES BARNARD, J. L. 1972a. Gammaridean Amphipoda of Australia, Part I. Smithson. Contr. Zool. 103: 1-333. BARNARD, J. L. 19726. The marine fauna of New Zealand: Algae-living littoral Gammaridea (Crustacea Amphipoda). Mem. N.Z. oceanog. Inst. 62: 7-216. BARNARD, K. H. 1916. Contributions to the crustacean Fauna of South Africa. 5.—The Amphipoda. Ann. S. Afr. Mus. 15: 105-302. 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. 65: 251-336. GriFFITHS, C. L. 1975. The Amphipoda of southern Africa. Part 5. The Gammaridea and Caprellidea of the Cape Province west of Cape Agulhas. Ann. S. Afr. Mus. 6: 91-181. WaLker, A. O. & A. Scott. 1903. Decapod and sessile-eyed crustaceans from Abd-el-Kuri. Nat. Hist. Sokotra and Abd-el-Kuri: 216-232. 7 dt ‘oe oo » ee aon pe. HRA G: @ 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, 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.’ 5 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. J. LAURENS BARNARD REDESCRIPTION OF PLIOPLATEIA K. H. BARNARD, A GENUS OF AMPHIPOD (CRUSTACEA) FROM SOUTH AFRICA , f Cone VOLUME 77 PART 4 DECEMBER 1978 ISSN 0303-2515 MUS. COMP. ZOOL. LIBRARY FEB 15 1979 HARV ARD 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. 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, 19694) 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. 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) ee Se ie ee — ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 77 Band December 1978 Desember Part 4 Deel CRETACEOUS FAUNAS FROM ZULULAND AND NATAL, SOUTH AFRICA A NEW GENUS AND SPECIES OF GASTROPLITINAE 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, 21-3, 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, 31(1-3), 32(5), 33 Copyright enquiries to the South African Museum Kopieregnavrae na die Suid-Afrikaanse Museum ISBN 0 908407 59 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 CRETACEOUS FAUNAS FROM ZULULAND AND NATAL, SOUTH AFRICA A NEW GENUS AND SPECIES OF GASTROPLITINAE FROM THE MZINENE FORMATION (ALBIAN) By WILLIAM JAMES KENNEDY Geological Collections, University Museum, Oxford & HERBERT CHRISTIAN KLINGER South African Museum, Cape Town (With 7 figures) LMS. accepted 20 September 1978] ABSTRACT Alopecoceras gen. nov., type species Alopecoceras ankeritterae sp. nov., is a unique Southern hemisphere homoeomorph of the Boreal genus Neogastroplites McLearn, 1930, which occurs in the low Middle Albian Mzinene Formation (Albian III) of the Mzinene River, Zululand. Whereas Neogastroplites evolved from Gastroplites McLearn, 1930, by acquisition of umbilical and ventral nodes and is of late Albian age, A/opecoceras is believed to be derived from Hatchericeras Stanton, 1910, by a similar acquisition of nodes. The genus also shows similarities to Tetrahoplitoides Casey, 1954. CONTENTS PAGE AMtrOdUCHION: so we ee ee ews) on) ST Systematic Palaeontology . . . . 58 Acknowledgements . . .. . . 68 References ay eee Pee ee Le 68 INTRODUCTION The Mzinene Formation exposed along the Mzinene River | 200 m north- east of the farm Amatis, north of Hluhluwe, Zululand (locality 35 of Kennedy & Klinger (1975: 28), 27°58’03”S, 32°18'34’E) has yielded a series of remarkable specimens which closely resemble the exclusively boreal late Albian ammonite genus Neogastroplites McLearn, 1930 (see Reeside & Cobban 1960 for extensive illustration of the genus) in association with a fauna of low Middle Albian age in part described by the authors in previous publications (Kennedy & Klinger 1977, 1978, in press; Klinger 1976; Klinger, Wiedmann & Kennedy 1975), and including Carinophylloceras collignoni Klinger, Wiedmann & Kennedy, Ammo- noceratites (Ammonoceratites) mahadeva (Stoliczka), Anagaudryceras buddha Ann, S. Afr. Mus. 77 (4), 1978: 57-69, 7 figs. 57 58 ANNALS OF THE SOUTH AFRICAN MUSEUM (Forbes), Protanisoceras (Rossalites) aff. superbus Collignon, Lyelliceras and Douvilleiceras species. Further study has shown that the material represents a new genus and species which is a heterochronous homoeomorph of Neogastroplites, and is described below as Alopecoceras ankeritterae sp. nov. All specimens are housed in the British Museum (Natural History) and the Geological Survey of South Africa, designated BMNH and SAS respectively. SYSTEMATIC PALAEONTOLOGY Superfamily HOPLITACEAE H. Douvillé, 1890 Family Hoplitidae H. Douvillé, 1890 Subfamily Gastroplitinae Wright, 1952 Genus Alopecoceras Kennedy & Klinger gen. nov. Type species Alopecoceras ankeritterae gen. et. sp. nov., low Middle Albian, Zululand. Derivation of name Alopex (Greek) fox. Diagnosis Initially depressed, whorl section trapezoidal with sparse conical or bullate umbilical nodes giving rise to pairs of ribs with additional ribs intercalated; all ribs bearing conical ventral tubercles and connected across the fastigiate venter by a much thickened rib. Whorl section becomes compressed and lanceolate in later growth, venter narrowly rounded and ornament reduced to low falcoid ribs with interspaces sometimes accentuated into feeble constrictions. Suture line with moderately incised bifid elements. Discussion Alopecoceras closely resembles Neogastroplites McLearn, 1930, but is of low Middle Albian age whereas that genus, a descendant of Gastroplites McLearn, 1930, is of late Albian age. Because of the great intraspecific variation in Neogastroplites (see Reeside & Cobban 1960) it is a little difficult to make overall statements of difference, but in general Alopecoceras is consistently more evolute, tubercles are lower on the flank, ribs fewer and less flexuous in juveniles, the ventral ribbing thicker and less convex. Neogastroplites typically develops a siphonal tubercle and is rostrate; neither of these features are seen in Alopecoceras, whilst the suture lines are more intricately subdivided. When adult, Neogastroplites may become smooth and very compressed (e.g. Reeside & Cobban 1960, pl. 17 (figs 17, 22), pls 18-19), but lack the CRETACEOUS FAUNAS FROM SOUTH AFRICA 59 flexuous branching ribs and feeble constrictions of A/opecoceras. Furthermore, these individuals have slender, feebly ornamented inner whorls; individuals of a similar degree of inflation to Alopecoceras ankeritterae retain strong ribs and nodes to a great size (Reeside & Cobban 1960, pl. 11 (figs 16, 20), pl. 12 (figs 11-12, 16), pl. 36 (figs 1, 5, 9-11)). Alopecoceras can be easily separated from Gastroplites McLearn, 1930 (see Reeside & Cobban 1960 pls 8-9), for that genus does not possess umbilical bullae and is usually more involute. Lemuroceras Spath, 1942, never develop strong umbilical nodes, shoulder tubercles or a fastigiate venter (see illustrations in Collignon 1963). Arcthoplites Spath, 1925 (see illustrations in Casey 1965, text-fig. 177), and Subarcthoplites Casey, 1954, both have strong, narrow ribs, round venters and lack tubercles, as does Cymahoplites Spath, 1922. Alopecoceras shows closer similarities to Tetrahoplitoides Casey, 1954 (see Fig. 7H—J); both have a trapezoidal whorl, although, in the type species at least, the whorls are slender, the umbilical tubercles are bullate and not conical as in Alopecoceras. The venter is narrower in Tetrahoplitoides, flat rather than fastigiate, and lacks ventrolateral tubercles. Evolutionary origins Alopecoceras occupies an isolated geographic position, and its evolutionary origins are far from obvious in the remaining faunas of both the South African and Madagascar Albian. C. W. Wright has, however, called our attention to its similarities to the genus Hatchericeras Stanton, 1901. Originally described from Patagonia, this genus was referred to the Neocomitinae Spath, 1924, in the Treatise, and regarded as of possibly Lower Hauterivian age. Subsequent publication by Leanza (1970) has show it to be an early Albian gastroplitinid. During middle growth (Fig. 7A—D) it differs from Alopecoceras very obviously in the lack of umbilical nodes and in possessing numerous flexuous ribs. There are, however, blunt thickenings of the ribs—incipient tubercles—at the ventro- lateral shoulder in some species, and a broad, blunt rib crosses the venter. Adults (Leanza 1970, fig. 3la—b) are compressed, with a rounded venter, feeble flexuous ribs and striae, plus feeble constriction-like folds. The sutures (Fig. 6C) of the two genera are also constructed on the same plan, with a similar degree of incision. The authors would conclude, therefore, that A/opecoceras is an offshoot of Hatchericeras which has developed prominent umbilical nodes, accentuated the ventral tubercles present in some of the latter, and developed a fastigiate rather than flattened venter: it is a homoeomorphous Southern hemisphere analogue of Neogastroplites rather than a close relative. Occurrence Alopecoceras is known only from the low Middle Albian of Zululand. 60 ANNALS OF THE SOUTH AFRICAN MUSEUM Alopecoceras ankeritterae gen. et. sp. nov. Figs 1-4, 5A-B, 6, 7E-G Holotype BMNH C81266 from locality 35, cliff and stream sections extending over several hundred metres along the Mzinene, approximately 1 200 m north-east of the farm Amatis, north of Hluhluwe, Zululand, 27°58’03”S 32°18’31"E Mzinene Formation, Albian III. Fig. 1. Alopecoceras ankeritterae sp. nov. The holotype, BMNH C81266. x 1. CRETACEOUS FAUNAS FROM SOUTH AFRICA Fig. 2. Alopecoceras ankeritterae sp. nov. Inner whorls of the holotype, BMNH C81266. 61 62 ANNALS OF THE SOUTH AFRICAN MUSEUM Paratypes BMNH C81267 to C81274, SAS H54a—b, SAS A584, all from the same horizon and locality as the holotype; SAS EM152 from locality 171, on the hill slopes west-south-west of Mlambongwenya Store, Zululand, Mzinene Formation, Albian II-III, and SAS A589 from locality 53, a derelict dam site on Indambana, south of Izwehelia Farm, north of Hluhluwe, Zululand, Mzinene Formation, Albian II. a Sa ey ae = 2 Fig. 3. Alopecoceras ankeritterae sp. nov. Body chamber of the holotype, BMNH C81266. x 1. CRETACEOUS FAUNAS FROM SOUTH AFRICA 63 Fig. 4. Alopecoceras ankeritterae sp. nov. Paratypes BMNH C81267-C81271. A-C, E-G, I-K, P-R x 2; D, H, L-O x 1. 64 ANNALS OF THE SOUTH AFRICAN MUSEUM C | A | D B Fig. SA-B. Whorl sections of Alopecoceras ankeritterae sp. nov. A. BMNH C81274. x 12. B. BMNH C81268. x 6. C—D. Early whorls of BMNH C81274. x 12. CRETACEOUS FAUNAS FROM SOUTH AFRICA 65 oo Fig. 6. A-B. Alopecoceras ankeritterae sp. noy. External sutures of BMNH C81268, x 12, and BMNH C81266, x 3. C. Hatchericeras semilaeve Leanza. External suture of a specimen in C. W. Wright's collection from Santa Cruz, Argentine. x 6. 66 ANNALS OF THE SOUTH AFRICAN MUSEUM Derivation of name The species is named for Anke Ritter of Gauting. Diagnosis As for genus. Dimensions All dimensions are in millimetres. D = diameter, Wb = whorl breadth, Wh = whorl height, U = umbilical diameter, R = ribs per whorl. Figures in parentheses are dimensions expressed as a percentage of diameter. D Wb Wh Wb:Wh U R Holotype BMNH C81266 .« §37,5(100) 35026) 61,2(45) 0,57 34,0(25) = at 97,9(100) 26,4(27) 42,8(44) 0,62 24,3(25) 32 at 67,2(100) 21,8(32). 30,0(45) 0,72 18,7(28) — SAS A584 .. «+ 41,9(100) 19,5(47) 18,5744) 1,05 10,224) @ BMNH C81267 .. 25,5(100) 11,8(46) 10,5(41) = ‘1,12 7,2(28) 26 SAS H54a => on “23,9(100) . 10,046) —10;9%46) — im 6,0(25) 26 BMNH C81268 .. 22,7(100) 10,4(46) 85738) - AL9 6,3(28) 26 Description The early developmental stages are seen in BMNH C81274. At a diameter of 3,8 mm, the juvenile shell is cadicone, the umbilicus comprising approxi- mately 30 per cent of the diameter, with a whorl breadth to height ratio of 1,4 (Fig. SC-D). Ornament consists of blunt, low, rounded umbilical nodes, which first appear at a diameter of c. 3 mm. These occur at a rate of seven per whorl up to a diameter of 6 mm, and in this interval, low, broad folds pass across the flanks and venter, although never developing into clearly delineated ribs. By 6 mm diameter, the whorl breadth to height ratio has become 1,2. From 6 mm (Fig. 4A—-R) the whorl section changes from depressed and reniform to trapezoidal. The umbilicus is shallow, and the wall slopes gently outwards. The greatest breadth is at the umbilical bullae, and the whorl! breadth to height ratio is commonly reduced to 1,!. In intercostal section the outer flanks are flattened, the ventrolateral shoulders broadly rounded and the venter very broadly rounded. In costal section, the flanks are concave at mid-flank and flattened on the outer flank, and the venter fastigiate. There are eight to ten prominent umbilical bullae per whorl, placed well out from the umbilical seam, and these give rise to groups of two or three broad ribs, weakened at mid-flank, where additional short ribs are intercalated, and recti- to feebly rursiradiate in some cases. All ribs bear a bullate to conical tubercle at the ventrolateral shoulder, and there are commonly twenty-five to twenty-eight ribs per whorl. The ribs pass across the venter with a faint con- vexity, broadening and diminishing in elevation over the siphonal line. i CRETACEOUS FAUNAS FROM SOUTH AFRICA 67 Fig. 7. A-D. Hatchericeras semilaeve Leanza. Inner whorls of a specimen in C. W. Wright's collection from Santa Cruz, Argentina. x |. E-G. Alopecoceras ankeritterae sp. nov. Para- type, SAS H93/3. x 1. H-—J. Tetrahopliloides stantoni (Anderson) cast of the holotype, from Texas Springs, Shasta County, California. H = 1, I-J x 2. 68 ANNALS OF THE SOUTH AFRICAN MUSEUM This general style of ornament extends to a diameter of c. 50 mm; as size increases, the whorls eventually change from depressed to compressed, the ribs coarsen, and, on the venter, become very thick (Fig. 2C). There is some variation in strength and number of ribs, bullae and whorl section at this stage, as can be seen from the figures. The features of later growth are shown by the holotype (Figs 1-3). Here, from 50 mm onwards, all ornament declines, the ribs become low, broad and flexuous, arising in pairs from the bullae, whilst the whorl becomes compressed and lanceolate in section with a narrowly rounded venter which is crossed by ribs and has, as a result, an undulose profile. On the mature body chamber (Fig. 3), coiling has become more evolute, with a low, outwards sloping, flattened umbilical wall. There is a somewhat abrupt umbilical shoulder and high, lanceolate whorls (whorl breadth to height ratio is 0,57 to 0,62) with a narrowly arched venter. Ornament consists of irregular falcoid ribs and striae, arising from feeble umbilical bullae, separated by interspaces which may be strengthened into feeble, irregular constrictions. The suture line (Fig. 6B) at maturity is moderately divided. E/L is broad and asymmetrically bifid, L broad and asymmetrically bifid. L/U, is large, although narrower than E/L, and asymmetrically bifid. U, is narrow and quite deeply divided. Discussion Features which separate Alopoceceras ankeritterae from species of other genera are fully covered in comments in the generic discussion on p. 58. Occurrence Low Middle Albian of Zululand only. ACKNOWLEDGEMENTS Our best thanks are due to Mr C. W. Wright for drawing our attention to the similarities between Alopecoceras and Hatchericeras, as well as providing useful discussion. We are also grateful to Dr M. R. Cooper (Oxford), Dr M. K. Howarth and Mr D. Phillips (British Museum, Natural History) and the staff of the Geological Collections, Oxford, and the South African Museum for technical assistance. REFERENCES Casey, R. 1965. A Monograph of the Ammonoidea of the Lower Greensand. Palaeontogr. Soc. (Monogr.) 6: 399-546. CoLLIGNoN, M. 1963. Atlas des Fossiles caractéristiques de Madagascar (Ammonites), Fasicule X (Albien). Tananarive: Service Géologique. KENNEDY, W. J. & Kuincer, H. C. 1975. 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 Phylloceratidae. Bull. Br. Mus. nat. Hist. (Geol.) 27: 347-380. CRETACEOUS FAUNAS FROM SOUTH AFRICA 69 KENNEDY, W. J. & KLINGER, H. C. 1978. Cretaceous faunas from Zululand and Natal, South Africa. The ammonite family Lytoceratidae Neumayr, 1875. Ann. S. Afr. Mus. 74: 257-333. 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.). KLINGER, H. C. 1976. Cretaceous heteromorph ammonites from Zululand. Mem. S. Afr geol. Surv. 69: 1-142. KLINGER, H. C., WIEDMANN, J. & KENNEDY, W. J. 1975. A new carinate phylloceratid from the early Albian (Cretaceous) of Zululand, South Africa. Palaeontology 18: 657-644. LeANZA, A. F. 1970. Ammonites nuevos o pocos conocidos del Aptiano, Albiano y Ceno- maniano de los Andes Australes, con notas acera de su posicion estratigraphica. Revta Assoc. geol. argent. 25: 197-261. ReesipE, J. B. & CoBBAN, W. A. 1960. Studies of the Mowry Shale (Cretaceous) and con- temporary formations in the United States and Canada. Prof. Pap. U.S. geol. Surv. 355: 1-126. 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, 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 NEW GENUS AND SPECIES OF GASTROPLITINAE FROM THE MZINENE FORMATION (ALBIAN) OLUME 77 PART 5 DECEMBER 1978 MUS. COMP. zoo. !SSN 0303-2515 LIBRARY FEB 15 1979 HARVARD UNIVERSITY ANNALS OF THE SOUTH AFRICAN MUSEUM U-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 ) 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; ie mar ‘As described (Haughton & Broom se ‘As described (Haughton ef a/. 1927) . Note: no comma separating name ak 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, 5, etc. to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 19694) 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) BuLLouGu, 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., DuvaLt, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. gen. 74: 627-634. Konun, A. J. 1960a. Ecological notes = ead (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309. Konn, A. J. 19606. Spawning hate te oe egg masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51. Tues, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: 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 77 Band December 1978 Desember Patt 5 Deel PPreR CRETACEOUS SEDIMENTS FROM THE IGODA RIVER MOUTH, EAST LONDON, SUU TE APRICA By HERBERT CHRISTIAN KLINGER & BRIAN E. LOCK 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.), 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), 32(5), 33 Copyright enquiries to the South African Museum Kopieregnavrae na die Suid-Afrikaanse Museum ISBN 0 908407 61 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, UPPER CRETACEOUS SEDIMENTS FROM THE IGODA RIVER MOUTH, EAST LONDON, SOUTH AFRICA By HERBERT CHRISTIAN KLINGER South African Museum, Cape Town & BRIAN E. LOCK, The University of Southwestern Louisiana, Lafayette (With 7 figures) [MS. accepted 20 September 1978] ABSTRACT The name Igoda Formation is proposed for a sedimentary sequence consisting mainly of calcareous sandstones and arenaceous limestones exposed on the bluffs overlooking the Igoda River Mouth near East London, South Africa. The invertebrate fauna indicates a Late Cretaceous age, probably Late Campanian to Early Maastrichtian. Faunal paucity precludes detailed comparison with other areas in southern Africa, but common faunal elements occur at Lower Needs Camp (here regarded as a lateral facies equivalent of the Igoda Formation), Pondoland, Zululand, Madagascar and Angola. Affinities with Madagascar are strongest. CONTENTS PAGE IMtTOMUCIONNE Ee kes ee ees ee fe) aes 71 The Igoda River Mouth section. . . ... . 73 ithe roadsection™. = = | me Mew eet ek is) Fauna and age of the Igoda Formation ... . 75 Comparison with other areas in southern Africa . . 81 SURIUMAY; Mee ooo SP ee a ty Gee a) ce. Bech 82 Acknowledgements =. . 5 6 4 «a 83 IREfEDeD CES Twat Pe un ei tcweils, Say Wee eters 83 INTRODUCTION Late Cretaceous fossiliferous limestones have long been known from the eastern Cape from one of the small quarries on the farm Needs Camp (the East or Lower Quarry) near East London (Lang 1908; Woods 1908; Chapman 1916). McGowran & Moore (1971) established a probable Upper Senonian (Campanian to Maastrichtian) age for this deposit on the basis of the micro- faunal content. Contrary to a report by King (1972), the second quarry at Needs Camp (the West or Upper Quarry) is excavated in limestones of Tertiary age (Lock 1973 and in preparation). Microfaunal investigations confirm this Ann. S. Afr. Mus. 77 (5), 1978: 71-83, 7 figs. 71 TZ ANNALS OF THE SOUTH AFRICAN MUSEUM (W. G. Siesser pers. comm.), although reworked Cretaceous material occurs in the limestones. Until 1974 no other outcrops of Late Cretaceous marine sediments had been reported from south of the Umzamba Estuary (Pondoland, Transkei). Mountain (1974: 19) briefly described a new fossiliferous locality less than 15 km from Needs Camp as follows: The rock occurs in a ‘tiny exposure on the East London—Kidd’s Beach road just west of the Goda River valley as the road climbs up on to the coastal plain. It can be seen in the present road cutting and consists of a friable limestone with abundant echinoid spines, Ostrea shells and gastropod casts. It is possibly Cretaceous but no one has examined the microfossils.’ Following landslides during the last few years, an extension of this occur- rence became exposed in the bluffs overlooking the western shore of the lagoon at Igoda Mouth, less than a kilometre from the roadside section mentioned by Mountain (Fig. 1). Tertiary and ?Pleistocene calcareous sands Igoda Formation Beaufort sedimentary rocks and Karroo dolerite Fig. 1. Locality map, Igoda Formation. A. Roadside exposure reported by Mountain (1974); note the new road under construction. B. Type section of the Igoda Formation. C. Reference section. (See also Fig. 2.) UPPER CRETACEOUS SEDIMENTS FROM THE IGODA RIVER MOUTH te THE IGODA MOUTH SECTION The succession in the new exposures is summarized in Figure 2. The Cretaceous sequence, here named the Igoda Formation, rests uncon- formably on sandstones and mudstones of the Permian/Triassic Beaufort Group, which have been intruded by dolerites of later Karoo age. The contact is one of some relief. The type section (sections BI, B2) is a composite one (see Fig. | for location and Fig. 2 for sections), and has at its base up to 50 cm of matrix- supported small-pebble conglomerate, comprising well-rounded and well- sorted brown-stained pebbles with a mean diameter of about 2 cm but with individual pebbles up to about 5 cm long. Pebble lithologies consist of vein- quartz and quartz arenites for the most part, although one pebble was a single, well-rounded crystal of orthoclase 1,2 cm long. The matrix is an arenaceous limestone with abundant glauconite. Some shell fragments are present in this limestone matrix. The basal conglomerate passes up into a sequence of arenaceous limestones and calcareous sandstones, all with a high content of glauconite. As this section is followed up the slope, exposures become very poor, and only a few of the more resistant calcareous beds form ledges. At an altitude some 20 m above the small-pebble conglomerate, a second unconformity, at the base of the overlying Alexandria Formation (of probable Tertiary age), is reached. The middle portion of the Igoda Formation is better exposed some 30 m to the south. In this second section a brown, small-pebble conglomerate, indistinguishable from that just described, lies at the base of a similar sequence of well-exposed glauconitic arenaceous limestones and calcareous sandstones. As in the first section, these strata are fossiliferous, yielding a shelly fauna dominated by ostreids which litter the surface. At this point, however, the calcareous sequence has a much reduced thickness of just over 7 m, and the small-pebble conglomerate overlies a 3-metre-thick sequence of white, poorly consolidated, unfossiliferous sandstone and white, small-pebble conglomerates which are in all other respects identical to the brown ones already described. These three lithological associations are regarded as informal members, and are known as the ‘white member’, ‘brown, small-pebble conglomerate member’ and ‘calcareous member’ respectively. The base of the Alexandria Formation is marked by another conglomerate consisting of much larger pebbles (up to 30 cm in diameter) mostly of Beaufort Group sandstone and siltstone. These pebbles are generally less resistant than those found in the conglomerates of the Igoda Formation. The matrix of the Alexandria Formation basal conglomerate is a coarse bioclastic limestone containing fossil gastropods, especially Patella sp. and Conus sp., usually as moulds. Glauconite is absent to rare. This unit is about 1,5 m thick and is overlain by 10 m of cross-bedded, well-indurated, coarse bioclastic limestones. Characteristically, the cross-bedding comprises a single tabular set, with seaward dips of about 25°. Above this scarp-forming unit is a considerable thickness (at least 30 m) of poorly consolidated calcareous aeolian sands. ANNALS OF THE SOUTH AFRICAN MUSEUM 74 C] ‘Sty Osye 99g) “y7eW J9}eM YsIY A[oJeUITXOIdde ‘[OAo] 19}eM UOOSR] SAO SOIJOU UI OB sopnyn[y “(MOjaq) suOT}OES S1Yde.ASIVvI}S pue (QAOQK) 1X9} Ul Poqiiosop Ssdoidjno JO UOTINGIAWsSIP SUIMOYS ‘UOCOSL] BPOST JO yURQ UID}SOAA “7 “BIT ~ 330) JOQGUIBUWI BLIWUM ——— saquaw a3e,0W046 “| -—uod uMojq sequiow snoasesjeo NOILVINHOA4 VdODOI PoeO| @3e190W0/6u09 ee ayiuase9je9 NOILVINHOS VIYQONVX3IT1V UPPER CRETACEOUS SEDIMENTS FROM THE IGODA RIVER MOUTH 75 A second section was measured, 500 m closer to the sea (section C). Here the white member is no longer present, and the calcareous member is even further reduced in thickness (5,5 m). In other respects the sequence is similar (see Fig. 2). THE ROAD SECTION The original outcrop described by Mountain (1974, see above) was identi- fied, and the adjective ‘tiny’ found to be very appropriate. Only a few tens of centimetres of very weathered glauconitic arenaceous limestone can be seen overlying Beaufort Group sandstones and merging upwards into soil and surface debris in the bank at the inland side of the road. The outcrop is very overgrown and easily overlooked. Broken specimens of Rhynchostreon decussata and Lopha spp., common forms in the main outcrop area, establish a correlation with the lagoon-side exposures. At the time of the most recent visit (May 1977), roadwork was in progress for the straightening and improving of the main coastal road. Where this new road reaches the approximate altitude of Mountain’s outcrop, it runs about 100 m inland of the latter, and cuts through a mass of dolerite. At the top of the cutting at this point, a few rounded pebbles of Beaufort Group sandstone in a calcareous cement were found. The size of these pebbles (20-30 cm) suggests that they are remnants of the conglomerate from the base of the Alexandria Formation. This would imply that the Igoda Formation pinches out at this point (area A on map, Fig. 1). FAUNA AND AGE OF THE IGODA FORMATION In comparison with the Cretaceous sediments of Zululand and Transkei (Kennedy & Klinger 1975 onwards), the invertebrate fauna of the Igoda Forma- tion is meagre and poorly preserved. Bivalves are the commonest group, especially the ostreid forms, which have been preserved by virtue of their unique shell mineralogy, followed by brachiopods, baculitid ammonites, echinoids, and rare, normally coiled, ammonites, in that order of abundance. At present, the material in the authors’ collections is too scant to merit formal description and discussion, but a preliminary examination of the fauna has revealed the presence of the following faunal elements: FORAMINIFERA Textularia sp. COELENTERATA “Caryophyllia’ cf. arcotensis Forbes (Fig. 3) ECHINODERMATA Unidentifiable cidarid with uniserial pore pairs (Fig. 4B) Crinoid stem ossicles of Pentacrinus type (Fig. 4A) 76 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 3. ‘Caryophyllia’ arcotensis Forbes (Geology Department, Rhodes University.) A-B. Lateral view. Scale bar 0,5 cm long. C. Dorsal view, stereopair. Scale bar 0,5 cm long. UPPER CRETACEOUS SEDIMENTS FROM THE IGODA RIVER MOUTH tif Fig. 4. A. Crinoid stem ossicle of the /socrinus type. B. Cidaris sp. indet. Scale bars 0,5 cm long. MOLLUSCA—CEPHALOPODA Baculites subanceps Haughton (Fig. 5) Eupachydiscus ? sp. (Fig. 6) Pachydiscid sp. indet (compressed) Saghalinites sp. cf. S. cala (Forbes) (Fig. 7A—B) MOLLUSCA—GASTROPODA Turritella (Zaria) cf. T. (Z.) besairiei Basse MOLLUSCA-BIVALVIA Rhynchostreon decussata (Goldfuss) Lopha (Actinostreon) schnaebelei Basse (Fig. 7C—H) ‘Trigonia sp. Spondylus douvillei Basse Panopea cf. orientalis (Forbes) ‘Inoceramus’ spp. BRYOZOA cf. Ceriopora micropora Goldfuss “Membranipora’ cf. plebicola Brydone BRACHIOPODA cf. Terebratulina relicta Stoliczka cf. Terebratula manuaensis Muir-Wood cf. Rhynchonella natuans Stoliczka Eolacazella affine (Bosquet) 78 ANNALS OF THE SOUTH AFRICAN MUSEUM J K Fig. 5. Baculites subanceps Haughton. A, C. SAM-PCI5721. x 1,2. B. SAM-PCI5723. « 1,0. D. SAM-—PCI5728. « 1,0. E-H. SAM-—PCI5906. x 1,0. I-L. SAM-PCI5720. x 1,0. UPPER CRETACEOUS SEDIMENTS FROM THE IGODA RIVER MOUTH 79 Fig. 6. Eupachydiscus ? sp. indet. SAM-—PCI5719. x 1,0. Of the species identified, only a few can be used for accurate dating of the Igoda Formation. Baculites subanceps s.s. has been firmly dated as Late Campanian in Angola (Howarth 1965), whilst the Pacific subspecies B. anceps pacificus occurs in the Late Campanian of Japan and California. Saghalinites ranges from the Santonian to Maastrichtian stages of the Late Cretaceous (Kennedy & Klinger 1977). S. cala, which the Igoda specimen resembles most, ranges from Cam- panian IV to Maastrichtian II (sensu Kennedy & Klinger 1975) in Zululand but is also known to occur in slightly older sediments in Pondoland (Transkei) 80 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 7. A-B. Saghalinites sp. cf. S. cala (Forbes). C-H. Lopha (Actinostreon) schnaebelei (Basse). C. SAM-PCI5773. x 1,0. D. SAM-PCI5749. x 1,0. E. SAM-PCI5736. x 1,0. F. SAM-PCI5750. x 1,0. G. SAM-PCI5752. x 1,0. H. SAM-PCI5744. x 1,0. UPPER CRETACEOUS SEDIMENTS FROM THE IGODA RIVER MOUTH 81 (Middle Santonian—Lower Campanian) and southern India (Santonian or Campanian). The ammonite genus Eupachydiscus is typically Late Cretaceous, ranging from Coniacian to Maastrichtian, but the Igoda material is too poorly preserved for specific identification. Ceriopora micropora Goldfuss has been described from the type locality of the Maastrichtian stage, but is also common in the Santonian and Campanian. M. plebicola is from the Maastrichtian. Eolacazella affine (Bosquet) occurs in the Maastrichtian of Western Europe, and was also found at the Lower Needs Camp Quarry now dated as Upper Campanian/Lower Maastrichtian by McGowran & Moore (1971). These data all seem to suggest an age of Campanian/Maastrichtian for the Igoda Formation with only a slight possibility of Santonian elements present. COMPARISON WITH OTHER AREAS IN SOUTHERN AFRICA The nearest onshore sediments of similar age occur at the Lower or Eastern Quarry at Needs Camp. Eolacazella affine occurs at both the Igoda River Mouth and at Lower Needs Camp, thus suggesting the two outcrops to be temporal equivalents, though of different lithologies. In places, the outcrops at Lower Needs Camp consist virtually of bryozoan limestone only. Exposures of similar lithologies have since been found by one of the authors (H. C. K.) in roadside excavations approximately half-way between the Upper and Lower Quarries, illustrating that the Lower Needs Camp lithology has a much greater aerial extent than previously suspected, and may yet prove to be a mapable unit to conform with the requisites for formal recognition as a separate Formation by the South African Committee for Stratigraphy. The Lower Needs Camp sediments probably grade laterally into the slightly deeper water facies of the Igoda Formation, but outcrops connecting the two areas are, as yet, unknown. Along the east coast of southern Africa, the closest Senonian sediments occur at the Umzamba Estuary (Transkei), which have been firmly dated as Mid-Santonian to Lower Campanian (Klinger & Kennedy 1977). Rare Saghali- nites cala in the Umzamba Formation are comparable with Saghalinites sp. in the Igoda Formation. The closest, and thus far only known, onshore Upper Campanian/Lower Maastrichtian sediments on the east coast of southern Africa occur in the False Bay/St Lucia region of Zululand (Kennedy & Klinger 1975). Here Saghali- nites cala occurs quite frequently with Nostoceras sp. and Eubaculites sp. in the Upper Campanian and Lower Maastrichtian respectively. S. cala appears to be the only identifiable faunal element in common between the two areas. Lower Maastrichtian sediments are known from Cheringoma further north in Mozambique, but to date only poorly-preserved Eubaculites species are known (Crick 1923). Much further north, the Campanian and Maastrichtian faunas of Mainti- rano, Madagascar, as described by Basse (1931), strongly resemble those from 82 ANNALS OF THE SOUTH AFRICAN MUSEUM Igoda especially as far as the thick-shelled ostreid forms are concerned, and to a lesser extent with the Late Senonian fauna of the east coast Province of Vatomandry described by Cottreau (1922), which is characterized by the paucity of ammonites, similar to the Igoda Formation. Towards the southern and western coast, the closest sediments of com- parable age are found in the off-shore Alphard Group between Cape St Francis and Cape Recife (Klinger ef a/. in prep.). Here, however, the dominant faunal element is Eubaculites latecarinatus indicative of Early Maastrichtian age. No macrofossils are common to both outcrops. From here westwards, the first known Cretaceous on-shore sediments occur near Bogenfels (South West Africa) (Klinger 1977), but these have been dated definitely as Cenomanian, although some doubt still exists as to whether Senonian strata are present on-shore or not. The closest comparable deposits occur at Carimba in Angola, the type area of Baculites subanceps, the latter being the only faunal element in common between Igoda and Angola. The Cretaceous sediments at Carimba have been firmly dated as Late Campanian by Howarth (1965). SUMMARY The name Igoda Formation is proposed for a sedimentary sequence consisting mainly of calcareous sandstones and arenaceous limestones exposed on the bluffs overlooking the Igoda River Mouth. The Formation is informally divided into ‘white member’, ‘brown, small-pebble conglomerate member’ and ‘calcareous member’. In comparison with the Cretaceous sediments of Natal and Transkei, the Igoda fauna is meagre, consisting mainly of thick-shelled ostreid bivalves, followed by brachiopods, baculitid ammonites, echinoids, and rare, normally coiled ammonites, in that order of abundance. These faunas point to an age of Late Campanian to Early Maastrichtian. Due to the paucity of the faunas, comparisons with other temporally equivalent areas are tenuous. The Lower Needs Camp Quarry deposits are of similar age, and probably represent a shallower water and more restricted lateral facies equivalent of the Igoda Formation. Saghalinites sp. cf. cala connects the Igoda Formation to the Cretaceous deposits on the east coast of southern Africa at the Umzamba Estuary and in the False Bay/Lake St Lucia region of Zululand. As far as the abundance of thick-shelled ostreids and relative scarcity of ammonites is concerned, the Cretaceous deposits of Maintirano and Vato- mandry in Madagascar show greatest affinity, but this should probably be ascribed to the depositional environment rather than to more favourable migration routes. On the west coast of southern Africa, comparable deposits occur at Carimba in Angola which provide a definite date for Baculites subanceps, i.e. Late Campanian. UPPER CRETACEOUS SEDIMENTS FROM THE IGODA RIVER MOUTH 83 ACKNOWLEDGEMENTS We wish to express our gratitude towards the following persons for identi- fying and commenting on some of the material: Mr I. McMillan (SOEKOR, Johannesburg), Mr D. Salmon (Geological Survey, Cape Town) (Foraminifera), Prof. Dr E. Voigt (Geologisch-Palaontologisches Institut, Hamburg—Bryozoa), and Dr E. Owen (British Museum (Natural History), London—Brachiopoda). Mr L. Matthews collected some of the specimens while a graduate student. REFERENCES Basse, E. 1931. Monographie Paléontologie De Crétacé de la Province de Maintirano, Mada- gascar. Tananarive. Gouv. Gén. Madagascar et Dépend., Service des Mines. CHAPMAN, F. 1916. Foraminifera and ostracoda from the Upper Cretaceous of Needs Camp, Buffalo River, Cape Province. Ann. S. Afr. Mus. 12: 107-118. CoTTREAU, J. 1922. Paléontologie de Madagascar. X. Fossiles crétacés de la cote orientale. Annls Paléont. 11: 111-192. Crick, G. C. 1923. Appendix A. On Upper Cretaceous Cephalopoda from Portuguese East Africa. In: Teale, E. O. The Geology of Portuguese East Africa between the Sambezi and Sabie Rivers. Trans. geol. Soc. S. Afr. 26: 130-140. HowartH, M. K. 1965. Cretaceous ammonites and nautiloids from Angola. Bull. Br. Mus. nat. Hist. (Geol.) 10: 337-412. KENNEDY, W. J. & KLINGER, H. C. 1975. Cretaceous faunas from Zululand and Natal, South Africa. Introduction, Stratigraphy. Bull. Br. Mus. nat. Hist. (Geol.) 25: 265-315. 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. Kina, L. C. 1972. Geomorphic significance of the Late Cretaceous limestones at Needs Camp, near East London. Trans. geol. Soc. S. Afr. 75: 1-3. K.uincer, H. C. 1977. Cretaceous deposits near Bogenfels, South West Africa. Ann. S. Afr. Mus. 73: 81-92. K.uincer, H. C. & KENNEDY, W. J. 1977. Upper Cretaceous ammonites from a borehole near Richards Bay, South Africa. Ann. S. Afr. Mus. 72: 69-107. LANG, W. D. 1908. Polyzoa and anthozoa from the Upper Cretaceous limestones of Needs Camp, Buffalo River. Ann. S. Afr. Mus. 7: 1-11. Lock, B. E. 1973. Tertiary limestones at Needs Camp, near East London. Trans. geol. Soc. S. Afr. 76: 1-5. McGowran, B. & Moore, A. C. 1971. A reptilian tooth and Upper Cretaceous microfossils from the Lower Quarry at Needs Camp. Trans. geol. Soc. S. Afr. 74: 103-105. Mountain, E. 1974. The geology of the area around East London, Cape Province. An explanation of sheet map 3227 D (East London), 3228 (Kei Mouth). Pretoria: Geological Survey. Woops, H. 1908. Echinoidea, Brachiopoda and Lamellibranchiata from the Upper Cretaceous limestone of Needs Camp, Buffalo River. Ann. S. Afr. Mus. 7: 13-19. s¢ - 5 siragst der 6 aie 6. SYSTEMATIC papers must conform to the J/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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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. 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. 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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 a of publication given in text, e.g.: “Smith (1969) describes . “Smith (1969: 36, fig. 16) A oe og “As described (Smith 1969a, 19695; "Jones arian ‘As described (Haughton & Broom “oie “As described (Haughton et a/. 1927)...’ Note: no comma separating name nee 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, 5, 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) BuLLouGn, 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. 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. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51. TurEte, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: 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 77 Band February 1979 Februarie Part. -@.- » Deel CRETACEOUS FAUNAS FROM ZULULAND AND NATAL, SOUTH AFRICA THE AMMONITE SUPERFAMILY HAPLOCERATACEAE ZITTEL, 1884 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.), 5(1-3, 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 Copyright enquiries to the South African Museum Kopieregnavrae na die Suid-Afrikaanse Museum ISBN 0 908407 63 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 CRETACEOUS FAUNAS FROM ZULULAND AND NATAL, SOUTH AFRICA THE AMMONITE SUPERFAMILY HAPLOCERATACEAE ZITTEL, 1884 By WILLIAM JAMES KENNEDY Geological Collections, University Museum, Oxford & HERBERT CHRISTIAN KLINGER South African Museum, Cape Town (With 20 figures) [MS. accepted 18 October 1978] ABSTRACT Representatives of the superfamily Haplocerataceae Zittel, 1884, are locally common in rocks of Barremian age in northern Zululand, where Sanmartinoceras (Sanmartinoceras) africanum sp. nov. occurs at all stages of development from larval shells to discs which are still septate at a diameter of 120 mm, and represent the largest known specimens of the genus. Sanmartinoceras (Theganeceras) nodosum sp. nov., of Lower Aptian age, Sanmartinoceras (Sinzovia) trautscholdi (Sinzow), and an Aconeceras sp. of Upper Aptian age are other repre- sentatives of the Aconeceratidae, whilst the Binneyitidae are represented by specimens of Borissiakoceras of Middle Cenomanian age. In addition to systematic descriptions, the problems of recognizing dimorphism in these genera are discussed, and certain resultant taxonomic problems noted, and an annotated list of species referred to the two families is included. CONTENTS PAGE Introduction . ‘ ‘ ‘ : SD Location of specimens : 3 : . 86 Field localities . ; é : : - 86 Dimensions of specimens . : : «286 Suture terminology . ‘ ; 5 ~ 3S6 Systematic palaeontology : 87 Annotated list of species referred to the Aconeceratidae and Binneyitidae . a bY Acknowledgements. : ; : ~ 1a9 References! \.., 228 aie cri tee » 120 INTRODUCTION The Haplocerataceae are the longest ranging of the Ammonitina, first appearing in the Middle Jurassic (Bajocian) and ranging to the Upper Cretaceous (Coniacian). As Casey has noted, two broad morphologies recur within the superfamily: firstly, smooth platycones, typified by the Haploceratidae and Binneyitidae, and secondly, variously ribbed or strigate oxycones with entire or 85 Ann. S. Afr. Mus. 77 (6), 1979: 85-121, 20 figs. 86 ANNALS OF THE SOUTH AFRICAN MUSEUM crenulate keels, typified by the Oppeliinae, Strigoceratidae and Aconeceratidae (Casey 1961a: 118). In the Cretaceous of Zululand both groups are represented, in the Upper Barremian and Aptian the Aconeceratidae are not uncommon at some levels, whilst the Binneyitidae occur as a great rarity in rocks of Middle Cenomanian age. Although only a few species are represented in the authors’ collections, they are of some significance, for their specimens of Aconeceratidae allow, for the first time, the description of the detailed ontogeny of the widely occurring genus Sanmartinoceras Bonarelli, 1921, as well as clarifying some aspects of its stratigraphic distribution, whilst the present records of the binneyitid Borissiakoceras extend the known distribution of the group. The following species are described below: Aconeceras sp., compared to A. walshense (Etheridge) ; Sanmartinoceras (Sanmartinoceras) africanum sp. Nov. ; Sanmartinoceras (Theganeceras) nodosum sp. nov.; Sanmartinoceras (Sinzovia) trautscholdi (Sinzow); and Borissiakoceras sp. To this is added an annotated list of genera, subgenera and species referred to the two families. LOCATION OF SPECIMENS The following abbreviations are used to indicate the repositories of the material studied: BMNH British Museum (Natural History), London GSM Institute of Geological Sciences, London MNHP Muséum d’Histoire Naturelle, Paris OUM = Oxford University Museum, Oxford SAM _ South African Museum, Cape Town SAS South African Geological Survey, Pretoria FIELD LOCALITIES Details of localities mentioned in the text are given by Kennedy & Klinger (1975); fuller descriptions of sections are deposited in the Palaeontology Department of the British Museum (Natural History), London; Geological Survey, Pretoria; and South African Museum, Cape Town. DIMENSIONS OF SPECIMENS All dimensions given below are in millimetres: D = diameter, Wb = whorl breadth, Wh = whorl height, U = umbilical diameter. Figures in parentheses are dimensions as a percentage of the total diameter. SUTURE TERMINOLOGY The suture terminology of Wedekind (1916), recently reviewed and dis- cussed by Kullman & Wiedmann (1970) is followed here: Is = Internal lobe with septal lobe, U = Umbilical lobe, L = Lateral lobe, E = External lobe. | , CRETACEOUS FAUNAS FROM SOUTH AFRICA 87 SYSTEMATIC PALAEONTOLOGY Phylum MOLLUSCA Class CEPHALOPODA Cuvier, 1797 Subclass AMMONOIDEA Zittel, 1884 Order AMMONITIDA Zittel, 1884 Suborder AMMONITINA Zittel, 1889 Superfamily HAPLOCERATACEAE Zittel, 1884 Family Aconeceratidae Spath, 1923 Discussion The Aconeceratidae Spath, 1923, is a group of small to medium-sized, compressed, involute, commonly oxycone ammonites with flat or convex sides and a fastigiate to keeled venter. Ornament consists of weak to strong flexuous, commonly biconvex ribs and growth striae and a keel which may be minutely crenulate. The following genera and subgenera have been placed in the family: Genus Protaconeceras Casey, 1954, Hauterivian Genus Aconeceras Hyatt, 1903, Barremian to Lower Albian Genus Sanmartinoceras Bonarelli, 1921 (in Bonarelli & Nagera 1921), Barremian to Lower Albian Subgenus Sanmartinoceras s.s., Barremian to Aptian Subgenus Sinzovia Sazonova, 1958, Aptian Subgenus Theganeceras Whitehouse, 1926, Lower Aptian Genus Doridiscus Casey, 1961, Aptian Genus Nothodiscus Casey in Collignon, 1962, Aptian Genus Gyaloceras Whitehouse, 1927, Upper Aptian. Aconeceratids have a wide geographic distribution extending from Greenland to Antarctica, although their occurrence is sporadic. In Europe at least, they occur in enormous numbers in some clay facies as in southern France, where the blue marls of the Fosse Vocontienne yield thousands of Aconeceras and Casey (19616: 122) regarded them as an open sea group which invaded the neritic zone only in periods favourable for their growth. As will be seen below, however, their local abundance in carbonaceous nearshore sand- stones of the South African Barremian is scarcely compatible with such a generalization. The Haplocerataceae as a whole are characterized by striking dimorphism (Makowski 1962, Callomon 1963, Kennedy & Cobban 1976 with references), but within the Aconeceratidae our knowledge is such that it remains most difficult to recognize dimorphic pairs. In Aconeceras itself, the many hundreds of specimens from the French Aptian examined are all pyritic phragmocones and it would appear that juveniles, at least, are most difficult to split into dimorphs, as Palframan (1969) found in some Jurassic haploceratids. The authors do know, however, of a few specimens referrable to the family in which 88 ANNALS OF THE SOUTH AFRICAN MUSEUM apertures are preserved: the original material of S. (Sanmartinoceras) groenlandicum Rosenkrantz, 1934 (in Bogvad & Rosenkrantz), illustrated here as Figure 1A-C, is an obvious microconch with prominent lappets and a rostrum. Equally, Gyaloceras smithi Whitehouse, 1927, appears to be a female or macroconch. The specimen is reproduced here as Figure 2A. In most cases, however, the shell ornament of aconeceratids (as opposed to apertural form) gives little clue to dimorphism and a markedly biconvex growth line, indicating a long rostrum and blunt lappets, occurs, in the present material at least, in both small and large (if not mature specimens). The authors would, however, suggest that the presence of a strong spiral depression may indicate the possible presence of a long lappet (Figs 1A-B, 5C). The topic is returned to below under the discussion of Sanmartinoceras. Genus Aconeceras Hyatt, 1903 Type species Ammonites nisus d’Orbigny, 1841. Discussion Aconeceras is represented by a single, poor specimen from the Aptian of Zululand ; in consequence the reader is referred to Casey’s (19615: 123) extensive remarks on the genus. Occurrence The genus ranges from Barremian to Lower Albian and is widely distributed from western Europe and the U.S.S.R. to east Africa, Madagascar, Nepal, Antarctica and eastern Australia. Aconeceras sp. Material One specimen only, BMNH C80002, from the Makatini Formation, Aptian III, locality 166, Mfongosi Spruit, northern Zululand. Description The specimen is an external mould of an oxycone individual with an original maximum whorl height of over 20 mm. Coiling is very involute with a small, pit-like steep-sided umbilicus. The whorl section is high, compressed with the greatest breadth well below mid-flank. The flanks are distinctly flattened and the venter fastigiate. The specimen is somewhat worn and there is no obvious trace of ornament preserved. Discussion Although poorly preserved, this specimen is clearly referrable to the genus Aconeceras. Amongst described species it is perhaps to be compared with the 89 CRETACEOUS FAUNAS FROM SOUTH AFRICA ‘TX = “(pE6I) ZULIYUOSOY wz peaseg jo AdoD “pEeG] ‘zZJUBIYUOSOY WndIpunjuaosd (SpsaIOUIJADULUDG) SDAaIOUIJADULUDS *| “BL{ 90 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 2. A. Gyaloceras smithi Whitehouse, 1926. B-D. Aceneceras walshense (Etheridge), 1892. After Whitehouse (1926, 1927a). x1. CRETACEOUS FAUNAS FROM SOUTH AFRICA 91 Fig. 3. A-C. Sanmartinoceras (Sanmartinoceras) fontinale (Hudleston), 1890. D-G. Sanmartinoceras (Sanmartinoceras) olenae (Tenison-Woods), 1883. After Whitehouse (1927a). x1. 92 ANNALS OF THE SOUTH AFRICAN MUSEUM F Fig. 4. Sanmartinoceras (Sanmartinoceras) africanum sp. nov., paratypes. A. SAS H54/41/e. B-D. BMNH C79988. E-H. BMNH C79982. x1. CRETACEOUS FAUNAS FROM SOUTH AFRICA 93 Australian Aconeceras walshense (Etheridge) (see Whitehouse 1927a: 114, pl. 16 (figs 2-3); text-figs 1, 6-7) (Fig. 2B—D herein). Occurrence Upper Aptian of Zululand. Genus Sanmartinoceras Bonarelli, 1921 (in Bonarelli & Nagera 1921) Type species Sanmartinoceras patagonicum Bonarelli, 1921 (in Bonarelli & Nagera 1921). Diagnosis Stout oxycones, initially smooth, with falciform striae and ribs, the latter weak on the inner flank, sometimes bunched, and strengthening into distinctive concave ribs on the outer flank. Keel finely denticulate. Discussion Sanmartinoceras was originally based upon crushed specimens from the Aptian of Lago San Martin, Argentina. Leanza (1970, text-fig. 14) (see Fig. 5SA-B herein) has figured uncrushed topotypes, whilst Howarth (1958) and Thomson (1974) have figured other material from the sub-Antarctic Islands. Specimens are illustrated here as Figure SC_D. Specimens of S. patagonicum with apertures preserved are unknown, but Rosenkrantz (in Bogvad & Rosenkrantz 1934: 20, pl. 4 (fig. 3), pl. 5 (figs 1-5)) figured a species, S. groenlandicum Rosenkrantz, with a distinctive rostrum and lappets. These features, indicative of a micro- conch, have been incorporated into the generic diagnosis. Examination of these examples shows that the lateral lappets are associated with a distinct spiral groove (Fig. 1A-B). In the Antarctic specimens (Fig. SC—D) similarly sized individuals show this groove developed to varying degrees at the same diameter, and the authors suspect it may prove a criterion for differentiation of immature males and females at similar diameters, in the type species at least. In the case of Australian Sanmartinoceras the problem is even more tantalizing. Whitehouse (1926; 1927a) described four aconeceratids from the Aptian of Walsh River. Two, Sanmartinoceras fontinale (Hudleston) (Fig. 3D-G, 7A—H) and S. olenae (Tenison-Woods) (Fig. 3A—C) have strong falcoid ribs and prominent rostra. The others are very feebly ornamented. Aconeceras walshense (Etheridge) (Fig. 2B—D) has falcoid growth striae, whilst Gyaloceras smithi Whitehouse (Fig. 2A) has an inflated whorl and fastigiate venter. 1he authors strongly Suspect that these species are dimorphs but again cannot resolve the problem fully. Within Sanmartinoceras, three subgenera have been recognized. In addition to Sanmartinoceras sensu stricto, Theganeceras Whitehouse, 1926, was treated as a subgenus by both Wright (1957) and Casey (19616). Only three species, S. (T.) falcatum (von Koenen) (Fig. 6I-J) S. (7.) scalatum (von Koenen) (Fig. 6H) from the Lower Aptian of northern Germany, and S. (7.) grande 94 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 5. Sanmartinoceras (Sanmartinoceras) paragonicum Bonarelli, 1921. A-B. Topotype, after Leanza (1970). C. BMNH C49055. D. BMNH C49057. C_D. from Alexander Land. x1. CRETACEOUS FAUNAS FROM SOUTH AFRICA 95 H 1 J Fig. 6. A. Sanmartinoceras (Sinzovia) aff. trautscholdi (Sinzow), 1870. B—D. Sanmartinoceras (Sinzovia) stolleyi Casey, 1961. BMNH C14365 and 14362, from the Upper Aptian of Bekum, near Hildesheim, Germany. E-—F. Sanmartinoceras (Sinzovia) trautscholdi (Sinzow), 1870. Copy of Trautschold (1865). G. Sanmartinoceras (Theganeceras) grande Thomson, 1974. After Thomson (1974). H. Sanmartinoceras (Theganeceras) scalatum von Koenen, 1902. I-J. Sanmartinoceras (Theganeceras) falcatum von Koenen, 1902. H-J after von Koenen (1902). All x1. 96 ANNALS OF THE SOUTH AFRICAN MUSEUM Thomson (Fig. 6G) from the Aptian of Antarctica, have thus far been ascribed to the subgenus, which Casey (19616: 132) diagnosed as lacking the smooth juvenile stage of Sanmartinoceras sensu stricto and having finer, more numerous dense ribs. As is described below, the single Zululand specimen referred to the subgenus suggests that dimorphism may also present taxonomic problems. The subgenus Sinzovia Sazonova, 1958, differs from Sanmartinoceras sensu stricto (according to Casey 19616: 133) in having a very low keel and absence or poor development of a spiral groove. It differs from Theganeceras in having a longer juvenile smooth stage and greater tendency to smoothness on the lower flank. The type species is illustrated here as Figure 6E-—F, related German species as Figure 6A—D, and the only South African representative, S. (S.) trautscholdi (Sinzow), as Figure 7I-K. Occurrence Sanmartinoceras and its subgenera range from Upper Barremian to Aptian and are known from Greenland, western Europe, the U.S.S.R., Zululand, Madagascar, Nepal, Australia, Antarctica, Argentina and Papua. Subgenus Sanmartinoceras sensu stricto Sanmartinoceras (Sanmartinoceras) africanum sp. nov. Figs 4A-H; 8A-F; 9A-I; 10A-F; 11A-B; 12A-C; 13A-F; 14A-—C; 15F-J; 16-17; 19D-E Sanmartinoceras Kennedy & Klinger 197: 274. Types The holotype is SAS H54; paratypes are BMNH C79977-C80001, SAS H54/4la-d, SAS ZO(i), SAS LJE13la, SAS LJE112, SAS H54/17, SAS H54/33, all from the Makatini Formation, Barremian I-II, locality 170, Mlambongwenya Spruit, northern Zululand. Diagnosis A large (up to 120 mm diameter phragmocone) species of Sanmartinoceras in which the inner ‘haft’ of the ribs is narrow, straight and prorsiradiate and the outer ‘blade’ broad and markedly concave. Dimensions D Wb Wh Wb: Wh U Holotype, SAS H54 —(—) 20,5(—) 42-5(R—) 0,48 —(—) C79985 we ne 27,3(100) 6,6(24) 14,8(54) 0,45 4,1(15) C79978 ae .. 120,0(100) 28,8(24) 69,8(58) 0,41 TA C79984 .. ~—..-—«:115,0(100) 24,0(21) 65,5(57) :0.37-~——«11,3(9, 8) CRETACEOUS FAUNAS FROM SOUTH AFRICA 97 K Fig. 7. Sanmartinoceras (Sanmartinoceras) fontinale (Hudleston), 1890. Holotype, BMNH C5306, Upper Aptian, Primrose Springs, north of Lake Eyre, south Australia. A-D x1; E-H x2. I-K. Sanmartinoceras (Sinzovia) trautscholdi (Sinzow, 1870). SAM-—PCZ5919. x1. ANNALS OF THE SOUTH AFRICAN MUSEUM 98 Fig. 8. Sanmartinoceras (Sanmartinoceras) africanum sp. nov. A-C. Holotype SAS 54; D-F paratype BMNH C79989. x1. CRETACEOUS FAUNAS FROM SOUTH AFRICA Fig. 9. Sanmartinoceras (Sanmartinoceras) africanum sp. nov., paratypes. A-D. SAS H54/416. E-H. SAS H54/41a. I. BMNH C79996. All x2. 99 100 ANNALS OF THE SOUTH AFRICAN MUSEUM C D E F Fig. 10. Sanmartinoceras (Sanmartinoceras) africanum sp. nov., paratypes. A-B. BMNH C€79984. C-F. SAS H54/41c. A-B x1; C-F x2. CRETACEOUS FAUNAS FROM SOUTH AFRICA 101 Fig. 11. Sanmartinoceras (Sanmartinoceras) africanum sp. noy., paratypes. A. BMNH C79984. B. SAS H54/41d. A x1; B x2. 102 ANNALS OF THE SOUTH AFRICAN MUSEUM Description The barrel-shaped protoconch is succeeded briefly by rather stout whorls, beyond which, up to a diameter of 0,5-0,8 mm, the shell is moderately involute, slightly compressed, with flattened flanks and a rounded venter. All the available specimens are partially or wholly exfoliated, and any ornament, if present, must have been weak. A few specimens begin to show traces of a ventral ridge, but not a keel at this diameter, although these features may be an artefact of preservation. From 0,8 mm onwards the shell is an oxycone, the coiling is very involute, with a tiny, crater-like umbilicus. The umbilical wall is flat and subvertical, with an abruptly rounded shoulder. The whorl section varies markedly; stout individuals have a whorl breadth to height ratio of 0,6; in slender individuals the figure is 0,4. Maximum whorl breadth is low on the flanks; the inner flanks are flattened, the outer flanks convergent; a distinct ventrolateral shoulder is developed, and there is a sharp keel which, when well preserved, can be seen to be minutely crenulated (Fig. 9E—-H). Ornament varies greatly. In some individuals (Figs 11B, 13C—D) there are only the finest growth striae. These arise at the umbilical seam, but are very weak. They pass forwards across the inner flank, strengthening as they do so, and sometimes splitting (Fig. 15G—J). They are at their maximum development at mid-flank, where they flex gently backwards, giving rise to a clear concavity which extends across the outer third of the flank (Fig. 13D). Striae decline on the ventrolateral shoulder and project strongly forwards to meet the keel at an acute angle. Individual striae correspond to individual crenulations on the keel. In robust specimens, e.g. BMNH C79983, striae are accompanied by distant, pronounced ribs, which are strongest at mid-flank; there is every intermediate between. Middle growth stages are also characterized by a wide range of variation (Figs 4E-H, 8A-F). The holotype demonstrates the features of a strongly ornamented individual. It is wholly septate (Fig. 8B) retaining traces of shell. Coiling is very involute, with most of the inner whorls being covered. The tiny umbilicus is deep, with a high, flat, subvertical wall, abruptly rounded shoulder, and faint umbilical carina. The whorl section is compressed (breadth to height ratio is 0,48) with faintly concave inner flanks, a weakly inflated mid-flank region converging to distinct shoulders, clearly demarcated from a high, septate ventral keel. In none of the medium-sized specimens does the keel bear serrations. Ornament consists of fine, rectiradiate striae on the inner part of the flank, effaced at a spiral mid-flank depression. The outer flank bears low, blunt, concave ribs, declining at the ventrolateral shoulder, but projected forwards to meet the keel. The largest available specimens are up to 120 mm in diameter, and are still wholly septate (Fig. 12A—C). In these, the whorls are quite robust, and ornament consists of striae on the inner flank with blunt, concave ribs on the outer flank. None of the present specimens has the aperture preserved. A few fragments show 103 CRETACEOUS FAUNAS FROM SOUTH AFRICA ‘TX °8L66LO HNWa edéjered “aou “ds wnuvatafo (Spsav0uljADUWUDS) SDsaIOUllADULUDS *Z} ‘SIq 4 mt dual ~ » —~ ? Se . —.- S & ” 7 104 ANNALS OF THE SOUTH AFRICAN MUSEUM F Fig. 13. Sanmartinoceras (Sanmartinoceras) africanum sp. NOV., paratypes. A-B, F. SAS Zo(i). C. SAS LJE 131a. D. SAS H54/41a. E. BMNH C79979. A-D, E-F x1; C-D x2. —— i re 105 CRETACEOUS FAUNAS FROM SOUTH AFRICA ‘TX A-d ‘1X O-V ‘9] ‘ou UOHI[OD J9d00D “YW “ds sosav0yvissiaog “4-C “Z11 ALT SVS edAjered “Aou ‘ds wnuvd1fo (spsavouljavUUDs) Sp4aI0UlJADULUDS *D-Y*pl “BLT V 106 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 15. A-E. Borissiakoceras sp., BMNH C80003. F—-J. Sanmartinoceras (Sanmartinoceras) africanum sp. nov., paratypes. F is SAS LJE 112; G-J is BMNH C79985. A-E, G-J x2; F x1. CRETACEOUS FAUNAS FROM SOUTH AFRICA 107 weak to strong strigations (Figs 4B—D, 13E). Where well preserved, this appears to be a feature of the dorsal shell layer, and not of the original external shell surface. That this occurs in some very large specimens suggests the species may have reached diameters close to 200 mm. The available specimens show the details of suture development to advantage, as shown in Figure 16. Mature sutures are deeply and intricately subdivided, as shown in Figure 17. Discussion The specimens show the ontogeny and range of intraspecific variation far better than in any previously described Sanmartinoceras whilst the specimens are the largest known for the genus. It is most unfortunate that none of the specimens retains apertures, but the rather distinct spiral flank depression present in the holotype suggests that it may be a microconch; the largest individuals (Figs 1OA—B, 11A, 12A—D) are probably macroconchs. Thomson (1974: 23) reviewed criteria used to differentiate species of Sanmartinoceras as follows: (i) strength and form of ribbing, (ii) the way in which ribs appear (i.e. abruptly or increasing gradually in strength), (iii) the size of the individual at which ribs first appear. In his discussion, however, he concluded that the form of the falcate ribbing alone was sufficient to separate species. Figure 18 compares the line of ribs and striae in the five described species; on this criterion alone it can be seen that the straight ‘haft’ and form of ‘sickle’ of the present species are quite different from the markedly biconcave rib of S. olenae, S. fontinale and S. patagonicum. A straight ‘haft’ characterizes S. groenlandicum (Fig. 18), but here the ‘blade’ is much more deeply concave with the inner half almost straight. As noted elsewhere, the authors do not regard the presence or absence of a lateral spiral groove as of specific significance, whilst the present specimens vary greatly in strength of ornament and growth stage at which it appears. The other feature which separates S. africanum from other species is its great size, but this may be no more than an artefact of preservation. Occurrence Upper Barremian of Zululand only. Subgenus Theganeceras Whitehouse, 1927 Theganoceras nodosum sp. nov. Figs 19A-C, 20 Holotype SAM-PCZ5708 from the Makatini Formation, Aptian I, locality 170, Mlambongwenya Spruit, northern Zululand. 108 ANNALS OF THE SOUTH AFRICAN MUSEUM A jf / / B 4 Vg / / Py Cc i itn “e$ Fig. 16. Suture development in Sanmartinoceras (Sanmartinoceras) africanum sp. nov. A-C. BMNH C79979. D-E. SAS H54/41A. All x 12,5. 109 CRETACEOUS FAUNAS FROM SOUTH AFRICA ‘STIX “Olb/PSH SWS ‘Aou ‘ds wnupotsfo (Spdad0UujJ ADU) SDABDOUILADUUDS JO SaINyNs oINIeY| ‘LI ‘31g 110 ANNALS OF THE SOUTH AFRICAN MUSEUM a ———— i A B Vite Fig. 18. Rib patterns in Sanmartinoceras (Sanmartinoceras) compared. A. S. (S.) olenae. B-C. S. (S.) fontinale. D. S. (S.) africanum. E. S. (S.) groenlandicum. F-G. S. (S.) patagonicum. Diagnosis A large species of Theganeceras with broad, flat, falcoid ribs on the inner whorls which bear umbilical bullae and incipient ventral clavi. Outer whorls ornamented by bullae, flexuous growth lines, folds and constrictions. Description The unique holotype of this species comprises a crushed body chamber and a nucleus. Growth of calcite between shell and sedimentary infilling has produced an unduly thick ‘shell’ of partly diagenetic origin. The inner whorl, at a diameter of approximately 38 mm, bears Santos to sixteen closely spaced, low, broad, sickle-shaped ribs which are straight and prorsiradiate on the inner flank, flexing backwards at mid-flank and becoming markedly concave on the outer flank. The ribs are strong and well developed down to the umbilicus where a distinct bulla is present, whilst on the ventro- lateral shoulder they strengthen into an incipient clavus. The ribs themselves and interspaces between bear dense striae, sometimes strengthened into riblets, giving some ribs a bunched, fasciculate appearance. The venter is fastigiate, and bears strongly projected striae. CRETACEOUS FAUNAS FROM SOUTH AFRICA 111 B Cc Fig. 19. A-C. Sanmartinoceras (Theganeceras) nodosum sp. nov., holotype, SAM-—PCZ5708. x 1. —E. S. (Sanmartinoceras) africanum sp. nov., paratype, BMNH specimen. x 1. 112 ANNALS OF THE SOUTH AFRICAN MUSEUM The outer body whorl shows coiling to have been involute with a moderately large, deep, crater-like umbilicus with a flat, outwards-sloping wall. The whorl section is compressed, lanceolate, so that the overall shell form was an oxycone. The whorl breadth to height ratio is estimated as approximately 0,38, the greatest breadth being close to the angular umbilical shoulder. The preserved fragment bears seven small, sharp, comma-shaped bullae at the shoulder. These give rise to bundles of fine, dense, flexuous striae which, being crowded close to the umbilicus, appear as ribs, which fade out at a short distance from the bulla. The striae flex forwards to mid-flank, then backwards, to form the very shallow blade of a sickle. They are strongly projected across the ventro- lateral shoulder. Paralleling these striae are irregular, low folds and constrictions. The imperfectly exposed suture is illustrated in Figure 20. Discussion The presence of well-developed ribs which extend to the umbilical shoulder indicates this specimen to be a Theganeceras. It differs from all previously described species in having broader, less markedly flexed ribs when young, as well as possessing umbilical bullae, which feature is unique for the subgenus. Only S. (Theganeceras) grande Thomson (1974, pl. 4 (fig. 9)) approaches this specimen in size; the unique holotype of that species is quite distinct, however, lacking bullae and having strongly flexed ribs and striae (Fig. 6G). Equally, no S. (Sinzovia) bears bullae, and the closest species, S. (S.) stolleyi Casey (Fig. 6A—D), has ribs which efface on the inner flank. The presence of bullae also distinguishes our specimen from described S. (Sanmartinoceras) at comparable dimensions. Occurrence Lower Aptian I of northern Zululand only. Subgenus Sinzovia Sazonova, 1958 Sanmartinoceras (Sinzovia) trautscholdi (Sinzow) Figs 4I-K, 6E-F Ammonites bicurvatus Trautschold (non Michelin), 1865: 22, pl. 3 (figs 17a-c). Oppellia Trautscholdi Sinzow, 1870: 118, pl. A (figs 1, la—b only). - Sinzovia trautscholdi (Sinzow): Sazonova, 1958: 128, pl. 6 (fig. 2), pl. 8 (figs 1, 1a, 3-7), pl. 10 (figs 4-5) (with synonymy). Druschchitz & Kudryavtseva, 1960: pl. 42 (figs 7a—b). Collignon 1962: 31, pl. 229 (fig. 974). ? Sanmartinoceras (Sinzovia) sp. cf. trautscholdi (Sinzow): Casey 1961b: 136, pl. 26 (fig. 7). Holotype Trautschold’s original of his plate 3 (fig. 17a-c), from the Lower Aptian of Simbirsk (now Polivna) in the Stalingrad area of Russia, on which Sinzow (1870: 118) based his species. The original figures are reproduced here as Figure 6E-F. 113 CRETACEOUS FAUNAS FROM SOUTH AFRICA ‘STLX “80LSZOd-WYS ‘odAjojoy 94} Jo oinjns yenszed “Aou ‘ds wnsopou (spsazaudsay [) SDAQIOUIJADUUDS! “QT “BLA or gts 114 ANNALS OF THE SOUTH AFRICAN MUSEUM Material SAM-PCZ5919 from the Makatini Formation, Aptian IV at locality 152, Mkuze Game Reserve, northern Zululand. Description The specimen is a beautifully preserved disc of which the last half of a whorl is body chamber. It retains extensive areas of replaced shell, in consequence of which the sutures are not fully decipherable. The dimensions are as follows: D Wb Wh Wb:Wh U 43,0 10,0(23,3) 22,0(51,2) 0,45 —-9,0(20,9) Coiling is very involute, oxycone, with two-thirds of the previous whorl being covered. The umbilicus is small and shallow (20° of diameter), with a flat, subvertical wall. The umbilical shoulder is sharp and angular. The whorl section is very compressed, with a breadth to height ratio of 0,45. The inner flank is flattened to concave, with gently rounded, convergent outer flanks, converging to an acute, delicately keeled venter. The inner, concave part of the flank is virtually smooth where shell is present, due to weathering. On the mould, however, there are delicate, concave, prorsiradiate, bunched striae which sweep forwards to a distinct spiral ridge at mid-flank and a parallel groove external to this. On the ridge and groove, the striae flex backwards in a marked convex projection. On the outer flank, they flex backwards to form a ‘blade’ corresponding to the bunches of ‘handles’ to what is an overall sickle-like ornament. The blades are strong, concave, blunt ribs, strong over most of the last whorl, although effacing on the venter, which is smooth on either side of a narrow, sharp ventral keel. The sutures cannot be seen. Discussion On the basis of weakness of inner flank ornament, and strength and style of ribs on the outer flank, the authors would refer their single specimen to Sinzow’s species. It is particularly close to the specimen from Ulyanovsk, Russia, illustrated by Casey (19616, text-fig. 143 (1a-b)). Sinzovia stolleyi Casey (19616, text-fig. 143(g)) is more distantly and narrowly ribbed, as can be seen from Figure 6A—D herein. Sinzovia aptianum (Sarasin) (1893: 155, pls 4-6 (fig. 12a—c); Casey 19615: 134, pl. 26 (fig. 6a—b)) has blunter, fewer ribs on the outer flank. Sanmartinoceras ? (Sinzovia?) sp. nov. of Casey (19616: 136, text-fig. 43(f)) is a Lower Albian species of uncertain affinity. Occurrence Lower Aptian of the U.S.S.R. and southern England; Upper Aptian of Madagascar and Zululand. CRETACEOUS FAUNAS FROM SOUTH AFRICA 115 Family Binneyitidae Reeside, 1927 The Binneyitidae are a family of micromorphs, characteristically platycone, with much reduced sutures. Two genera, Borissiakoceras Arkhangelsky, 1916, and Binneyites Reeside, 1927, were referred to the family by Wright (1957) who suggested derivation from the Acanthocerataceae, but subsequent work has revealed the presence of passage forms linking the family to the Aconeceratidae via Falciferella Casey, 1954. This Middle to Upper Albian genus has a vestigal keel when young and can be linked to the platycone but still feebly carinate Aptian aconeceratids Doridiscus Casey, 1961, and Nothodiscus Casey, 1962 (in Collignon 1962). The evolutionary origins of the family thus involved a change from oxycone to platycone with corresponding loss of keel, and sutural simplification. Genus Borissiakoceras Arkhangelsky, 1916 Type species Borissiakoceras mirabile Arkhangelsky, 1916. Diagnosis Small, compressed, moderately involute to moderately evolute platycones. Flanks typically smooth, sometimes bearing narrow, falcoid ribs. A few species bear ventrolateral tubercles. Suture simple with narrow, bifid lobes and broader, bifid or trifid saddles. Discussion Borissiakoceras is a distinctive genus, differing obviously from the later (Coniacian) Binneyites Reeside, 1927, which has stronger lateral and ventro- lateral ornament, sharp ventrolateral shoulders and more auxiliary elements in the suture. Johnsonites Cobban, 1961, has a simpler suture and a flat or concave venter. Borissiakoceras is clearly descended from Falciferella, species of which extend to the Upper Albian. It is the only binneyitid in which dimorphism has been demonstrated (Kennedy & Cobban 1976: pl. 1 (figs 3-4)); males have stronger, more markedly biconcave growth striae and ribs, reflecting a similar aperture, with a short rostrum. Occurrence Borissiakoceras is best known from North America, ranging from Texas (Stephenson 1952, 1955) to Kansas and Colorado (Morrow 1935). Other United States occurrences are summarized by Cobban (1961). It also occurs in British Columbia (Warren & Stelck 1958) and Alaska (Cobban & Gryc 1961). In these regions it ranges from Middle Cenomanian to Lower Turonian. The type species comes from Turkestan and is probably of late Cenomanian age; 116 ANNALS OF THE SOUTH AFRICAN MUSEUM Bodylevsky & Shulinga (1958) record the genus from the Turonian—Coniacian of the northern U.S.S.R., and Kennedy & Juignet (1973) record it from the Middle Cenomanian of Normandy, whilst the present authors have seen a fragment from the Lower Cenomanian of Sarthe (Sorbonne collections). Wright (1963) records a doubtful species from the Middle Cenomanian of northern Australia, and the Engonoceratidae gen. et sp. nov. of Henderson (1973: 106, pl. 14 (fig. 8)) belongs here, as do the Middle Cenomanian specimens from Zululand described below. Borissiakoceras sp. Figs 14D-F, 15A-E Material Two specimens, BMNH C80003, and no. 16 in M. R. Cooper’s collection, University of Natal, Durban, both from the Middle Cenomanian (Cenomanian III) Locality 62, the Skoenberg, Zululand. Description The specimens comprise a small phragmocone and a body chamber with a maximum whorl height of 7,4 mm. Coiling is involute with a small, shallow umbilicus comprising approximately 20 per cent of the diameter. The overall form is platycone, the whorl breadth to height ratio being approximately 0,5, with a low umbilical wall, flattened inner, and slightly convergent outer flanks, abruptly and narrowly rounded ventrolateral shoulder, and somewhat flattened venter. Both specimens are corroded, so that no trace of any original ornament remains. The suture is poorly exposed, but much simplified, with such elements as are visible resembling those of B. mirabile. Discussion Overall shell form and suture indicate these specimens to be Borissiakoceras, the first representatives of the genus, and indeed the Binneyitidae, to be described from Africa. Because of poor preservation it is not possible to identify them fully. They most closely resemble feebly ornamented species such as B. mirabile (see Kennedy & Juignet 1973) and B. orbiculatum (see Cobban, 1961: 750, pl. 88 (figs 15-41), text-figs 5a-f). Occurrence Middle Cenomanian of Zululand. CRETACEOUS FAUNAS FROM SOUTH AFRICA 117 ANNOTATED LIST OF SPECIES REFERRED TO THE ACONECERATIDAE AND BINNEYITIDAE Family Aconeceratidae Spath, 1923 Genus Protaconeceras Casey, 1954 Type species: Oppelia patagoniensis Favre, 1908, by original designation. Protaconeceras patagoniense (Favre), 1908: 634, pl. 34 (fig. 7), pl. 37 (figs 3-5), text-fig. 6. Lower Hauterivian of Patagonia. Protaconeceras spp. nov. Casey, 1954: 270, pl. 7 (fig. 7), text-fig. 2. Upper Hauterivian, England. Genus Aconeceras Hyatt, 1903 (= Adolphia Stolley, 1907; Adolphites Hennig, 1932) Type species: Ammonites nisus d’Orbigny, 1841, by monotypy. Aconeceras nisus (d’Orbigny), 1841: 184, pl. 55 (figs 7-9). Casey 19616: 128. Upper Aptian of western Europe, notably Gargasian clays of the Vocontian Trough. Also recorded from Madagascar and elsewhere, although many records are dubious. Aconeceras haugi (Sarasin), 1893: 156, pls 4—6, text-fig. 1la-c. Lower Aptian of western Europe, Nepal (Bordet ef a/. 1971), and Madagascar (Collignon 1962) where it is said to be of Upper Aptian age. Aconeceras neonisoides Casey, 19616: 129, pl. 26 (figs 1, 9-10), text-fig. 41d—e. Lower Albian of southern England and north Africa (Sornay 1955; Dubourdieu 1956). Aconeceras australonisoides Brunnschweiler, 1959: 11, pl. 1 (fig. la—b). Aptian of western Australia. Aconeceras whitehousei Brunnschweiler, 1959: 12, pl. 1 (fig. 2a—b). Aptian of western Australia. Aconeceras walshense (Etheridge), 1892: 493, pl. 42 (figs 10-11). Whitehouse 1926: 203, pl. 34 (fig. 1), pl. 37 (fig. 3); 1927a@: 114, pl. 16 (figs 2-3), text-figs 1, 6-7. Aptian of Queensland. Aconeceras nisoides (Sarasin), 1893: 155, pls 4-6 (fig. 10a—c), text-figs 3, 5. Lower Aptian of western Europe. Aconeceras luppovie (Sazonova), 1958: 130, pl. 8 (fig. 2). Aptian of the U.S.S.R. Aconeceras saratoviensis (Sazonova), 1958: 130 (= Oppelia trautscholdi Sinzow, 1898, pl. A (figs 4-5 only)). Lower Aptian of the U.S.S.R. Genus Sanmartinoceras Bonarelli, 1921 Type species: Sanmartinoceras patagonicum Bonarelli, 1921, by monotypy. Sanmartinoceras (Sanmartinoceras) patagonicum Bonarelli, 1921, in Bonarelli & Nagera 1921: 27, pl. 5 (figs 3-6). See also Howarth 1958: 5, pl. 1 (figs 6-10); Leanza 1970: 215, fig. 14; Thomson 1974: 24, pl. 4b-f, text-fig. 7a. Aptian of Argentina and Antarctica. (See Fig. 6A—-D herein.) Sanmartinoceras (Sanmartinoceras) groenlandium Rosenkrantz, 1934, in Bogvad & Rosen- a 1934: 20, pl. 4 (fig. 3), pl. 5 (figs 1-5). Aptian of east Greenland. (See Fig. 1A—C erein.) Sanmartinoceras (Sanmartinoceras) olenae (Tenison-Woods), 1883: 150, pl. 7 (fig. 8), pl. 8 (fig. 1). Whitehouse 1926: 205, pl. 41 (fig. 3); 1927a: 117, pl. 17 (fig. 6), text-figs 3, 4, 9. Aptian of Australia. (See Fig. 3A—C herein.) Sanmartinoceras (Sanmartinoceras) fontinale (Hudleston), 1890: 241, pl. 9 (fig. 1). Whitehouse, 1927a: 116, pl. 17 (figs 2-5). Aptian of Australia. (See Figs 3D-G, 7A-H herein.) Sanmartinoceras (Sanmartinoceras) africanum Kennedy & Klinger, 1978 sp. nov. (See p. 96.) Upper Barremian of Zululand. Subgenus Sinzovia Sazonova, 1958 Type species: Ammonites trautscholdi Sinzow, 1870 (= Ammonites bicurvatus Trautschold, 1865 non Michelin, 1838) by original designation. Sanmartinoceras (Sinzovia) trautscholdi (Sinzow), 1870: 118-119. See also Ammonites bicurvatus Trautschold, 1865: 22, pl. 3 (fig. 17a—-c) (non Michelin); Casey, 19615: text-fig. 135a—c. Lower Aptian of the U.S.S.R. and southern England; Upper? Aptian of Madagascar; Upper Aptian of Zululand. 118 ANNALS OF THE SOUTH AFRICAN MUSEUM Samartinoceras (Sinzovia) aptianum (Sarasin), 1893: 155, pls 4-6, fig. 12a—c. Casey 19616: 134, pl. 26 (fig. 6a—b), text-fig. 43d—e. Aptian of France, southern England and elsewhere in Europe; Nepal (Bordet et a/. 1971). (See Fig. 6E—F herein.) Sanmartinoceras (Sinzovia) stolleyi Casey, 19616: 133, 136; text-fig. 135g—h. Upper Aptian of Germany. (See Fig. 6A—D herein.) Sanmartinoceras ? (Sinzovia ?) sp. nov. Casey, 19616: 136, text-fig. 43f. Aptian of England. vnon Sinzovia luppovie Sazonova, 1958: 130, pl. 8 (fig. 2) = S. luppovi Casey, 19616: 130. Lower Aptian of the U.S.S.R. An Aconeceras according to Casey (19616: 133). ?non Sinzovia saratoviensis Sazonova, 1958 (= Oppelia trautscholdi Sinzow, 1898, pl. A (figs 4-5a only). Lower Aptian of the U.S.S.R. Also an Aconeceras according to Casey (19616: 133). Subgenus Theganeceras Whitehouse, 1926 Type species: Oppelia scalata von Koenen, 1902, by original designation. Sanmartinoceras (Theganeceras) grande Thomson, 1974: 25, pl. 4g. Lower Aptian of Alexander Island. (See Fig. 6G herein.) Sanmartinoceras (Theganeceras) scalatum (von Koenen), 1902: 54, pl. 45 (fig. 6). Lower Aptian of north Germany. (See Fig. 6H herein.) Sanmartinoceras (Theganeceras) falcatum (yon Koenen), 1902: 48, pl. 45 (figs 7-8). See also Casey 19616: 132, pl. 26 (fig. 2). Lower Aptian of north Germany and England. (See fig. 6I-J.) Sanmartinoceras (Theganeceras) (?) sp. Thomson, 1974: 26, pl. 4k. Lower Aptian of Alexander Island. Sanmartinoceras (Theganeceras) nodosum Kennedy & Klinger sp. nov. (See p. 107.) Lower Aptian of Zululand. Genus Gyaloceras Whitehouse, 1927 Type species: Gyaloceras smithi Whitehouse, 1927 by original designation. Gyaloceras smithi Whitehouse, 1927a: 115, pl. 17 (fig. 1), text-fig. 8. Aptian of Australia. (See Fig. 2A.) Gyaloceras ibo Reyment, 1955: 15, pl. 2 (figs 1-3), text-figs 3-4. Upper Albian of Nigeria. Casey (19616: 139) has suggested that this species is ‘a completely new development whose relationship to the Aconeceratidae is doubtful. Possibly it is congeneric with the Aconeceras ? described by Haas (1942: 165) from the Upper Albian of Angola’. ? Genus Eofalciferella Brunnschweiler, 1959 Type species: Eofalciferella condoni Brunnschweiler, 1959, by original designation. Eofalciferella condoni Brunnschweiler, 1959: 13, pl. 1 (figs. 3-4). Aptian of Australia. The only illustrations of this species are pencil sketches of the unique holotype, a crushed specimen from the Windalia Radiolarite. The species and genus are best treated as nomen dubia; Casey (19615: 131) implies in his discussion of the genus that its affinities may be with Sanmartinoceras. Family Binneyitidae Reeside, 1927 Genus Falciferella Casey, 1954 Type species: Falciferella milbournei Casey, 1954, by original designation. Falciferella milbourni Casey, 1954: 274, pl. 7 (figs 1-5), text-fig. 3. Middle Albian of southern England. Falciferella malandiandrensis Collignon, 1962: 32, pl. 229 (fig. 975). Upper Aptian of Madagascar. This species is keeled to a diameter of approximately 45 mm. It may be an Aconeceras, although Collignon (1962) indicates that it has a Falciferella-like suture. Genus Doridiscus Casey, 1961 Type species: Doridiscus rotulus Casey, 1961, by original designation. Doridiscus rotulus Casey, 1961b: 139, pl. 26 (fig. 8a—b), text-fig. 44c-e. Low Upper Aptian of southern England. Doridiscus sp. nov. indet? Casey, 19615: 140, text-fig. 44f. Upper Lower Aptian of southern England. CRETACEOUS FAUNAS FROM SOUTH AFRICA 119 Genus Nothodiscus Casey in Collignon, 1962 Type species: Nothodiscus planus Casey, in Collignon, 1962, by original designation. Nothodiscus planus Casey in Collignon, 1962: 32, pl. 229 (fig. 976). Upper Aptian of Madagascar. Genus Borissiakoceras Arkhangelsky, 1916 Type species: Borissiakoceras mirabile Arkangelsky, 1916: 55, pl. 8 (figs 2-3), Lower Turonian of Turkestan. Kennedy & Juignet, 1973: 900, text-figs 1-2, Middle Cenomanian of France. Borissiakoceras compressum Cobban, 1961: 747, pl. 87 (figs 19-33); pl. 89 (figs 1-9), text- fig. 4a—k. Middle Cenomanian of the United States Western Interior. Borissiakoceras reesidei Morrow, 1935: 463, pl. 49 (fig. 7a—b), pl. 50 (fig. 5), text-fig. 8. Cobban, 1961: 749, pl. 88 (figs 1-14), text-fig. 3h-k. Middle Cenomanian of the United States Western Interior. Borissiakoceras orbiculatum Stephenson, 1955: 64, pl. 6 (figs 1-4). Cobban, 1961: 750, pl. 88 (figs 15-41), text-fig. Sa-f. Middle/Upper Cenomanian of Texas and the United States Western Interior. Borissiakoceras cf. B. orbiculatum Stephenson, 1955; Cobban 1961: 753, pl. 89 (figs 10-14), text-fig Sg, i. Uppermost Cenomanian of the Black Hills, United States Western Interior. Borissiakoceras inconstans Cobban & Gryc, 1961: 187, pl. 38 (figs 30-37), text-fig. 2i-l. Latest? Cenomanian of Alaska. Borissiakoceras ashurkoffae Cobban & Gryc, 1961: 188, pl. 38 (figs 38-43), text-fig. 2j-k, m. Lower Turonian of Alaska. Borissiakoceras (?) sp. Wright, 1963: 602, pl. 89 (fig. 5). Middle Cenomanian of Bathurst Island, Australia. Borissiakoceras ? sp. Lower Cenomanian of Sarthe, France (Sorbonne collections). Borissiakoceras sp. Kennedy & Klinger, 1978. (See p. 116.) Middle Cenomanian of Zululand. Genus Johnsonites Cobban, 1961 Type species: Johnsonites sulcatus Cobban, 1961, by original designation. Johnsonites sulcatus Cobban, 1961: 743, pl. 87 (figs 1-18), text-fig. 3a—g. Middle Cenomanian of Wyoming and Colorado in the United States Western Interior. ? Johnsonites sp., the original of Stephenson’s (1952: 198, pl. 45 (figs 5—6)) Euhoplites ? sp. from the Middle Cenomanian of Texas may also belong to this genus. Genus Binneyites Reeside, 1927 Type species: Binneyites parkensis Reeside, 1927, by original designation. Binneyites parkensis Reeside, 1927: 5, pl. 3 (figs 1-10). Cobban, 1961: 754, pl. 89 (figs 32-37), text-fig. Ss, t. Coniacian of Wyoming and Utah in the United States Western Interior. Binneyites carlilensis Cobban, 1961: 755, pl. 89 (figs 15-22), text-fig. Sh, j-m. Mid-Turonian of South Dakota and Wyoming in the United States Western Interior. Binneyites aplatus (Morrow), 1935: 465, pl. 49 (fig. 5), pl. 50 (fig. 6), text-fig. 7. Mid-Turonian of Kansas in the United States Western Interior. Binneyites rugosus Cobban, 1961: 756, pl. 89 (figs 26-31), text-fig. Sn—p. The Engonoceratidae gen. et sp. nov. of Henderson, 1973: 106, fig. 14 (no. 8), text-fig. 15, is either a Borissiakoceras or Binneyites of Turonian age. 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), Mr C. W. Wright and Dr M. R. Cooper (Oxford), for their advice and useful discussions. The assistance of the staff of the Geological Collections at Oxford University Museum, and South African Museum, Cape Town, is gratefully acknowledged as is the financial support of the Trustees of the Sir Henry Strakosh Bequest, the Royal Society, and Natural Environment Research Council. 120 ANNALS OF THE SOUTH AFRICAN MUSEUM REFERENCES ARKHANGELSKY, A. D. 1916. Les mollusques du Crétacé Supérieur du Turkestan. Trudy geol. Kom. n.s. 152: 1—57. BopyLevsky, V. I. & SHULINGA, N. I. 1958. Jurassic and Cretaceous faunas of the lower Yenisei River region, U.S.S.R. Trudy nauchno-issled. Inst. Geol. Arkt. 93: 1-196 (In Russian.) BoGvAD, R. & ROSENKRANTZ, A. 1934. Beitrage zur Kenntnis der unteren Kreide Ostgronlands. Medadr. Gronland 93: 15-28. BONARELLI, G. & NAGERA, J. J. 1921. Observaciones geologicas en las immendiaciones del Lago San Martin (Territorio de Santa Cruz). Boln Dir. gen. Minas Geol. Hidrol., B. Aires ser. B. (Geoldgia) 27: 1-39. BorbDetT, P., COLCHEN, L., KRUMMENACHER, D., LE Fort, P., MOUTERDE, R., & Remy, M. 1971. Recherches Géologiques dans Himalaya du Népal, Région de la Thakkhola. Paris: Centre National de la Recherche Scientifique. BRUNNSCHWEILER, R. O. 1959. New Aconeceratinae (Ammonoidea) from the Albian and Aptian of Australia. Bull. Bur. Miner. Resour. Geol. Geophys. Aust. 54: 1-19. CALLOMON, J. H. 1963. Sexual dimorphism in Jurassic ammonites. Trans. Leicester lit. phil. Soc. 57: 21-56. Casey, R. 1954. Falciferella, a new genus of Gault ammonites, with a review of the family Aconeceratidae in the British Cretaceous. Proc. Geol. Ass. 65: 262-277. Casey, R. 196la. A monograph of the Ammonoidea of the Lower Greensand. Palaeontogr. Soc. (Monogr.) part 2: 45-118. Casey, R. 19615. A monograph of the Ammonoidea of the Lower Greensand. Palaeontogr. Soc. (Monogr.) part 3: 119-216. CosBan, W. A. 1961. The ammonite family Binneyitidae Reeside in the Western Interior of the United States. J. Paleont. 35: 176-190. CosBaNn, W. A. & Gryc, G. 1961. Ammonites from the Seabee Formation (Cretaceous) of northern Alaska. J. Paleont. 35: 176-190. COLLIGNON, M. 1962. Atlas des fossiles carctéristiques de Madagascar (Ammonites) LX (Aptien). Tananarive: Service Géologique. DrusHcuitz, V. V. & KupryAvtTsevA, M. P. 1960. Atlas of Lower Cretaceous fauna of the northern Caucasus and Crimea. Moscow: Gostoptekhizdat. (In Russian.) DusourDieu, G. 1956. Etude géologique de la région de l’OQuenza (Confins Algéro-Tunisiens). Publs Serv. Carte géol. Algér. 10: 1-659. ETHERIDGE, R. 1892. In: Jack, R. L. & ETHERIDGE, R. The geology and palaeontology of Queensland and New Guinea. Brisbane & London. Favre, F. 1908. Die Ammoniten der unteren Kreide Patagoniens. Neues Jb. Min. Geol. Beil. Bd. 35: 601-647. Haas, O. 1942. The Vernay collection of Cretaceous (Albian) ammonites from Angola. Bull. Amer. Mus. nat. Hist. 81: 1-224. HENDERSON, R. A. 1973. Clarence and Raukumara Series (Albian- ?Santonian) Ammonoidea from New Zealand. J] R. Soc. N.Z. 3: 71-123. HENNIG, E. 1932. Wesen und Wege der Paldontologie. Berlin. Howarth, M. K. 1958. Upper Jurassic and Cretaceous ammonite faunas of Alexander and Graham Land. Scient. Rep. Falkld Isl. Depend. Sury. 21: 1-16. HUuDLEsTON, W. H. 1890. Further notes on some Mollusca from South Australia. Geol. Mag. (3) 7: 241-246. Hyatt, A. 1903. Pseudoceratites of the Cretaceous. STANTON, T. W. ed. Monogr. U.S. geol. Surv. 44: 1-351. KENNEDY, W. J. & CoBBAN, W. A. 1976. Aspects of ammonite biology, biogeography & biostratigraphy. Spec. Pap. Palaeont. 8: 1-133. KENNEDY, W. J. & JuIGNET, P. 1973. First record of the ammonite family Binneyitidae Reeside, 1927 in western Europe. J. Paleont. 47: 900-902. KENNEDY, W. J. & KLINGER, H. C. 1975. Cretaceous faunas from Zululand and Natal, South Africa. Introduction, stratigraphy. Bull. Br. Mus. nat. Hist. (Geol.) 25: 263-315. KoENEN, A. von. 1902. Die Ammonitiden des Norddeutschen Neocom. (Valanginien, Hauterivien, Barrémien und Aptien). Abh. preuss. geol. Landesanst. n.s. 24: 1-451. ANNALS OF THE SOUTH AFRICAN MUSEUM 121 KULLMAN, J. & WIEDMANN, J. 1970. Significance of sutures in phylogeny of Ammonoidea Paleont. Contr. Univ. Kans. 47: 1-32. LEANZA, A. F. 1970. Ammonites nuevos 0 pocos conocidos del Aptiano, Albiano y Cenomanino de los Andes australes con notas acera de su posicion estratigrafica. Revfa Soc. geol. argent. 25: 197-261. MAkowskI, H. 1962. Problems of sexual dimorphism in ammonites. Pa/aeont. pol. 12: 1-92. Morrow, A. L. 1935. Cephalopods from the Upper Cretaceous of Kansas. J. Paleont. 9: 463-473. OrBiGNy, A. D’. 1840-1842. Paléontologie francaise: Terrains crétacés. 1. Céphalopodes. Paris: Masson. PALFRAMAN, F. F. B. 1969. Taxonomy and sexual dimorphism in ammonites. Morphogenetic evidence in Hecticoceras brightii (Pratt). In: WESTERMANN, G. E. G. ed. Sexual dimorphism in fossil Metazoa and taxonomic implications: 126-154. Stuttgart: Schweizerbart’sche Verlagsbuchhandlung. REESIDE, J. B. 1927. Cephalopods from the lower part of the Cody Shale of Oregon Basin, Wyoming. Prof. pap. U.S. geol. Surv. 150-A: 1-19. REYMENT, R. A. 1955. The Cretaceous Ammonoidea of southern Nigeria and the Southern Cameroons. Bull. geol. Surv. Nigeria 25: 1-112. SaRASIN, C. 1893. Etude sur les Oppelia du groupe de nisus et les Sonneratia du groupe du bivurvatus et du raresulcatus. Bull. Soc. géol. Fr. (4) 21: 149-164. SAZONOVA, I. G. 1958. Lower Cretaceous deposits of the Russian Platform. Jn: FLerovot, O. V. Mesozoic and Tertiary deposits of the central regions of the Russian Platform. Vses. Nauchno-Issled. Geol. Razved. Neft. Inst. Moscow: 31-136. (In Russian.) Sinzow, I. 1870. A geological sketch of the Saratow region. Bull. Soc. Mem. St. Petersb. (2) 5: 103-161. (In Russian.) Sinzow, I. 1898. Bemerkungen iiber einige Ammoniten des Aptien. Odessa. Sornay, J. 1955. Ammonites nouvelles du Crétacé de la région des Monts du Méllegue (Constantine). Bull. Serv. Carte géol. Alger (1) Paléontologie 18: 1-40. SpATH, L. F. 1923. A Monograph of the ammonoidea of the Gault. Part 1. Palaeontogr. Soc. Monogr. 76: 1-72. STEPHENSON, L. W. 1952. Larger invertebrate fossils of the Woodbine Formation (Cenomanian) of Texas. Prof. pap. U.S. geol Surv. 242: 1-226. STEPHENSON, L. W. 1955. Basal Eagle Ford fauna (Cenomanian) in Johnson and Tarrant counties, Texas. Prof. pap. U.S. geol. Surv. 274-C: 53-67. STOLLEY, E. 1907. Ueber ein norddeutsches aequivalent der Clansayes Fauna Siidfrankreichs und der Schweiz. Centrlbl. Min. Geol. u Pal. 1907: 266-270. TENISON-Woops, J. E. 1883. On some Mesozoic fossils from the Palmer River, Queensland. J. Proc. R. Soc. N.S.W. 16: 147-154. THomson, M. R. A. 1974. Ammonite faunas of the Lower Cretaceous of south-eastern Alexander Island. Scient. Rep. Falkld Isl. Depend. Surv. 80: 1-44. TRAUTSCHOLD, H. 1865. Der Inoceramen-Thon von Simbirsk. Bull. Soc. Imp. Nat. Moscou. 34: 432-457. WarrEN, P. S. & STELCK, C. R. 1958. Lower Cenomanian Ammonoidea and Pelecypoda from Peace River area, western Canada. Bull. geol. Div. Res. Coun. Alberta 2: 36-51. WEDEKIND, R. 1916. Uber Lobus, Suturallobus und Inzision. Zentbl. Miner. Geol. Paléont. 1916: 185-195. WHITEHOUSE, F. W. 1926. The Cretaceous Ammonoidea of eastern Australia. Mem. Qd Mus. 8: 195-242. WuiteHouse, F. W. 1927a. Additions to the Cretaceous ammonite fauna of eastern Australia. Part 1. (Simberskitidae, Aconeceratidae and Parahoplitidae). Mem. Qd Mus. 9: 109-120. WuiteHouse, F. W. 1927b. Additions to the Cretaceous ammonite fauna of eastern Australia. Part 2. Mem. Qd Mus. 9: 200-206. WRIGHT, C. W. 1957. Mesazoic Ammonoidea. In: Moore, R. C. ed. Treatise on Invertebrate Palaeontology, Part L, Mollusca 4: Kansas; New York: University of Kansas Press and Geological Society of America. WriGHT, C. W. 1963. Cretaceous ammonites from Bathurst Island, northern Australia. Palaeontology 6: 597-614. 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. 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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. Turere, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: 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 77 Band February 1979 February Part 7 . Deel A NEW GENUS AND SPECIES OF THE PENAEOID FAMILY SOLENOCERIDAE (CRUSTACEA, DECAPODA) FROM SOUTH-EAST AFRICAN WATERS By ANTONIO J. DE FREITAS 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, 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.), 714), 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 Copyright enquiries to the South African Museum Kopieregnavrae na die Suid-Afrikaanse Museum ISBN 0 908407 64 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 A NEW GENUS AND SPECIES OF THE PENAEOID FAMILY SOLENOCERIDAE (CRUSTACEA, DECAPODA) FROM SOUTH-EAST AFRICAN WATERS By ANTONIO J. DE FREITAS Oceanographic Research Institute, Durban (With | figure and 1 table) [MS accepted 31 October 1978] ABSTRACT A new genus, Cryptopenaeus, is proposed for a new solenocerid species, Cryptopenaeus catherinae, which is described and illustrated. The new genus is related to Hymenopenaeus and Haliporoides formerly belonging to the genus Hymenopenaeus, sensu lato. C. catherinae has so far been found only in three localities off southern Mozambique at depths of 310-500 metres. CONTENTS PAGE Introduction ae ant. Pe Cryptopenaeus gen. nov. . : : 2 2s Cryptopenaeus catherinae sp. nov. . a0 125 Acknowledgements . . . ... 130 References..." eae ok! Bee 180 INTRODUCTION In June 1969, while collecting penaeoid shrimps aboard a trawler operating in deep water off southern Mozambique, one male specimen representing a new genus and species was caught in 350 metres of water. Further male specimens were subsequently found in the same area in June 1973, and the only female available was caught in September 1976 at a depth of 500 metres. All specimens were caught together with the commercially important pink or knife prawn Haliporoides triarthrus, as well as the less common Aristeomorpha foliacea and Penaeopsis balssi. Genus Cryptopenaeus gen. nov. Diagnosis Body robust; carapace elongate; integument firm. Rostrum short, not reaching distal margin of first antennular segment; ventral margin moderately convex; armed only with dorsal teeth; epigastric tooth and first rostral separated by interval equal to or only slightly greater than that between first and second rostral teeth. Orbital, suprahepatic and branchiostegal spines absent; antennal, postorbital, hepatic and pterygostomian spines present; cervical sulcus deep, long but not reaching mid-dorsum of carapace; hepatic sulcus deep and long, bending anteroventrally from horizontal posterior part and almost reaching 123 Ann. S. Afr. Mus. 77 (7), 1979: 123-131, 1 fig., 1 table. 124 ANNALS OF THE SOUTH AFRICAN MUSEUM base of pterygostomian spine; orbito-antennal groove shallow and wide; branchiocardiac carina distinct but not sharp; mid-dorsal abdominal carina present on segments two to six. Telson with pair of short fixed spines ; no movable marginal spines. Prosartema narrow, long, extending beyond end of eye. Antennular flagella similar, subcylindrical and equal to, or slightly longer than, carapace. Mandibular palp two jointed; articles subequal in length, distal one narrower than basal and tapering to rounded apex. Exopodites on all maxillipeds and pereiopods. Lateral ramus of uropod with very small, blunt distolateral spine. Petasma with ventrolateral lobule entirely occupied by ventral costa and distally free from dorsolateral lobule; both ventrolateral and dorsolateral lobules heavily sclerotinized ; appendix masculina and appendix interna present. Thelycum simple, of open type. Podobranch on maxilliped II only; epipodites on maxillipeds II and III and on pereiopods I-IV. Type species Cryptopenaeus catherinae sp. nov. Etymology The generic name is derived from the prefix crypto, from the Greek kryptos meaning hidden, in combination with the generic name Penaeus, denoting the fact that this shrimp has been hidden from science until now; gender masculine. Taxonomic status and comments From the works of Bate (1881, 1888), Bouvier (1906), and Burkenroad (1936), it is clear that the generic complex forming the then accepted subfamily Solenocerinae presented many taxonomic difficulties. This subfamily consisted of three genera: Solenocera, shrimps with concave antennular flagella (Lucas 1849; Wood-Mason & Alcock 1891; Barnard 1950); Haliporus, shrimps with subcylindrical antennular flagella and movable lateral spines on the telson anterior to fixed pair (Bate 1881; Kensley 1968); and Hymenopenaeus, shrimps with subcylindrical antennular flagella and lacking lateral spines on the telson (Smith 1882; Wood-Mason & Alcock 1891; Barnard 1950). It is apparent from the literature, however, that Burkenroad (1936) was somewhat unhappy with the taxonomic status of Hymenopenaeus and went to the point of dividing the genus into four superspecies ‘. . . according to the presence or absence of branchiostegal or pterygostomian spines and to the nature of the postrostral armature’. Pérez Farfante (1977) revised the subfamily and, besides proposing that the subfamilies hitherto accepted should be elevated to the category of families of the superfamily Penaeoidea, divided Hymeno- penaeus into five genera partly based on the superspecific groups elaborated by Burkenroad (1936). In doing so Pérez Farfante takes into consideration the ‘... Shape of the antennular flagella and rostrum, proportions of the carapace, number and comparative size of the articles of the mandibular palp, presence or absence of certain carinae on the carapace, relative dimensions of the posterior A NEW GENUS AND SPECIES OF THE PENAEOID FAMILY SOLENOCERIDAE 125 two pairs of pereiopods, location of the distolateral spine of the lateral ramus of the uropod, structure of the petasma and degree of development of the arthrobranchia on somite VII’, as well as those characteristics originally used by Burkenroad. The system presented is very comprehensive and covers the previously known species extremely well. However, whereas the six specimens caught in southern Mozambique belong clearly to the family Solenoceridae and would have fitted into the genus Hymenopenaeus sensu lato, they do not belong to any of the five genera established by Pérez Farfante (1977), and have therefore been placed in a new genus, Cryptopenaeus. Table 1 sets out the similarities and differences between Cryptopenaeus and the other five genera, Hymenopenaeus, Haliporoides, Pleoticus, Hadropenaeus, and Mesopenaeus. The new genus is closely allied to Hymenopenaeus and Haliporoides but differs from the former in the arrangement of the rostral teeth, the absence of branchiostegal spines, the presence of a mid-dorsal carina on abdominal segments 2 and 3 and also by the short rostrum with a strongly convex ventral margin. Cryptopenaeus differs from Haliporoides in the arrange- ment of the rostral teeth, presence of a dorsal carina on the second abdominal segment, by the absence of a suprahepatic spine and by the short rostrum with a strongly convex ventral margin. The petasma of Cryptopenaeus differs from that of the other five genera in having the ventrolateral lobule entirely occupied by the ventral costa. Cryptopenaeus catherinae sp. nov. Fig. 1 Material Holotype: SAM-A16148 in the South African Museum, Cape Town, 3, 46,7 mm carapace length, caught off Cape Santa Maria in southern Mozam- bique (26°06’S 33°08’E) at a depth of 350 metres, on 16 December 1969. Alloptype: SAM-A16149 in the South African Museum, Cape Town, 2, 63,2 mm carapace length, caught off Monte Bello in southern Mozambique (25°00’S 35°21’E) at a depth of 500 metres, in September 1976. Paratypes: 4 gg, 44,5-47,7 mm carapace length, caught off southern Mozambique at a depth of 310 metres, in June 1973. One male paratype is in the National Museum of Natural History in Washington, D.C., and the remaining three will be sent to the British Museum (Natural History). Description Rostrum. Slightly downwardly directed, reaching to or just beyond end of first antennular segment; ventral margin convex; rostral teeth 74°; epigastric and three other teeth situated behind postorbital margin of carapace; adrostral carina short, just reaching postorbital margin; postrostral carina very well developed, long, almost reaching posterior margin of carapace and with conspicuous notch behind epigastric tooth; median groove absent. ANNALS OF THE SOUTH AFRICAN MUSEUM 126 xOAU09 xOAUO09 pur }10Y4§ A[SUO01S juosolg quosqy JUosold juosqy juosqy juesqy quosqy yuasold quosold quosqy juosqy quosqy X9AU09 SABDUOD OF} A[3uO.NS WYsIeNS juasolg juasqe 10 JUdSI.1g juasolg juosqy 9-C one S]PAIOJUI S[BA1O}UL SUISBOIDOp SuIsvaIDOp Ajae[nso. Ajavpnso. Aq poyeredag Aq poyeredog BIOUIT Po}LjoI A[OSO[D DAY OY} WIJ “AOU “Ud SnapuadojddsD Burysinsurjsip sonstiojovsreyO S[BAIOJUL S[BAIOJUI SUISvoIDOp SUISvOIDOp Ajre[nso. Ajarepnso. Aq poyeredag Aq poyeredag snanuadospoyy SnINo0ad y}00} [e11sOl 1S] WOJJ poyeiedos oLsesidg saplosodyopy] ‘ . [ a1avL SABIUOD 0} WysIeI}s ‘SuOT WYsIeNS juasqe 10 jUdSa1d quosold 9-1 puz woul poyeredos 4199} [B1}SOI 1ST pure o1ynsesidq snapuadouaud FT WINJ}sOJ JO UISIVU [RUD A, & BUIIeO [BUISIeWUGnS * BULIeS SRIPIBOOIyouLI_ ouids onedoyeidngs ouids [eIGIO ee ourlds UPIWIOJSOSAIN sf curds [esojsorpourig BUIIvO [eSIOp [eUILIOpqy 4199} [eIIsOI pue o1I\sesidq A NEW GENUS AND SPECIES OF THE PENAEOID FAMILY SOLENOCERIDAE 127 Carapace. Uniformly glabrous and lightly punctate; gastrofrontal and postocular grooves and horizontal suture absent; no supra-orbital spine; cervical sulcus and carina very well developed and long but not reaching dorsal midline; cervical carina terminating anteroventrally in prominent hepatic spine; gastro-orbital carina absent; postorbital spine prominent; antennal spine present but relatively small; antennal carina absent; orbito-antennal groove restricted to wide depression extending from below postorbital spine to below hepatic spine; hepatic carina sharp, anteroventrally directed and situated just anterior and below hepatic spine; hepatic sulcus wide and deep; extending posteriorly below hepatic spine; branchiocardiac carina distinct but not very sharp. Branchiostegal spine absent; pterygostomian spine prominent and sharp; submarginal carina long and sharp; no vertical suture or carina. Antennule. Flagella subequal in length; about 2,5 times length of antennular peduncle; subcylindrical; mesial flagella twice as thick as lateral flagella; prosartema flexible with pointed apex, reaching just beyond distal end of first antennular peduncle, copiously provided with long setae; stylocerite sharply pointed distally, straight and reaching just beyond end of eye or to end of first antennular article; distolateral spine prominent and long; parapenaeid spine absent. Scaphocerite. Distolateral spine reaching just beyond distal end of anten- nular peduncle; apex of lamella extending beyond distolateral spine; basicerite with single broad blunt tooth distally. Mandibular palp. Reaching to about basal one-third of carpocerite; proximal article 1,8 times as long as wide; distal article subequal to proximal, tapering to rounded apex. Maxilliped III. Endopodite not exhibiting sexual dimorphism; reaching distal end of scaphocerite; exopodite short, reaching less than half-way along merus of endopodites; epipodite present. Pereiopods. Exopodites present on all pereiopods, well developed, longest on first periopod and shortest on fifth; epipodites present on pereiopods I-IV; basipodites of first, second and third with prominent spines; ischial spine present on first pereiopod only; distinct coxal spine on fifth pereiopod. Pereiopod IV reaching to apex of mandibular palp; pereiopod V reaching to distal end of antennular peduncle. Extended laterally lengths of pereiopods in ascending order are: first, second and fourth, third and fifth. Abdomen. Uniformly glabrous; mid-dorsal carina present from posterior half of second segment to end of sixth where it terminates in short spine; short vertical groove on pleura of first segment; lateral carinae absent. Telson. Slightly longer than sixth segment; about as long as mesial ramus of uropod; median groove deep, occupying only anterior half of telson; apical spine somewhat blunt; pair of inconspicuous, very short, fixed subapical spines present; movable marginal spines lacking. Thelycum. Simple open structure; anterior portion formed by vertical posterior face of sternite between fourth pereiopods; posterior face with short 128 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 1. Cryptopenaeus catherinae sp. noy. A. Lateral view, holotype ¢ 46,7 mm carapace length. B. Right prosartema. C. Telson. D. Left mandibular palp, ventral view. E. Left appendix masculina and interna, ventral view. F. Appendix masculina, dorsal view. G. Distal subquadrate process of right half of petasma. H. Petasma, ventral view. I. Thelycum. A NEW GENUS AND SPECIES OF THE PENAEOID FAMILY SOLENOCERIDAE 129 median groove and low lateral ridges; covered with short setae and obscured by coxal protuberances of fourth pereiopods. Posterior portion (between fifth pereiopods) consisting of elongate plate; broad central ridge occupying slightly more than anterior two-thirds of plate; well-defined lateral ridges extending from posterior sternal process to anterior margin of somite; two suboval, setose, boss-like structures present between anterior third of lateral ridges and median ridge; anterior margin of posterior sternal process interrupted by deep median sulcus. Petasma. Simple and very slightly involuted; dorsomedian lobule about two-fifths of total length of petasma; entirely united along midline; ventro- median lobule elongate and subtriangular; inner membranous section folded slightly on itself; heavily sclerotinized central ridge running length of lobule terminating distally in thick subquadrate process; distal margin smooth; proximolateral angle of quadrate process beak-like with four or five blunt teeth; ventral face of process concave; dorsal face convex; dorsolateral lobule elongate and subtriangular; apex situated under proximolateral angle of distal ventro- median process; lower inner angle forms small proximal process; ventral surface sparsely covered with long setae; ventral costa (occupying entire ventrolateral lobule) extending along lateral margin of petasma reaching half way into distal ventromedian process; apex free and bi-lobed; dorsal lobe longer than ventral lobe; ventral lobe with one to four minute teeth. Appendix masculina. Dorsally convex, ventrally concave, roughly trapezoid in shape; distal margin with row of short, stout setae; appendix interna subequal in length to appendix masculina, half fitting into concave face of latter; elongate with concave median surface; apex with short stout setae. Basal segment of endopodite of pleopod II as wide as long, its distolateral portion concave, subtriangular and produced into long, blunt spur. Colour in life. Body generally red, carapace with broad white stripe running from below hepatic sulcus to almost posterior margin of carapace on each side; this stripe wider in posterior half of carapace; on abdominal segments brighter red patch running anteroventrally on each pleuron; distinct white longitudinal stripe on dorsal carina of abdominal segments 4-6; telson and uropods pinkish white becoming red along posterior edges; lateral margins of scaphocerite red; rostral crest and pereiopods pinkish white; basal segments of pleopods grey to white; pleopodal endopodites greyish white becoming bright red distally; marginal setae of pleura, pleopods and uropods orange while those of scapho- cerite and antennules white. Distribution Known only from the type locality. Ecological notes All the specimens were collected from the same general area, namely the Limpopo Bight in southern Mozambique. This is an important fishing ground 130 ANNALS OF THE SOUTH AFRICAN MUSEUM where the main species of interest are the spiny lobster, Palinurus delagoae, caught in about 280-350 metres, the langoustine, Nephrops andamanicus, trawled in 320-380 metres, and the pink or knife prawn, Haliporoides triarthrus, found in 300-500 metres. In this area, the continental slope is relatively gentle down to 500 metres after which it drops very steeply to about 3 000 metres. The substrate consists of muddy sand to sandy mud and the temperature at 300-500 metres depth recorded in April and September 1964 was 11-13°C (Instituto Hidrografico Lisbon 1965, 1967). Although Cryptopenaeus catherinae has so far been found only in the type locality, it seems feasible to expect that, as its associate species, Haliporoides triarthrus, Aristeomorpha foliacea and Penaeopsis balssi are found off the coast of Natal, the distribution of C. catherinae could possibly extend southward as well. ACKNOWLEDGEMENTS I wish to thank the Fisheries Development Corporation and the South African Association for Marine Biological Research for their financial and administrative assistance; Dr Isabel Pérez Farfante of the Systematics Laboratory, National Marine Fisheries Service, Washington D.C., U.S.A., for having checked the identity of this species and for the meticulous way in which she read and criticized my manuscript; Professor A. E. F. Heydorn, Director of the Oceanographic Research Institute, Durban, for his encouragement and for his criticism of the manuscript; Dr P. F. Berry for reading and criticizing the paper; and finally my wife, after whom this species is named, for her encourage- ment, patience and understanding. REFERENCES BARNARD, K. H. 1950. Descriptive catalogue of South African decapod crustacea. Ann. S. Afr. Mus. 38: 1-837. Bate, C. S. 1881. On the Penaeidae. Ann. Mag. nat. Hist. (5) 8: 169-196. Bate, C. S. 1888. Report on the Crustacea Macrura collected by H.M.S. Challenger during the years 1873-1876. Rep. Sci. Results Voyage H.M.S. Challenger 1873-76, Zool. 24: 1-942. Bouvier, E. L. 1906. Observations sur les Peneides du genre Haliporus Bate. Bull. Mus. Oceanogr., Monaco 81: 1-10. BURKENROAD, M. D. 1936. The Aristaeinae, Solerocerinae, and pelagic Penaeinae of the Bingham Oceanographic Collection. Bull. Bingham Oceanogr. Coll. 5(2): 1-151. INstiITUTO HiprRoGRAFICO LisBon. 1965. Resultados das observagées oceanograficas no Canal de Mocambique, Cruzeiro AL 1/64: Abril-Maio 1964 1: 1-73. INstrTuTO HipROGRAFICO LisBoN. 1967. Resultados das observagées oceanograficas no Canal de Mocgambique, Cruzeiro AL 2/64: Setembro-Outubro 1964 3: 1-107. KENSLEY, B. F. 1968. Deep sea decapod Crustacea from west of Cape Point, South Africa. Ann. S. Afr. Mus. 50: 283-323. Lucas, P. H. 1849. Genus Solenocera Lucas. Rev. Mag. Zool. (2) 1: 300. PEREZ FARFANTE, I. 1977. American solenocerid shrimps of the genera Hymenopenaeus, Haliporoides, Pleoticus, Hadropenaeus new genus, and Mesopenaeus new genus. Fish. Bull. 75: 261-346. A NEW GENUS AND SPECIES OF THE PENAEOID FAMILY SOLENOCERIDAE 131 SmiTH, S. I. 1882. Report on the results of dredging, under the supervision of Alexander Agassiz, on the East Coast of the United States, during the summer of 1880 by the U.S. Coast Survey Steamer ‘Blake’. 17: Report on the Crustacea, 1: Decapoda. Bull. Mus. comp. Zool., Harv. 10: 1-108. Woop-Mason, J. & Atcock, A. 1891. Natural history notes from H.M. Indian Marine Survey steamer ‘Investigator’, Commander R. F. Hoskyn, R.N., commanding. No 21. Note on the results of the last season’s deep-sea dredging. Ann. Mag. nat. Hist. (6) 8: 268-286. ere a g , a at ae ry) ’ - ar lseorw is - a 2? Ath 4. Vi! 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: Leda bicuspidata: Nicklés, 1950: is, 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. ANTONIO J. DE FREITAS A NEW GENUS AND SPECIES OF THE PENAEOID FAMILY SOLENOCERIDAE (CRUSTACEA, DECAPODA) FROM SOUTH-EAST AFRICAN WATERS Bf nkleirw VOLUME 77 PART 8 FEBRUARY 1979 ISSN 0303-2515 [32 A me fe "Ac Y = APR 1 3 1979 ( OUTH AFRICAN Sa a MUSEUM JAPE 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 (th) 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, 19696; Jones 1971)’ ‘As described (Haughton & Broom 1927)...’ ‘As described (Haughton ef 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) BuLLouGu, 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. 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) ee 2S 6 ES <= PS es = Ow ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 77 Band February 1979 Februarie Part 8 _ Deel YPE SPECIMENS OF HYDROIDA (COELENTERATA) IN THE SOUTH AFRICAN MUSEUM 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), 3(1-2, 4-5, 8, t.-p.i.), 5(1-3, 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 Copyright enquiries to the South African Museum Kopieregnavrae na die Suid-Afrikaanse Museum ISBN 0 908407 62 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 TYPE SPECIMENS OF HYDROIDA (COELENTERATA) IN THE SOUTH AFRICAN MUSEUM By N. A. H. MILLARD South African Museum, Cape Town [MS. accepted 31 October 1978] ABSTRACT A list is given of seventy-nine types of Hydroida housed in the South African Museum, including spirit material, whole mounts and microscope sections. CONTENTS PAGE Introduction : : : : He yile's Species list . : Ge gt: : co! 134 References . : : : 2 . 149 INTRODUCTION This paper lists the type holdings of Hydroida in the South African Museum, and includes not only material preserved in alcohol, but also whole mounts and sections prepared for microscopic examination. The arrangement of families, genera and species is the same as that used by Millard (1975). Each specimen has a South African Museum registered number (prefixed by the letters SAM-H), and in most cases a second number is quoted too, which is that of the coilector’s catalogue. For the latter the following abbreviations are used: UCT University of Cape Town; material from the Ecological Survey of the intertidal rocky shore, the estuaries and the benthic region of the South African seas. It should be noted that most of the U.C.T. collection of hydroids is now housed in the South African Museum. PF Material collected by the old Government Survey vessel, the R.S. Pieter Faure. The positions given in the Pieter Faure catalogue were not very reliable, but have been converted here as accurately as possible into latitude/longitude positions. MD _ Duplicate material from the Museum National d’Histoire Naturelle, Paris, collected during the cruise MD.03 of the Marion-Dufresne (Millard 1977a). MT Duplicate material from the collection of the Musée Royal de I’Afrique Centrale, Tervuren, Belgium, collected by J. Bouillon (Millard & Bouillon 1973, 1974, 1975). 133 Ann. S. Afr. Mus. 77 (8), 1979: 133-150. 134 ANNALS OF THE SOUTH AFRICAN MUSEUM SM __— Material collected by the R.V. Meiring Naude off the east coast of South Africa (Millard 19775). TME Duplicate material from the Universitetets Zoologiske Museum, Copen- hagen, collected during Th. Mortensen’s Java—South Africa Expedition, 1929-1930 (Millard 1968). SPECIES LIST Suborder ATHECATA Family Myriothelidae Monocoryne minor Millard, 1966: 435, fig. 1. Holotype: SAM-—H417 (UCT SCD 215C). One fertile specimen detached from substratum. Locality: Agulhas Bank, 34°03’S 25°58’E; 78 m; 25 November 1960. Collected by: University of Cape Town. Mpriothela tentaculata Millard, 1966: 437, fig. 2. Holotype: SAM-—H418 (UCT WCD 7Q). One male individual on encrusting bryozoan. Locality: Off west coast of Cape Peninsula, 34°09,3’S 18°17,5’E; 43 m; 24 March 1959. Paratype: SAM-H2915 (UCT WCD 7Q). Microscope sections of male individual; locality as holotype. Collected by: University of Cape Town. Family Corynidae Bicorona elegans Millard, 1966: 441, fig. 3. Holotype: SAM-H419 (UCT SB 161X). Rich male and female colonies; plus one whole mount; plus microscope sections. Locality: Saldanha Bay, 33°02,5’S 18°02’E; littoral; September 1957. Collected by: University of Cape Town. This is the type species of the genus Bicorona Millard, 1966. Family Eudendriidae Eudendrium deciduum Millard, 1957: 184, fig. 2. Holotype: SAM-H98 (UCT FAL 52V). Male and female colonies; plus one whole mount; plus microscope sections. Locality: False Bay, from 34°09,3’S 18°49,6’E to 34°09’S 18°50,1’E; 18 m; 25 June 1952. Collected by: University of Cape Town. TYPE SPECIMENS OF HYDROIDA IN THE SOUTH AFRICAN MUSEUM 135 Eudendrium ritchiei Millard, 1975: 87, fig. 30. Holotype: SAM-H1803. Abundant male and female colonies; plus two whole mounts. Locality: Saunders Rocks, Sea Point; littoral; 5 March 1973. Collected by: N. A. H. Millard. Family Bougainvilliidae Clavopsella quadranularia Thiel, 1962: 227, figs 1-28. = Clavopsella navis: Millard, 1975: 100, fig. 34A—D. (See also Rhizorhagium navis.) Schizoparatypes: SAM-H1774, H1775 (presented by Dr H. Thiel 1972). Locality: Kiel Canal. Rhizorhagium navis Millard, 1959b: 244, fig. 2. = Clavopsella navis: Millard, 1975: 100, fig. 34A—D. (See also Clavopsella quadranularia.) Holotype: SAM-—H124 (UCT SH 429B). Male and female colonies on weed and other hydroids; plus one whole mount. Locality: Cape Town docks, on ship’s hull; 10 February 1958. Collected by: N. A. H. Millard. Silhouetta uvacarpa Millard & Bouillon, 1973: 25, fig. 3A—D, pls 2-3. Schizoholotype: SAM-H2916 (MT 2664). Three whole mounts only, bearing medusa buds. Locality: Silhouette Island, Seychelles; 1966. Collected by J. Bouillon. This is the type species of the genus Silhouetta Millard & Bouillon, 1973. Family Hydractiniidae Clavactinia multitentaculata Millard, 1975: 106, fig. 35C-G. As Hydractinia sp.: Millard, 1968: 255. Holotype: SAM-—H389 (PF station 10724). Male colony on shell of gastro- pod Melapium lineatum (Lamarck), occupied by hermit Dardanus arrosor (Herbst); plus two whole mounts; plus microscope sections. Locality: off Natal, approx. 29°54’S 31°11’E; 99 m; 14 December 1900. Collected by: R.S. Pieter Faure. 136 ANNALS OF THE SOUTH AFRICAN MUSEUM Hydractinia altispina Millard, 1955: 215, fig. 1. Lectotype: SAM-—H87 (UCT F 274). Male colony on gastropod Thais squamosa (Lamarck); plus microscope sections. Locality: St James, False Bay; littoral; 1 August 1936. Paralectotypes: SAM-—H88 (UCT CP 258). Infertile colonies on two specimens of Thais squamosa; plus one whole mount; St James, False Bay; littoral; April, 1944. SAM-H89 (UCT B 92). Colonies with very young gonophores on two specimens of Thais squamosa; Lambert’s Bay; littoral; 30 July 1938. Collected by: University of Cape Town. Hydractinia canalifera Millard, 1957: 179, fig. 1. Holotype: SAM—H97 (UCT CP 332). Female colony on weed; plus one whole mount; plus microscope sections. Locality: Clovelly, False Bay; littoral; 23 August 1949. Collected by: N. A. H. Millard. Hydractinia diogenes Millard, 1959a: 305, fig. 2. Holotype: SAM-H123 (UCT MOR SIH). Male and female colonies on five gastropod shells occupied by hermits Diogenes costatus (Fabricius); plus three whole mounts; plus microscope sections. Locality: off mouth of Rio Coche, Mozambique; 3—5 m; 21 January 1954. Collected by: University of Cape Town. Hydractinia kaffraria Millard, 1955: 217, fig. 2. Lectotype: SAM—H90 (UCT BRE 111A). Female colony on gastropod Nassarius kraussianus (Dunker). Locality: Breede River estuary; 3 February 1952. Paralectotypes: SAM-—H92 (UCT SUN 3N). Colonies on five specimens of Nassarius kraussianus; plus one whole mount; Sunday’s River estuary; 7 January 1950. SAM-H91 (UCT HAM 3Q). Microscope sections of female colony; Keiskama River estuary, The Haven; 9 January 1950. Collected by: University of Cape Town. Hydractinia marsupialia Millard, 1975: 113, fig. 38. Holotype: SAM-H1854 (UCT TB 13). Female colony on gastropod Nassarius speciosus Adams; plus microscope sections of male and female colonies. Locality: Table Bay; 9-18 m; 4 August 1946. TYPE SPECIMENS OF HYDROIDA IN THE SOUTH AFRICAN MUSEUM 137 Paratypes: SAM-H1855 (UCT TB 14 plus 15). Nine colonies, male and female, on Nassarius speciosus; plus one whole mount; Table Bay; 17-20 m; 25 October 1946 and 11 February 1947. Collected by: University of Cape Town. Family Cytaeidae Podocoryne nassa Millard, 1959a: 307, fig. 3. = Cytaeis nassa: Rees, 1962: 390, figs 8-9, pl. 11. Holotype: SAM-H122 (UCT IN 112). Fertile colonies on two shells of gastropod Nassarius fenestratus Marrat; plus one whole mount; plus microscope sections. Locality: Punta Torres, Inhaca Island, Mozambique; littoral; 18 September 1955. Collected by: University of the Witwatersrand. Suborder THECATA Family Campanulinidae Aequorea africana Millard, 1966: 461, fig. 8. Holotype: SAM—H420 (UCT MB 70G). Fertile colony on empty gastropod shell; plus two whole mounts. Locality: Mossel Bay, 34°08,9’S 22°07,9’E; 18 m; 19 January 1956. Collected by: University of Cape Town. Egmundella amirantensis Millard & Bouillon, 1973: 40, fig. SA—D. Schizoholotype: SAM-—H2917 (MT 2665). One whole mount only of fertile material. Locality: Amirante Island, Seychelles; 1966. Collected by: J. Bouillon. Lineolaria gravierae Millard, 1975: 134, fig. 43H. As Lineolaria sp.: Gravier, 1970: 144, figs 11, 13A. Millard & Bouillon, 1974: 22, fig. 2D. Schizoholotype: SAM-—H1955. One whole mount (infertile) only. Locality: Barreira Vermelha, Inhaca, Mozambique, 26°02’S 32°54’E; 1969. Collected by: J. Bouillon. 138 ANNALS OF THE SOUTH AFRICAN MUSEUM Lovenella chiquitita Millard, 1957: 198, fig. 7. Holotype: SAM-—H103 (UCT FAL 288J). One whole mount only of fertile colony epizootic on Eudendrium deciduum Millard. Locality: False Bay, 34°09,6’S 18°49,8’E; 18 m; 15 October 1897. Collector unknown. Paratypes: SAM-H104 (UCT FB 131K). Two whole mounts (infertile), one epizootic on Sertularella polyzonias falsa Millard; False Bay, 34°09’S 18°26,7’E; 5-8 m; 13 December 1949. SAM-H105 (UCT FAL 1080). One whole mount (infertile) epizootic on Sertularella polyzonias falsa; False Bay, 34°09,3’S 18°51’E; 8-12 m; 23 January 1953. Collected by: University of Cape Town. Family Haleciidae Halecium dufresneae Millard, 1977a: 8, fig. 2A—D. Schizoholotype: SAM-—H2785 (MD station 26/63). Fragments of female colony; plus one whole mount. Locality: Crozet Island, Chenal des Orques, 46°21,5’S 51°55’E; 230 m; 20 April 1974. Collected by: R.V. Marion Dufresne. Halecium inhacae Millard, 1958: 168, fig. 1. Holotype: SAM-—H114 (UCT IN 140H). Female colony on weed; plus two whole mounts. Locality: Inhaca Island, Mozambique, east shore rocks; 20 July 1956. Collected by: University of the Witwatersrand. Halecium parvulum, var. magnum Millard, 1957: 190, fig. 4B—O. Since incorporated in Halecium delicatulum Coughtrey, 1876 (large form). Holotype: SAM-—H99 (UCT FAL 274R). Male and female colonies; plus three whole mounts. Locality: False Bay, 34°10,9’S 18°27,2’E; 14-17 m; 21 September 1954. Collected by: University of Cape Town. , Paratypes: SAM-H100 (UCT FAL 159L). Infertile colony; plus one whole mount; False Bay, 34°12,5’S 18°28’E; 0-3 m; 21 April 1953. Collected by: University of Cape Town. SAM-HI11 (PF station 405). Male colony; plus two whole mounts; off Buffels Bay, False Bay; depth unknown; 8 October 1898. SAM-H30 (PF station 16287). Female colony; plus two whole mounts; off Cape Point, approx. 34°20’S 18°32’E; 59 m; 9 December 1902. Collected by: R.S. Pieter Faure. TYPE SPECIMENS OF HYDROIDA IN THE SOUTH AFRICAN MUSEUM 139 Hydrodendron sympodiformis Millard & Bouillon, 1974: 25, fig. 5. Schizoholotype: SAM-H1732 (MT station InPT 42). Part of a female colony on weed; plus one whole mount. Locality: Punta Torres, Inhaca, Mozambique; August 1969. Collected by: J. Bouillon. Family Lafoeidae Hebella furax Millard, 1957: 200, fig. 8. Holotype: SAM-H34 (PF station 18293). Infertile colony parasitic and epizootic on Lytocarpus filamentosus (Lamarck); plus two whole mounts. Locality: False Bay, approx. 34°05,5’S 18°39’E; 15— 18 m; 7 December 1903. Collected by: R.S. Pieter Faure. Paratype: SAM-H106 (UCT FAL 58Y). Infertile colony on Lytocarpus filamentosus; one whole mount only; False Bay, 34°09,4’S 18°50,4’E; 12 m; 25 June 1952. Collected by: University of Cape Town. Hebella muscensis Millard & Bouillon, 1975: 10, fig. 3A—B. Schizoholotype: SAM-—H2918 (MT station SEY 20K). Two whole mounts only of fertile colony epizootic on Synthecium sp. Locality: Anse la Mouche, Seychelles; 1972. Collected by: J. Bouillon. Hebella urceolata Millard, 1964: 11, fig. 2A. Since incorporated in Hebella scandens (Bale, 1888) Holotype: SAM-H410 (UCT SCD 154H). Infertile colony epizootic on Halecium beanii (Johnston); plus one whole mount. Locality: Agulhas Bank, 34°03’S 25°59’E; 84 m; 25 November 1960. Collected by: University of Cape Town. Scandia tubitheca Millard & Bouillon, 1975: 10. As Scandia corrugata: Millard & Bouillon, 1973: 60 (pro parte), fig. 8D-F. Schizoholotype: SAM-—H2919 (MT station Am IP). Male colony epizootic on Synthecium dentigerum Jarvis; two whole mounts only. Locality: Amirante Island, Seychelles; 30 August 1966. Collected by: J. Bouillon. Zygophylax cornucopia Millard, 1955: 219, fig. 3. = Hydrodendron cornucopia: Millard, 1973: 33, fig. 6A—E (transferred to Haleciidae). Holotype: SAM—H93 (UCT FB 131B). Male and female colonies epizootic on Antennella quadriaurita Ritchie; plus one whole mount. 140 ANNALS OF THE SOUTH AFRICAN MUSEUM Locality: False Bay, 34°09’S 18°26,7’E; 5-8 m; 13 December 1949. Paratypes: SAM-H94 (UCT TB 1B). Infertile colonies epizootic on A. quadriaurita; plus one whole mount; Table Bay, 33°47,5’S 18°24,3’E; 19-20 m; 11 February 1947. SAM-—H95 (UCT FAL 78S). Fertile colonies epizootic on A. quadriaurita: plus one whole mount; False Bay, several positions; 4-17 m; 19 August 1952-23 September 1954. SAM-H96 (UCT FAL 217N); one whole mount of infertile colony epizootic on A. quadriaurita; False Bay, 34°07’S 18°32,5’E; 18 m; 9 September 1953. Collected by: University of Cape Town. Zygophylax crozetensis Millard, 1977a: 15, fig. 4. Schizoholotype: SAM—H2779 (MD station 26/64). Part of a large fertile colony; plus one whole mount. Locality: Crozet Island, Chenal des Orques, 46°24’S 51°59’E; 180 m; 20 April 1974. Collected by: R.V. Marion Dufresne. Zygophylax cnigmatica Millard, 1964: 19, fig. SA—F. = Hydrodendron gracilis (Fraser, 1914): Millard, 1973: 33, fig. 6F—G (trans- ferred to Haleciidae). Holotype: SAM-H411 (UCT WCD 12E); female colony epizootic on Nemertesia ramosa Lamouroux; two whole mounts only. Locality: off Cape Town, 34°09,4’S 18°16,5’E; 75 m; 24 March 1959. Collected by: University of Cape Town. Zygophylax geminocarpa Millard, 1958: 177, fig. 4D-G. Holotype: SAM-HS59 (PF station 12308). About eight fertile stems; plus two whole mounts. Locality: off Natal, approx. 30°53’S 30°28’E; 66 m; 14 March 1901. Collected by: R.S. Pieter Faure. Zygophylax inconstans Millard, 1977b: 117, fig. 5. Holotype: SAM-H1975 (SM station 23J). A fertile, tangled colony on a bryozoan; plus one whole mount. Locality: off Natal, 27°44,4’S 32°42,8’E; 400-450 m; 26 May 1975. Collected by: R.V. Meiring Naude. Zygophylax infundibulum Millard, 1958: 180, fig. 4B—C. Holotype: SAM-—H36 (PF station 10781). Several infertile stems; plus two whole mounts. Locality: off Natal, approx. 29°53’S 31°11’E; 155 m; 17 December 1900. Collected by: R.S. Pieter Faure. TYPE SPECIMENS OF HYDROIDA IN THE SOUTH AFRICAN MUSEUM 141 Family Campanulariidae Campanularia laminacarpa Millard, 1966: 472, fig. 12E-K. Holotype: SAM-—H421 (UCT TRA 32A). A female colony epizootic on Thyroscyphus aequalis Warren; plus two whole mounts. Locality: Agulhas Bank, 34°49’S 20°21,5’E; approx. 91 m; 9 November 1947. Collected by: commercial trawler. Campanularia morgansi Millard, 1957: 195, fig. 6. Holotype: SAM-H24 (PF station 15675). Female colonies epizootic on Corhiza scotiae (Ritchie) and Sertularella arbuscula (Lamouroux); plus one whole mount. Locality: False Bay, approx. 34°28’S 18°32’E; 73 m; 9 October 1902. Collected by: R.S. Pieter Faure. Paratypes: SAM-H7 (PF station 351). Infertile colony epizootic on Salacia articulata (Pallas); plus one whole mount; False Bay, approx. 34°19’S 18°31’E; 58-62 m; 28 September 1898. SAM-H32 (PF station 18232). Fertile colony epizootic on Eudendrium ?deciduum Millard; False Bay, approx. 34°27’S 18°45’E; 110 m; 11 November 1903. Collected by: R.S. Pieter Faure. SAM-H101 (UCT FB 119L plus FAL 289B). Male and infertile colonies epizootic on Salacia articulata (Pallas); plus two whole mounts; False Bay, 34°08,5’S 18°34,5’E; 27 m; 21 April 1947; and another locality in False Bay, exact location and collector unknown; 40 m; September 1897. Collected by: University of Cape Town. SAM-H102 (UCT FAL 26L). Infertile colony on empty tubes; one whole mount only; False Bay, 34°13’S 18°29’E; 15-21 m; 5 March 1952. Collected by: J. C. Morgans. Campanularia pecten Gow & Millard, 1975: 1, fig. 1. Holotype: SAM-H1659. Male colony growing on sea-grass Caulerpa filiformis (Suhr); plus four whole mounts. Locality: St James, False Bay; littoral; 29 March 1968. Collected by: C. Gow. Campanularia roberti Gow & Millard, 1975: 3, fig. 2. Holotype: SAM-H1660. Female colony growing on weed Sargassum longifolium (Turner) attached to the kelp Ecklonia maxima (Osbeck); plus four whole mounts. Locality: Partridge Point, False Bay; 24 March 1968. Collected by: R. W. Day. 142 ANNALS OF THE SOUTH AFRICAN MUSEUM Campanularia subantarctica Millard, 1971: 403, fig. 5. Holotype: SAM—H2920. Two whole mounts only of fertile colony epizootic on Symplectoscyphus marionensis Millard. Locality: Near mouth of Soft Plume River, Marion Island; littoral; 7 February 1965. Collected by: N. R. Fuller. Clytia latitheca Millard & Bouillon, 1973: 55, fig. 7H-L. Schizoholotype: SAM—H2921 (MT 2666). Two whole mounts only of fertile colony. Locality: Praslin Island, Seychelles; 1966. Collected by: J. Bouillon. Clytia serrata Millard, 1958: 173, fig. 3C, H. = Clytia gravieri (Billard): Millard & Bouillon, 1973: 51, fig. 7E-G. Holotype: SAM-H115 (UCT MOR 216C). One whole mount only of infertile colony epizootic on Eudendrium carneum Clarke. Locality: On wreck at Linga Linga, Morrumbene estuary, Mozambique; 15 July 1954. Collected by: University of Cape Town. Family Syntheciidae Hincksella corrugata Millard, 1958: 181, fig. 5. Holotype: SAM-H85 (PF station 12456). Infertile colony on weed; plus one whole mount. Locality: off Natal, approx. 30°32’S 30°38,5’E; 46 m; 22 March 1901. Collected by: R.S. Pieter Faure. Hincksella indiana Millard, 1967: 178, fig. 3D-G. Holotype: SAM-H1646 (UCT AFR 1235A). One infertile stem in three parts; plus one whole mount of hydrocladia. Locality: south of Madagascar, 27°48’S 47°19’E; 875 m; 22 June 1961. Collected by: R.V. Africana II. Synthecium hians Millard, 1957: 204, fig. 9A-C. Holotype: SAM-—H107 (UCT FAL 214G). Several infertile stems; plus two whole mounts. Locality: False Bay, 34°12,4’S 18°43,5’E; 42 m; 10 September 1953. Collected by: University of Cape Town. TYPE SPECIMENS OF HYDROIDA IN THE SOUTH AFRICAN MUSEUM 143 Family Sertulariidae Salacia disjuncta Millard, 1964: 31, fig. 1OA-F. Holotype: SAM-H412 (UCT SCD 37K). One whole mount only (infertile). Locality: Agulhas Bank, 32°15,2’S 28°57,7’E; 50 m; 19 May 1958. Paratype: SAM-H1782 (UCT SCD 296H). One whole mount only (infertile); Agulhas Bank, 33°09’S 28°02’E; 84 m; 6 February 1962. Collected by: University of Cape Town. Sertularella agulhensis Millard, 1964: 35, fig. 12A. Holotype: SAM-—H413 (UCT TRA 151F). Several infertile stems; plus one whole mount. Locality: Agulhas Bank, 34°51’S 19°55’E; 22 m; 6 March 1958. Collected by: commercial trawler. Sertularella capensis Millard, 1957: 210, fig. 10H. Holotype: SAM-H108 (UCT FB 114A). A rich infertile colony; plus one whole mount. Locality: False Bay, 34°07,5’S 18°31’E; 27-28 m; 22 February 1947. Paratypes: SAM-H109 (UCT FB 115D). A fairly rich infertile colony; False Bay, 34°08’S 18°31,5’E; 27-28 m; 22 February 1947. SAM-H110 (UCT FAL 64L). A fairly rich infertile colony; plus one whole mount; False Bay, 34°17,3’S 18°48,7’E; 37-38 m; 29 July 1952. Collected by: University of Cape Town. Sertularella capensis delicata Millard, 1964: 38, fig. 12B—D. Since incorporated in Sertularella annulaventricosa Mulder & Trebilcock, 1915. Holotype: SAM-H414 (UCT NAD 22F). Several infertile stems; plus one whole mount. Locality: off Natal, 29°58’S 31°02’E; 49 m; 12 August 1958. Collected by: University of Cape Town. Sertularella congregata Millard, 1964: 39, fig. 13A—D. Holotype: SAM-H415 (UCT SCD 254Q). Fertile colony; plus one whole mount. Locality: Agulhas Bank, 33°07,3’S 28°01’E; 88 m; 16 July 1961. Collected by: University of Cape Town. Paratype: SAM-H18S. Infertile colony; plus one whole mount; Agulhas Bank, 33°09’S 28°03’E; 86 m; 28 December 1898. Collector unknown. 144 ANNALS OF THE SOUTH AFRICAN MUSEUM Sertularella dubia, var. magna Millard, 1958: 189, fig. 7A. = Sertularella dubia magna Millard. Holotype: SAM-HS54 (PF station 12028). Several infertile stems; plus one whole mount. Locality: off Natal, approx. 28°41’S 32°22’E; 62 m; 27 February 1901. Collected by: R.S. Pieter Faure. Sertularella falsa Millard, 1957: 211, figs 1OF, 11D. = Sertularella polyzonias falsa Millard. Holotype: SAM-H111 (UCT FB 119C). A fairly rich fertile colony; plus one whole mount. Locality: False Bay, 34°08,5’S 18°34,5’E; 27 m; 21 April 1947. Paratypes: SAM-—H112 (UCT FB 131H). Rich fertile colonies from several localities in False Bay and on various dates; plus one whole mount. SAM-H113 (UCT CP 333B). Fertile colony; plus one whole mount; Dalebrook, False Bay; littoral; 25 August 1949. Collected by: University of Cape Town. Sertularella gilchristi Millard, 1964: 44, fig. 12E, G-H. Holotype: SAM-H416 (UCT SCD 85J). A fairly rich male colony; plus one whole mount. Locality: Agulhas Bank, 33°03’S 27°55’E; 27 m; 17 July 1959. Collected by: University of Cape Town. Sertularella mediterranea var. asymmetrica Millard, 1958: 191, fig. 7B. = Sertularella mediterranea asymmetrica Millard. Holotype: SAM-H116 (UCT IN 49K). Several female stems; plus two whole mounts. Locality: Inhaca Island, Mozambique; littoral; 1954. Collected by: University of the Witwatersrand. Sertularella natalensis Millard, 1968: 271, fig. 4E-G. Schizoholotype: SAM—H2922 (TME 23M). One whole mount only. Locality: off Natal, 29°47,5’S 31°11,8’E; 64 m; 22 August 1929. Collected by: Th. Mortensen’s Java—South Africa Expedition. Sertularia linealis var. longa Millard, 1958: 197, fig. 8E. = Sertularia longa: Millard & Bouillon, 1974: 33, fig. 7D, J. Holotype: SAM-H117 (UCT IN 140E). Infertile colony on weed; plus one whole mount. Locality: East Shore Rocks, Inhaca Island, Mozambique; 20 July 1956. Collected by: University of the Witwatersrand. TYPE SPECIMENS OF HYDROIDA IN THE SOUTH AFRICAN MUSEUM 145 Symplectoscyphus marionensis Millard, 1971: 405, fig. 7. Holotype: SAM—H2923. One whole mount only of fertile colony. Locality: near Soft Plume River, Marion Island; littoral; 1965. Collected by: N. R. Fuller. Uniscyphus fragilis Millard, 19776: 122, fig. 7E-G. Holotype: SAM-H1982 (SM station 43). Two hydrothecae; plus one whole mount of three hydrothecae. Locality: off Natal, 28°45,5’S 32°24,5’E; 360-420 m; 29 May 1975. Collected by: R.V. Meiring Naude. This is the type species of the genus Uniscyphus Millard, 1977. Family Plumulariidae Subfamily Halopterinae Corhiza bellicosa Millard, 1962: 275, fig. 2A—E. Holotype: SAM-—H364 (UCT SCD 84S). Several infertile stems; plus one whole mount. Locality: Agulhas Bank, 33°03’S 27°55’E; 27 m; 17 July 1959. Paratype: SAM-H1947 (UCT SCD 153J). One infertile stem; plus one whole mount; Agulhas Bank, 34°03’S 25°59’E; 84 m; 25 November 1960. Collected by: University of Cape Town. Corhiza mortenseni Millard, 1968: 274, fig. SA—D. Schizoholotype: SAM-—H2914 (TME 24). Two whole mounts only. Locality: off Natal, 29°48,5’S 31°18’E; approx. 219 m; 22 August 1929. Collected by: Th. Mortensen’s Java—South Africa Expedition. Corhiza pannosa Millard, 1962: 278, fig. 3A—B, D-G. Holotype: SAM-—H362 (UCT SCD 5H). One infertile colony; plus one whole mount. Locality: Agulhas Bank, 34°15’S 25°05’E; 11 m; 19 April 1958. Collected by: University of Cape Town. Gattya tropicalis Millard & Bouillon, 1973: 79, fig. 1OA—D. Schizoholotype: SAM—H2924 (MT 2667). One whole mount only. Locality: Beau Vallon, Mahé Island, Seychelles; 1966. Collected by: J. Bouillon. 146 ANNALS OF THE SOUTH AFRICAN MUSEUM Halopteris gemellipara Millard, 1962: 282, fig. 4A—F. Holotype: SAM-—H308 (PF station 13810). An infertile colony; plus one whole mount. ; Locality: Agulhas Bank, approx. 33°54’S 26°51’E; 120 m; 10 September 1901. Collected by: R.S. Pieter Faure. Halopteris pseudoconstricta Millard, 1975: 355, fig. 114D-G. As Halopteris constricta: Millard, 1957: 227, fig. 14A. Holotype: SAM—H542 (UCT CP 740D). Numerous pinnate stems bearing female gonophores and a few infertile simple stems; plus one whole mount. Locality: Melkbosstrand, Table Bay; littoral; 17 November 1967. Collected by: University of Cape Town. Halopteris rostrata Millard, 1975: 357, fig. 114A-C. Holotype: SAM-—H543 (UCT NAD 89H). Eleven infertile stems and several damaged ones growing on skeleton of antipatharian; plus two whole mounts. Locality: off Natal, 29°11’S 32°02’E; 70 m; 30 July 1964. Collected by: University of Cape Town. Monostaechas faurei Millard, 1958: 204, fig. 11. Holotype: SAM-H58 (PF station 12028). Six male stems; plus two whole mounts. Locality: off Natal, approx. 28°41’S 32°22’E; 62 m; 27 February 1901. Collected by: R.S. Pieter Faure. Monostaechas natalensis Millard, 1958: 206, fig. 12. Holotype: SAM-H79 (PF station 12456). Many stems, male and female; plus two whole mounts. Locality: off Natal, approx. 30°32’S 30°38,5’E; 46 m; 22 March 1901. Paratypes: SAM-H48 (PF station 11803). One infertile stem; plus one whole mount; off Natal, approx. 29°00’S 31°49’E; 24 m; 8 February 1901. SAM-H76 (PF station 12392). Several infertile stems; plus one whole mount; off Natal, approx. 31°02,5’S 30°18’E; 46 m; 14 March 1901. Collected by: R.S. Pieter Faure. TYPE SPECIMENS OF HYDROIDA IN THE SOUTH AFRICAN MUSEUM 147 Subfamily Plumulariinae Plumularia antonbruuni Millard, 1967: 185, fig. 5. Holotype: SAM-H1647 (UCT ABD 14C). Three male stems; plus one whole mount. Locality: off Natal, 29°45’S 31°40’E; 440 m; 8 September 1964. Collected by: R.V. Anton Bruun. Plumularia irregularis Millard, 1958: 210, fig. 13A—C. = Kirchenpaueria irregularis: Millard, 1975: 370, fig. 118D-G (transferred to subfamily Kirchenpaueriinae). Holotype: SAM-H119 (UCT DBN 70Q). Three infertile stems; plus one whole mount. Locality: Salisbury Island, Durban Bay; littoral; 22 July 1950. Collected by: University of Cape Town. Plumularia mossambicae Millard, 1975: 393, fig. 123E-G. Holotype: SAM-H1866 (UCT PED 4V). One whole mount only (infertile stem). Locality: off Mozambique, 24°46’S 35°18’E; 110 m; 18 August 1964. Collected by: University of Cape Town. Plumularia pennycuikae Millard & Bouillon, 1973: 85, fig. 1ON-P. As Plumularia sp.: Pennycuik, 1959: 183, pl. 3 (fig. 7). Schizoholotype: SAM-—H2925 (MT 2668). One whole mount only. Locality: Silhouette Island, Seychelles; 1966. Collected by: J. Bouillon. Subfamily Aglaopheniinae Cladocarpus crepidatus Millard, 1975: 417, fig. 130A-C. Holotype: SAM-—H537 (UCT SST 6GG). One female stem; plus one whole mount. Locality: Agulhas Bank, 35°22’S 22°31’E; 200 m; 20 June 1972. Collected by: University of Cape Town. Cladocarpus natalensis Millard, 1977b: 127, fig. 10. Holotype: SAM-—H2861 (SM station 86). Five fertile stems; plus one whole mount. Locality: off Natal, 27°59,5’S 32°40,8’E; 550 m; 22 May 1976. Collected by: R.V. Meiring Naude. 148 ANNALS OF THE SOUTH AFRICAN MUSEUM Cladocarpus paries Millard, 1975: 483, fig. 143. Holotype: SAM-—H1914. One infertile stem; plus two whole mounts. Locality: Agulhas Bank, 33°25’S 27°29’E; 70 m; 23 January 1975. Collected by: South African Museum. Cladocarpus unicornus Millard, 1975: 429, fig. 133E-F. Holotype: SAM—H538 (UCT PED 4S). One infertile stem; plus one whole mount. Locality: off Mozambique, 24°46’S 35°18’E; 110 m; 18 August 1964. Collected by: University of Cape Town. Halicornaria africana Millard, 1958: 215, fig. 1SA—C. = Gymnangium africanum (Millard). Holotype: SAM-H120 (UCT AFR 1028B). One infertile colony; plus one whole mount. Locality: off Natal, 28°28’S 32°25,8’E; 27 m; 15 May 1948. Collected by: R.V. Africana IT. Halicornaria arcuata var. epizootica Millard, 1958: 218, fig. 15F. = Gymnangium arcuatum (Lamouroux, 1816), epizootic form. Holotype: SAM-H73 (PF station 12392). One whole mount only, infertile, epizootic on Thecocarpus formosus (Busk). Locality: off Natal, approx. 31°02,5’S 30°18’E; 46 m; 14 March 1901. Collected by: R.S. Pieter Faure. Halicornaria exserta Millard, 1962: 309, fig. 11A—E. = Gymnangium exsertum (Millard). Holotype: SAM-H169 (PF station 596). One fertile stem; plus one whole mount. Locality: Agulhas Bank, approx. 33°49’S 25°56’E; depth unknown; 11 November 1898. Collected by: R.S. Pieter Faure. Halicornaria exserta epizootica Millard, 1962: 309, fig. 11F—H. = Gymnangium exsertum (Millard), epizootic form. Holotype: SAM-—H320 (PF station 13915). Colony epizootic on Thecocarpus flexuosus umbellatus Millard; plus one whole mount. Locality: Agulhas Bank, approx. 34°05’S 26°34’E; 115 m; 23 September 1901. Collected by: R.S. Pieter Faure. TYPE SPECIMENS OF HYDROIDA IN THE SOUTH AFRICAN MUSEUM 149 Thecocarpus flexuosus umbellatus Millard, 1962: 316, fig. 12B, M. Holotype: SAM—H363 (UCT SCD 36M). Large infertile colony; plus one whole mount. Locality: Agulhas Bank, 32°15,2’S 28°57,7’E; 49,5 m; 19 May 1958. Collected by: University of Cape Town. Thecocarpus giardi var. solidus Millard, 1958: 222, fig. 16B—C. = Thecocarpus flexuosus solidus Millard. Holotype: SAM-H121 (UCT AFR 1028A). Large fertile colony; plus two whole mounts. Locality: off Natal, 28°28’S 32°25,8’E; 27 m; 15 May 1948. Collected by: R.V. Africana II. REFERENCES BaLe, W. M. 1888. On some new and rare Hydroida in the Australian Museum collection. Proc. Linn. Soc. N.S.W. (2) 3: 745-799. CouGuTrey, M. 1876. Critical notes on the New Zealand Hydroida. Trans. Proc. N.Z. Inst. 8: 298-302. Fraser, C. M. 1914. Some hydroids of the Vancouver Island region. Trans. R. Soc. Can. (3) 8: 99-216. Gow, C. & MILLARD, N. A. H. 1975. Two new species of campanularian hydroids from South Africa. Ann. S. Afr. Mus. 67: 1-6. GrAVIER, N. 1970. Etude des Hydraires epiphytes des Phanérogames marines de la région de Tulear (sud-oest de Madagascar). Rec! Trav. Stn mar. Endoume-Marseille 10: 111-161. Lamouroux, J. V. F. 1816. Histoire des polypiers coralligénes flexibles, vulgairement nommés zoophytes. Caen: Poisson. MILLARD, N. A. H. 1955. New species of Hydrozoa from South Africa. Ann. S. Afr. Mus. 41: 215-222. MILLARD, N. A. H. 1957. The Hydrozoa of False Bay, South Africa. Ann. S. Afr. Mus. 43: 173-243. MILLARD, N. A. H. 1958. Hydrozoa from the coasts of Natal and Portuguese East Africa. Part I. Calyptoblastea. Ann. S. Afr. Mus. 44: 165-226. MILLARD, N. A. H. 1959a. Hydrozoa from the coasts of Natal and Portuguese East Africa. Part II. Gymnoblastea. Ann. S. Afr. Mus. 44: 297-313. MILLARD, N. A. H. 19596. Hydrozoa from ships’ hulls and experimental plates in Cape Town docks. Ann. S. Afr. Mus. 45: 239-256. MILLARD, N. A. H. 1962. The Hydrozoa of the south and west coasts of South Africa. Part I. The Plumulariidae. Ann. S. Afr. Mus. 46: 261-319. MILLARD, N. A. H. 1964. The Hydrozoa of the south and west coasts of South Africa. Part II. The Lafoeidae, Syntheciidae and Sertulariidae. Ann. S. Afr. Mus. 48: 1-56. MILLARD, N. A. H. 1966. The Hydrozoa of the south and west coasts of South Africa. Part III. The Gymnoblastea and small families of Calyptoblastea. Ann. S. Afr. Mus. 48: 427-487. MILLARD, N. A. H. 1967. Hydroids from the south-west Indian Ocean. Ann. S. Afr. Mus. 50: 169-194, MILLarD, N. A. H. 1968. South African hydroids from Dr Th. Mortensen’s Java—South Africa Expedition, 1929-1930. Vidensk. Meddr dansk naturh. Foren. 131: 251-288. MILLARD, N. A. H. 1971. Hydrozoa. In: ZINDEREN BAKKER, E. M. VAN, WINTERBOTTOM, J. M. & Dyer, R. A., eds. Marion and Prince Edward Islands: 396-408. Cape Town: A. A. Balkema. MILLARD, N. A. H. 1973. Auto-epizoism in South African hydroids. Proc. Second Internat. Symp. Cnidaria; Publs Seto mar. biol. Lab. 20: 23-34. 150 ANNALS OF THE SOUTH AFRICAN MUSEUM MILLARD, N. A. H. 1975. Monograph on the Hydroida of southern Africa. Ann. S. Afr. Mus. 68: 1-513. MILLARD, N. A. H. 1977a. Hydroids from the Kerguelen and Crozet shelves, collected by the cruise MD.03 of the Marion-Dufresne. Ann. S. Afr. Mus. 73: 1-47. MILLARD, N. A. H. 19776. The South African Museum’s Meiring Naude cruises. Part 3. Hydroida. Ann. S. Afr. Mus. 73: 105-131. Miiiarpb, N. A. H. & BourLton, J. 1973. Hydroids from the Seychelles (Coelenterata). Annls Mus. r. Afr, cent. Sér. 8 vo (Sci. zool.) 206: 1-106. MILLARD, N. A. H. & BourILLon, J. 1974. A collection of hydroids from Mocgambique, East Africa. Ann. S. Afr. Mus. 65: 1-40. MILLARD, N. A. H. & BoumLton, J. 1975. Additional hydroids from the Seychelles. Ann. S. Afr. Mus. 69: 1-15. Mu.per, J. E. & Trepitcock, R. E. 1915. Victorian Hydroida. With description of new species. Part V. Geelong Nat. 1915: 51-S9. PENNYCUIK, P. R. 1959. Faunistic records from Queensland. Part VY. Marine and brackish water hydroids. Pap. Dep. Zool. Univ. Qd 1: 141-210. Rees, W. J. 1962. Hydroids of the family Cytaeidae L. Agassiz, 1862. Bull. Br. Mus. nat. Hist. (Zool.) 8: 381-400. TureL, H. 1962. Clavopsella quadranularia nov. spec. (Clavopsellidae nov. fam.), ein neuer Hydroidpolyp aus der Ostsee und seine phylogenetische Bedeutung. Z. Morph. Okol. Tiere 51: 227-260. 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, 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. N. A. H. MILLARD TYPE SPECIMENS OF HYDROIDA (COELENTERATA) IN THE SOUTH AFRICAN MUSEUM j - a 52 . A ) MUS. COMP. ZOOL. LIBRARY JUL 2 3 407 HARVARD OF THE SOUTH vi. RN 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 ) 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, 19696; Jones 1971)’ “As described (Haughton & Broom 1927)...’ ‘As described (Haughton ef a/. 1927)...’ Note: no comma separating name and year Pagination indicated by colon, not p. names of joint authors connected by ampersand re 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, 6, 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) BuLLouGuH, 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. Fiscuer, 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 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. TurELe, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: 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 77 Band May 1979 Mei Part 9 Deel THE UNIONACEA (MOLLUSCA, LAMELLIBRANCHIATA) OF SOUTH-CENTRAL AFRICA By C.<. APPLETON 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, 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 Copyright enquiries to the South African Museum Kopieregnavrae na die Suid-Afrikaanse Museum ISBN 0 908407 71 8 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 UNIONACEA (MOLLUSCA, LAMELLIBRANCHIATA) OF SOUTH-CENTRAL AFRICA By C. C. APPLETON* Bilharzia Field Research Unit, South African Medical Research Council, Nelspruit (With 8 figures and | table) [MS. accepted 11 December 1978] ABSTRACT Twelve species of the Unionacea (Mollusca, Lamellibranchiata), five Unionidae and six Mutelidae and one Etheriidae, are recorded from the south-central region of Africa. Their distribution within this region, which is taken to embrace the catchments of the Kunene, Cuvelai, Okavango, Linyanti and Zambezi River systems, is updated. Attention is drawn to the paucity of ecological data relevant to the freshwater bivalves of Africa and the detailed locality notes made by several collectors are therefore included. CONTENTS PAGE Introdiction a=” 2] 2 Se os: 151 Systematic discussion ae eee aso General discussion . . . .-. 168 Acknowledgements . . . . . 172 Refererices 9% | ae seep eee Bc 2 INTRODUCTION Knowledge of the Unionacea of the southern central African region embracing the Kunene, Cuvelai, Okavango and Zambezi River systems (i.e. between latitudes 9° and 21°S) has hitherto been limited, particularly with regard to the Kunene and Okavango. During the past decade, however, a considerable amount of material has been collected from these drainages and through the co-operation of a number of individuals and institutions the author has been able to examine examples of eleven of the twelve recorded unionacean species, five Unionidae, five Mutelidae and one Etheriidae, per- mitting an updating of knowledge on the composition and distribution of this bivalve fauna. Unfortunately no material is available from the middle and lower Zambezi, east of Lake Kariba. However, the bivalve fauna of Lake Macllwaine on the Hunyani River, a tributary to the middle Zambezi, has been discussed by Marshall (1975) and the malacofauna of Lake Malawi has been dealt with by Crowley et a/. (1964) and Mandahl-Barth (1972). In addition, the bivalves of the adjacent Bangweulu—Luapala basin, the Congo River system, and Mozam- bique have been variously discussed by Mandahl-Barth (1968), Pilsbry & * Present address: School of Environmental & Life Sciences, Murdoch University, Murdoch, Western Australia 6153. 151 Ann. S. Afr. Mus. 77 (9), 1979: 151-174, 8 figs, 1 table. 152 ANNALS OF THE SOUTH AFRICAN MUSEUM Bequaert (1927), Azevedo et al. (1961), and Braga (1952). In the present contri- bution the systematic arrangement of Connolly (1939) has been followed, though changes in nomenclature made by Haas (1969), Mandahl-Barth (1968, 1972) and Heard & Vail (1976) have been incorporated. Figure 1 shows the watercourses referred to in the text. These comprise both large, perennial rivers such as the Kunene and Okavango with volumes that may exceed 6 000 m/s during high flood which can drop to less than 15 m3/s in dry years (Stengel 1963), as well as largely seasonal floodplain or deltaic environments such as the Okavango delta, Linyanti swamp, eastern Caprivi floodplain and the Etosha Pan. Where measurements of shell height (altitude) are given, maximum height is used (except where indicated) in preference to umbonal height because in many cases the umbones were eroded to such an extent that considerable inaccuracies would have been introduced. The following abbreviations are used: SAM — South African Museum, Cape Town NM Natal Museum, Pietermaritzburg ™ Transvaal Museum, Pretoria MMK McGregor Memorial Museum, Kimberley SMN — State Museum, Windhoek Where no abbreviations are given, the material has been lodged in the collection of the South African Museum, Cape Town. Lake Malawi 2 vate FR. a | - - ; | ss | a = ii. es od ae " » if — Vee 7 (ae —_ _ : —_ i me : = 7 : a — _ a a - q hi > i : : : : 1 § ¥ ime | yy s ‘ a - i es in | ] { ; . i : 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, 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. C. C. APPLETON THE UNIONACEA (MOLLUSCA, LAMELLIBRANCHIATA) OF SOUTH-CENTRAL AFRICA S- Hy - ajcetow ste VOLUME 77 PART 10 JUNE 1979 ISSN 0303-2515 MUS. COMP. ZOOL, LIBRARY AUG 15 1979 HARVARD UNIVERSITY | ANNALS OF THE SOUTH AFRICAN MUSEUM 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. 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, 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. gen. 74: 627-634. Kon, 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, i sa aes and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): Turee, J. 1910. Mollusca: B. Seiatemainon: 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 77 Band June 1979 Junie Part 10 £Deel UPPERMOST ALBIAN (STOLICZKAIA DISPAR ZONE) AMMONITES FROM THE ANGOLAN LITTORAL By M. R. COOPER & W. J. KENNEDY 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 gelang van die 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, 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 Copyright enquiries to the South African Museum Kopieregnavrae aan die Suid-Afrikaanse Museum ISBN 0 908407 76 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 UPPERMOST ALBIAN (STOLICZKAIA DISPAR ZONE) AMMONITES FROM THE ANGOLAN LITTORAL By M. R. CooPER Queen Victoria Museum, Salisbury & W. J. KENNEDY Geological Collections, University Museum, Oxford (With 82 figures) LMS. accepted 11 December 1978] ABSTRACT Rich, well-preserved collections of late Albian (Stoliczkaia dispar Zone) ammonites from the Angolan littoral provide the basis for discussion of the ontogenetic and intraspecific variation within S. (Stoliczkaia) tenuis Renz, Tetragonites jurinianus (Pictet), Eogaudryceras italicum Wiedmann & Dieni, Desmoceras latidorsatum perinflatum subsp. nov., Phylloceras (Hypophylloceras) seresitense Pervinquiére, Anisoceras haasi sp. nov., and Mortoniceras (Durnovarites) collignoni sp. noy. Other important elements present include Jdiohamites dorsetensis Spath, I. pygmaeus sp. nov., I. cf. elegantulus Spath, Anisoceras armatum (J. Sowerby), A. perarmatum Pictet & Campiche, A. phillipsi sp. nov., Mortoniceras (Durno- varites) perinflatum (Spath), M. (D.) subquadratum Spath, M. (Angolaites) simplex (Choffat), M. (A.) gregoryi (Spath), Cantabrigites? curvatum Renz, Hysteroceras? cf. ootaturense (Stoliczka) and Borissiakoceras sp. nov. ? aff. reymenti (Brunnschweiler). The faunas form the basis for discussion of the subdivisions of the Stoliczkaia dispar Zone, and a return to the simple division into subzones of S. (Faraudiella) blancheti below and M. (Durnovarites) perinflatum above is proposed. CONTENTS PAGE Introduction . : : : 2 : : ; «. Lib Systematic palaeontology . : P : : ee i 8 Family Phylloceratidae p , : : Re ae) WS i Genus Phylloceras . : F : ‘ eo TT Family Tetragonitidae . : , : : St Genus Tetragonites : : : : . 181 Family Gaudryceratidae . : ; ‘ . 489 Genus Eogaudryceras . ‘ : : ; 189 Family Ancyloceratidae : ‘ A : Oo Genus Anisoceras . : A , ; ‘ 196 Genus Idiohamites . : ‘ ; ; ee: Family Hamitidae . : 4 . 5 : at 26 Genus Hamites . P : , ‘ » tees Family Turrilitidae : : : : ; < eee Genus Mariella. ; ; : ; . 228 Family Desmoceratidae . ‘ ‘ : a 28 Genus Puzosia ; : : , ; ee Genus Desmoceras ~ : : : 7) ma 175 Ann. S. Afr. Mus. 77 (10), 1979: 175-308, 82 figs. 176 ANNALS OF THE SOUTH AFRICAN MUSEUM Systematic palaeontology (cont.) PAGE Family Lyelliceratidae . : : = out el Genus Stoliczkaia . . ; : : . 243 Family Brancoceratidae ‘ : : ; ~ 265 Genus Hysteroceras? . : : : » 265 Genus Mortoniceras . ‘ j : . 269 Subgenus M. (Durnovarites) : F . 269 Subgenus M. (Angolaites) . : : - 285 Genus Cantabrigites? . : : : . 290 Genus Drakeoceras : ‘ ; : e293 Family Binneyitidae . : ‘ ‘ : . 296 Genus Borissiakoceras . : : : » . 296 Age of the fauna . ; P : : , : z - 298 Summary : : F ; : : : , = » S01 Acknowledgements : : : : ; : . 303 References. : : ‘ : : : : = 303 INTRODUCTION Although Albian ammonites have long been known from Angola (Szajnocha 1884; Meunier 1887; Choffat in Choffat & De Loriol 1888), Paul Choffat was the first to record uppermost Albian ammonites when he described Ammonites dispar Choffat (non d’Orbigny) (= Stoliczkaia tenuis Renz) from the environs of Catumbela. Subsequently, many authors have reported on the presence of strata with Ammonites dispar (Mouta & Borges 1926, 1928; Mouta 1937, 1954, 1956; Mouta & O’Donnell 1933) and the presence of uppermost Albian strata in Angola is now well established. In contrast, the faunal constituents of these rocks are poorly known. The collections which form the basis of this paper are from Porto Amboim, Cabo Ledo and Praia-Egito. The material from Porto Amboim comes from the lime kiln at the south side of the boat landing, and was collected mostly by C. W. Washburn on 22 July 1914, although some material from this locality, in the South African Museum, was described by Haughton (1924, 1925). Elements of this fauna were also described subsequently by Haas (1942), whose localities R28, R30 and 3041 have yielded an identical fauna in like preservation, with recrystallized test preserved. The fauna from Egito was collected by the senior author and comes from the coastal exposures at Praia-Egito where thinly-bedded limestone-shale alternations have yielded a rich fauna, mostly preserved as composite internal moulds. Faunas collected by the senior author from the Quissama Ridge, the promontory on which stands the Farol de Cabo Ledo, were obtained from road gravels and may represent more than one faunal horizon. These faunas include a number of new species, and also populations, of S. (Stoliczkaia), Mortoniceras (Durnovarites), E. (Eogaudryceras), Phylloceras (Hypophylloceras), D. (Desmoceras), Anisoceras and Tetragonites which provide unique information on the intraspecific variation, development and morphology of these ammonites. Furthermore, the composition of the faunas provides an opportunity for discussion of the stratigraphic and geographic distribution of the late Albian Stoliczkaia dispar Zone fauna. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 177 SYSTEMATIC PALAEONTOLOGY The following abbreviations are used to indicate the source of material: BM _ British Museum (Natural History), London SAM_ South African Museum, Cape Town USNMNH USS. National Museum of Natural History, Washington D.C. Measurement abbreviations are as follows: D, diameter; H, whorl! height; W, whorl width (intercostal unless otherwise specified); U, diameter of umbilicus between umbilical seams; “/H, whorl width/height ratio. All measurements are given in millimetres and dimensions, as a percentage of the diameter, follow in parentheses. Class CEPHALOPODA Cuvier, 1797 Subclass AMMONOIDEA Zittel, 1884 Order PHYLLOCERATIDA Arkell, 1950 Superfamily PHYLLOCERATACEAE Zittel, 1884 Family Phylloceratidae Zittel, 1884 Subfamily Phylloceratinae Zittel, 1884 Genus Phylloceras Suess, 1865 Subgenus Hypophylloceras Salfeld, 1924 Type species Phylloceras onoense Stanton, 1895 Phylloceras (Hypophylloceras) seresitense Pervinquiére, 1907 Figs 1-2, 31 Ammonites velledae Pictet & Campiche (non Michelin), 1860: 268, pl. 36 (fig. 8). Stoliczka, 1865: 116, pl. 59 (figs 1-3). Phylloceras velledae (Michelin) Kossmat 1895: 12, pl. 1 (fig. 3). Boule, Lemoine & Thévenin, 1906: 7, pl. 1 (figs 6, 11), fig. 2. Crick, 1907: 166, pl. 10 (figs 10-11). Bése, 1923: 119, pl. 7 (figs 15-17). Spath, 1925b: 180. Besairie, 1936: 164, pl. 16 (fig. 1). Venzo, 1936: 66, pl. 5 (fig. 4). Haas, 1942: 146, pl. 27 (fig. 1), pl. 44 (fig. 1), fig. 19. Matsumoto, 1942: 676. Almela & Revilla, 1957: 17, pl. 3 (fig. 2). Anderson, 1958: 180, pl. 16 (fig. 4). Phylloceras velledae var. seresitense Pervinquiére, 1907: 52; 1910: 9, pl. 1 (figs 1-3), fig. 2. Phylloceras tanit Pervinquiére, 1907: 53, pl. 3 (figs 3-9), fig. 5. Phylloceras angolaense Haughton, 1924: 85, pl. 1 (figs 1-2); 1925: 267, pl. 12 (figs 1-2). Phylloceras seresitense Pervinquiére, Spath, 1923: 18, pl. 1 (fig. 2), pl. 2 (fig. 1). Collignon (in Besairie) 1936: 190, pl. 21 (figs 1-2). Fabre, 1940: 211, pl. 5 (fig. 1). Phylloceras boulei Collignon, 1928: 144, pl. 15 (fig. 5), fig. 1. Hyporbulites seresitensis (Pervinquiére) Breistroffer, 1947: 82. Phylloceras (Hyporbulites) seresitense Pervinquiére, Collignon 1950: 66; 1963: 4, pl. 241 (fig. 1038), pl. 242 (fig. 1041). 178 ANNALS OF THE SOUTH AFRICAN MUSEUM Hyporbulites seresitensis var. raynaudiensis Collignon, 1956: 16, pl. 4 (fig. 1). Neophylloceras seresitense (Pervinquiére) Matsumoto, 1959: 55, pl. 12 (figs 4-5), fig. 3. ? Phylloceras (Euphylloceras) vohipalense Collignon, 1962: 1, pl. 215 (fig. 940). ? Phylloceras sp. (Ph. velledae ?) da Silva, 1962: 26, pl. 15 (figs 1-3). Hypophylloceras seresitense seresitense (Pervinquiére) Wiedmann, 1962a: 142, pl. 8 (figs 1-2) fig. 8; 19625: 249, pl. 16 (fig. 1). Hypophylloceras seresitense tanit (Pervinquiére) Wiedmann, 1962a: 142; 1962b: 250, fig. 2. Phylloceras (Hypophylloceras) seresitense seresitense Pervinquiére, Wiedmann, 1964: 221, pl. 15 (fig. 4), pl. 21 (fig. 1), fig. 52. Wiedmann & Dieni, 1968: 26. Kennedy & Klinger, 1977a: 364, pl. 4 (fig. 6), pl. 6 (fig. 4), pl. 7 (fig. 4), pl. 9. Renz, 1968: 17, pl. 1 (fig. 1). Phylloceras (Hypophylloceras) seresitense tanit Pervinqui¢re, Wiedmann, 1964: 226, pl. 21 (figs 2-3), fig. 54. Wiedmann & Dieni, 1968: 26, pl. 1 (fig. 6), pl. 3 (figs 1-2). Phylloceras (Hypophylloceras) seresitense boulei Collignon, Wiedmann, 1964: 224, pl. 20 (figs 2-3), fig. 53. Renz, 1968: 18, pl. 1 (fig. 2), figs 6a, 7c. Forster, 1975: 140, pl. 1 (fig. 1), fig. 27. ? Phylloceras (Hypophylloceras) seresitense vohipalense Collignon, Forster, 1975: 139, fig. 26. ? Phylloceras serum var. perlobata Zwierzycki (non Sayn), 1913: 323, figs a-c. ? Phylloceras ex aff. ramosi Meek, Collignon, 1928: 1, pl. 1 (figs 2-4). ? Phylloceras aff. tanit Pervinquiére, Matsumoto, 1942: 674, fig. 2. Phylloceras cf. seresitense Pervinquiére, Wright & Wright, 1951: 12. ? Phylloceras cfr. semistriatum Choffat (non d’Orbigny), 1903: 17, pl. 1 (fig. 10). Material Ten specimens, SAM-6527, USNMNH 236897-236904, 237013, all with recrystallized shell preserved, and all from Porto Amboim. ~ Description The coiling is very involute, with a narrow, crater-like umbilicus (6-7% of the diameter). The whorl section is strongly compressed, elliptical (W/~ = 0,55-0,67). The flanks are slightly convex, with maximum width just below midflank, converging to a narrow, evenly rounded venter. Ornament comprises fine, flexuous lirae, very faint or completely effaced on the inner half of the flanks (faint growth striae suggest they were prorsiradiate there), recurving at about midflank so as to pass straight or slightly backwards across the venter. On USNMNH 236897 there are about 128 lirae per half-whorl. None of the present specimens shows the sutures. Measurements No. D H W W/H U USNMNH 236897 37 21(57) 13(35) 0,62 2,6(7) » 26,5 15,5(58) +8,5(32) 0,55 USNMNH 236898 a2 30,5(59) 17,5(34) 0,57 3(6) » 35 19(54) 11(31) 0,58 it USNMNH 236900 15 8(53) 5(33) 0,63 1,1(7) » 11 5,5(50) 3,7(34) 0,67 ? USNMNH 236901 | OF, 10(59) 6,5(38) 0,65 1,0(6) USNMNH 236902 || 12(57) TMD 0,64 ? UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 179 Fig. 1. Phylloceras (Hypophylloceras) seresitense Pervinquitre. A-B. USNMNH 236899. C-E. USNMNH 236904. F-H. USNMNH 236897. I-K. USNMNH 236903. L-N. USNMNH 236898. O-P. USNMNH 236902. Q-S. USNMNH 236901. T-V. USNMNH 236900. x1. 180 ANNALS OF THE SOUTH AFRICAN MUSEUM Intraspecific variation Wiedmann (1964: 221) recognized three subspecies within P. (H.) seresitense: P. (H.) seresitense seresitense—whorl section moderately compressed, whorl width to height ratio up to 0,65. Umbilicus very narrow. P. (A.) seresitense boulei Collignon—whorl section very compressed (W/H = 0,50-0,57), with very narrow umbilicus. P. (H.) seresitense tanit—whorl section compressed, but with moderately open umbilicus. The most noteworthy variation in the population before the writers is in the degree of inflation of the whorls. As can be seen from Figure 2 the variation in this respect (¥/H = 0,55-0,67) includes individuals of all of Wiedmann’s (1964) subspecies and, since they are all broadly contemporaneous (ranging from Aptian to Cenomanian), appear to have little biological significance. The authors find, therefore, the application of the subspecies boulei and tanit inappropriate to the present collection. Discussion The holotype of Phylloceras angolaense Haughton is in the South African Museum, SAM-6527. The specimen is entirely septate and is replaced by crystalline calcite. The shell is strongly compressed and very involute, with a very narrow, deep umbilicus. The flanks are slightly convex, almost flat, and converge towards the narrow, evenly rounded venter (Fig. 31). Maximum width is about one-quarter of the way up the flanks. Ribbing is very faint on the inner half of the flanks, although growth striae suggest it was prorsiradiate here. On the outer half of the flanks, the ribs recurve so as to become almost radial. The ribbing is very fine, with fourteen ribs in a distance of 10 mm along the venter on the adoral quarter of the outer whorl. There are no features by which P. angolaense may be satisfactorily dis- tinguished from P. seresitense and consequently the authors follow Wiedmann (1962a, 19625) in regarding it as a junior subjective synonym of Pervinquiére’s species. Phylloceras velledae (Michelin) (Wiedmann 1964: 209, pl. 11 (fig. 1), pl. 13 (fig. 4), pl. 21 (fig. 4), fig. 49) typically differs from the present species in having more convex flanks, with much coarser, more flexuous lirae, and a suture which shows diphyllic saddles in immaturity rather than tetraphyllic saddles as in P. (H.) seresitense. Hypophylloceras yeharai Nakai & Matsumoto (1968: 4, pl. 1 (figs 1-3), pl. 3 (fig. 1)) is based upon crushed material said to differ from P. (H.) velledae in its more compressed whorls (? enhanced by crushing) and less numerous and broader lirae. Kennedy & Klinger (1977a) have recently demonstrated the wide range of variation within contemporaneous populations of P. (H.) velledae and UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 181 WIDTH YAEIGHT oO 0 30 60 90 120 SAE) ————— Fig. 2. Plot of inflation (whorl width/height ratio) versus diameter for Phylloceras seresitense Pervinquiére, showing the unsatisfactory separation of subspecies. Circles = present Angolan material; dots = P. seresitense tanit Pervinquiére; square = holotype of P. angolaense Haughton; open triangles = P. seresitense seresitense Pervinquiére; black triangles = P. seresitense boulei Collignon. (Data after Stoliczka 1865, Spath 1923, Wiedmann 1962, Collignon 1963, Wiedmann & Dieni 1968, Renz 1968, Forster 1975, and Kennedy & Klinger 1977.) figured examples (1977a: pls 10-11) as coarsely ribbed as the Japanese material, Suggesting that P. (H.) yeharai may be within the limits of variation of P. (A..) velledae. Occurrence Phylloceras (Hypophylloceras) seresitense ranges from Upper Aptian to Middle Cenomanian, and is known from southern France, Switzerland, southern England, Spain, Tunisia, Algeria, Sardinia, the Balearics, southern India, Japan, Alaska, California, Mexico, Angola, Zululand, possibly Tanzania, and Madagascar. Order LYTOCERATIDA Hyatt, 1889 Superfamily LYTOCERATACEAE Neumayr, 1875 Family Tetragonitidae Hyatt, 1900 Genus Tetragonites Kossmat, 1895 Types species Ammonites timotheanus Pictet, 1848 Discussion Wiedmann (1973) has provided the most recent discussion of the Albian to Cenomanian members of this genus, recognizing three species groups. The group of 7. rectangularis possesses persistent constrictions and a straight umbilical suture, and includes 7. rectangularis Wiedmann, T. subtimotheanus 182 ANNALS OF THE SOUTH AFRICAN MUSEUM Wiedmann, 7. kitchini (Krenkel), T. subbeticus Wiedmann, T. marrei Thomel and 7. blaisoni Collignon. In the group of 7. timotheanus (Pictet), constrictions are present only at an early growth stage, whilst the umbilical suture is retracted. To this group Wiedmann (1973) assigned 7. timotheanus (Pictet), T. balmensis Breistroffer, and 7. nautiloides (Pictet). Forms lacking constrictions at all growth stages are referred to the group of 7. jurinianus (Pictet). Tetragonites (Tetragonites) collignoni Breistroffer, 1940 Ammonites timotheanus Stoliczka (non Mayor), 1865: 146, pl. 73 (figs 3-4, 6). Tetragonites collignoni Breistroffer, 1940: 110. Murphy, 1967a: 66, pl. 5 (figs 2-5), fig. 36. Forster, 1975: 147, pl. 1 (fig. 5) (with synonymy). Tetragonites subtimotheanus Wiedmann, 1962a: 131; 1973: 592, pl. 1 (fig. 5), pl. 2 (fig. 2), pl. 3 (figs 1-5), pl. 7 (fig. 8), fig. 2 (with synonymy). Tetragonites blaisoni Collignon, 1964: 31, pl. 324 (fig. 1448). Wiedmann, 1973: 601, pl. 1 (fig. 4), pl. 6 (figs 5-7), fig. 7 (with synonymy). Material One specimen, USNMNH 236916, preserved as an internal mould, from Porto Amboim. Description Shell involute, somewhat inflated, with a depressed, trapezoidal whorl section (W/H = 1,21). Umbilicus narrow (22% of the diameter), deep, with steep, almost vertical umbilical walls. Umbilical shoulder evenly rounded, with flattish flanks converging towards a broad, slightly convex venter which rounds somewhat in maturity. There are two distinct constrictions in a distance equal to the whorl height on the last portion of the outer whorl (representing part of the body chamber), although the number of constrictions per whorl is unknown. The constrictions are markedly prorsiradiate across the inner flanks, recurving strongly in the region of the ventrolateral angulations to cross the venter with a prominent concave sinus. Measurements No. D H W W/H U USNMNH 236916 44 21,5(49) +26(59) 1,21 ++9,5(22) Discussion Problems concerning the specific identification of constricted tetragonitids become very clear from the recent literature concerning this group (Wiedmann 1962a, 1962b, 1973; Collignon 1963, 1964; Murphy 1967a, 1967b; Wiedmann & Dieni 1968; McLearn 1972; Forster 1975; Kennedy & Klinger 19776). Forster (1975) has suggested that 7. subtimotheanus and T. blaisoni are junior subjective synonyms of T. collignoni. Wiedmann (1973) rejected the latter name because it UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 183 was based upon a pyritic nucleus only 14 mm in diameter (although he retains T. kitchini and T. subbeticus as valid species even though the types are only 18 and 16 mm in diameter respectively), considering it a nomen dubium. Because the authors are at present unhappy with Wiedmann’s (1973) fine delimitation of T. subtimotheanus Wiedmann, T. rectangularis Wiedmann, T. blaisoni Collignon, and 7. kitchini (Krenkel), all of which are broadly contemporaneous, they have some sympathy with Forster’s (1975) view in assigning the present material to T. collignoni. Pot) Fig. 3. A-D. Mortoniceras (Angolaites) simplex (Choffat). A. SAM-PCA4613. B. SAM- PCA4774. C. SAM-PCA4590. D. SAM-PCA4609. E-F. Drakeoceras cf. dellense Young. E. SAM-PCA4662. F. SAM-—PCA4800. G. Mortoniceras (Durnovarites) perinflatum (Spath), SAM-PCA4587. H. Mariella (Mariella) cf. oehlerti (Pervinquiére), SAM-—PCA4798. I. Phylloceras (Hypophylloceras) seresitense (Pervinquiére), the holotype of P. angolaense Haughton, SAM-6527. x1. 184 ANNALS OF THE SOUTH AFRICAN MUSEUM Occurrence Tetragonites collignoni ranges from the Lower Albian to the Middle Cenomanian, and is known from Madagascar, southern India, Zululand, Alaska, British Columbia, and Oregon. Tetragonites (Tetragonites) kitchini (Krenkel, 1910) Fig. 4 ? Ammonites timotheanus Whiteaves (non Mayor), 1876: 41, pl. 3 (fig. 2). Desmoceras (Puzosia; Latidorsella ?) kitchini Krenkel, 1910: 226, pl. 22 (fig. 8). Tetragonites kitchini (Krenkel), Wiedmann 1962a: 171. Murphy, 1967a: 33, pl. 2 (figs 11-14), figs 15-16. Wiedmann, 1973: 599, pl. 1 (fig. 7), pl. 6 (figs 1, ?2-4), figs 5, 26. Tetragonites hulenensis Murphy, 1967a: 54, pl. 6 (figs 16-19), pl. 7 (figs 3, 6-8, 10), figs 28-30. Murphy, 19675: pl. 4 (figs 8-9). ? Tetragonites aff. kitchini (Krenkel), McLearn 1972: 26, pl. 4 (figs 4-5). Material A single specimen, SAM-PCA3125, from low in the coastal cliffs immediately north of the estuary at Praia-Egito (low dispar Zone). Description The shell is moderately involute (umbilicus 31,5°% of the diameter) and partially retains recrystallized shell. The whorl section is subtrapezoidal, almost subquadrate, as wide as high (W/n = 1,00), with slightly converging flanks and a flattish, slightly convex venter. The umbilicus is rather narrow and deep, with steep umbilical walls and subrounded umbilical shoulders. The ventrolateral shoulders are evenly rounded. Maximum width is at the umbilical shoulder. Fig. 4. Tetragonites kitchini (Krenkel). Lateral and ventral views of SAM-PCA3125, partially retaining recrystallized test. x1. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 185 There would appear to have been about seven slightly flexuous (prorsi- radiate concave in the terminology of Murphy (1967a)), strongly prorsiradiate constrictions on the outer whorl, which cross the venter with a concave-adoral sinus. The suture line was not observed. Measurements No. D H W W/H U SAM-PCA3125 54 25(46) 25(46) 1,00 17(31,5 %) Discussion Tetragonites kitchini differs from all other contemporaneous species of constricted Tetragonites in having less strongly inflated whorls with an almost quadrate whorl section. However, there appear to be intermediate forms to T. subtimotheanus Wiedmann and T. rectangularis Wiedmann, and the population structures of these species require investigation to fully validate their specific separation. Occurrence Tetragonites kitchini is known from the upper Lower Albian of Tanzania, California, Alaska, and perhaps British Columbia, and from the Middle or Upper Albian of Madagascar, and the uppermost Albian of Angola. Tetragonites (Tetragonites) jurinianus (Pictet, 1847) Figs 5-6 Ammonites jurinianus Pictet (in Pictet & Roux), 1847: 297, pl. 3 (fig. 3). D’Orbigny, 1850: 124. Ammonites timotheanus Pictet & Campiche (zon Mayor), 1860: 289 (pars). Lytoceras (Tetragonites) epigonum Boule, Lemoine & Thévenin (non Kossmat), 1906: 186, pl. 3 (fig. 1). ? Lytoceras (Tetragonites) timotheanum Pervinquiére (non Mayor), 1907: 74 (pars.), pl. 3 (fig. 24 only). Lytoceras (Tetragonites) jurinianum (Pictet) Jacob, 1908: 19, pl. 1 (fig. 12). Latidorsella latidorsata Jacob (non Michelin), 1908: 35, pl. 5 (fig. 1 only). Lytoceras (Tetragonites) zacatecanus Bose, 1923: 127, pl. 9 (figs 11-17). Tetragonites jurinianus (Pictet) Spath, 1923: 26. Roman, 1938: 43. Breistroffer, 1940: 112. Wiedmann, 1962a: 176, pl. 14 (fig. 2), figs 37-38. Almela & Revilla, 1957: 20, pl. 4 (fig. 3). Murphy, 1967a: 23, pl. 2 (figs 1-4); figs 10-11. Renz, 1968: 19, pl. 1 (figs 6-7), figs 6d, 7Ja—b. Wiedmann & Dieni, 1968: 48, pl. 4 (fig. 4), pl. 5 (fig. 4). Wiedmann, 1973: 608, pl. 8 (figs 3-4). ? Tetragonites brazoensis Bose, 1927: 203, pl. 1 (figs 2-7). Tetragonites jurinianus var. angolana Haas, 1942: 170, pl. 44 (fig. 3), pl. 45 (fig. 1). Tetragonites timotheanus Matsumoto (non Mayor), 1959: 78, fig. 16. Tetragonites jurinianus angolanus Haas, Wiedmann, 1973: 609. Material Ten specimens, USNMNH 236915, 236917-21, 236977-79, all with recrystallized shell preserved, and all from Porto Amboim. 186 ANNALS OF THE SOUTH AFRICAN MUSEUM Description Shell inflated, involute, with a slightly depressed, trapezoidal whorl section in juveniles (“/q = 1,00-1,29), the venter becoming rounded in maturity. Umbilicus narrow (16-30% of the diameter), deep, with almost vertical walls and evenly rounded umbilical shoulders. The flanks are flattened, with maximum width close to the umbilical shoulder, and converge towards the venter. Up to 25 mm diameter, the venter is flattened, very slightly convex, whereafter it becomes rounded and the ventrolateral angulations disappear. At this stage the whorl section is broadly ovate. Constrictions are lacking at all observed growth stages. Measurements No. D H WwW W/H U USNMNH 236915 32:5 25,5(49) +28(53) 1,10 10(19) USNMNH 236917 38,5 20(52) +22(57) 1,10 +9(23) Pe 30 14,5(48) 16(53) 1,10 9(30) USNMNH 236918 43 19(44) 22,3(52) 1,17 9,8(23) USNMNH 236919 16 7,5(47) 8,5(53) 1,13 4(25) USNMNH 236920 18 8(44) 10(55) 125 +4(22) USNMNH 236921 21 10(48) +10(48) 1,00 5(24) USNMNH 236977 17,5 6,8(39) 8,5(49) 1,25 4,5(26) USNMNH 236978 iW 7(41) 9(53) 1,29 4,7(28) T. jurinianus angolanus (holotype) 105 56(53) 51(48) 0,91 17(16) ES 63 32(51) 34(54) 1,06 ? Intraspecific variation The intraspecific variation seen in Tetragonites jurinianus shows features which have an important bearing on the classification of tetragonitids in general. The wide range of inflation within juveniles far exceeds the range of variation admitted by Wiedmann (1973) within the constricted tetragonitids as a whole, and population studies may show his subdivisions of the group to be utilitarian rather than biologically significant. As can be seen from Figure 6, there is not only a distinct decrease in the umbilical ratio with growth, but there is also an ontogenetic increase in the height of the whorls, which become progressively less inflated. Wiedmann (1973) maintained 7. jurinianus angolanus as a separate sub- species by virtue of its narrower umbilicus (16% versus 23% of the diameter in T. jurinianus jurinianus), high-oval whorl section and large size. It appears, however, that these differences result from a comparison of different onto- genetic stages, and the authors include 7. jurinianus angolanus in the synonymy of T. jurinianus sensu stricto. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 187 Fig. 5. Tetragonites (Tetragonites) jurinianus (Pictet). A-B. USNMNH 236918. C-D, G. USNMNH 236921. E-F. USNMNH 236978. H-J. USNMNH 236977. K-M. USNMNH 236917. N-P. USNMNH 236919. Q-S. USNMNH 236920. x1. 188 ANNALS OF THE SOUTH AFRICAN MUSEUM Discussion As diagnosed by Wiedmann (1973: 606), Tetragonites nautiloides (Pictet) does not have constrictions beyond 10 mm diameter and thus closely approaches T. jurinianus. Indeed, the present material all exceeds this diameter and could, therefore, equally well be assigned to 7. nautiloides. However, Murphy (1967a) has noted that constrictions are present to 27 mm diameter in topotype material of T. nautiloides when there is considerable difficulty in distinguishing this 1,50 Wi 0,70 0 30 60 90 DIAMETER (mm) ————> UMBILICUS(%) 0 40 70 110 DIAMETER (mm) ————— Fig. 6. Plot of inflation (whorl width/height ratio) and umbilical ratio against diameter for examples of Tetragonites jurinianus (Pictet). Dots = present Angolan material; squares = the holotype of 7. jurinianus angolanus Haas; circles = material described by Wiedmann (1962, 1973), Murphy (1967a), Wiedmann & Dieni (1968). UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 189 species from some constricted tetragonitids, e.g. 7. rectangularis Wiedmann. Wiedmann’s (1973) diagnosis of T. nautiloides is difficult to support, since some of his figured specimens (pl. 8 (figs 2, 5)) clearly show weak constrictions at 22 and 27 mm diameter respectively, suggesting that this species may better be included in the T. rectangularis group. Occurrence Tetragonites jurinianus ranges from Middle Albian to Lower Cenomanian, and is recorded from Switzerland, France, Sardinia, Mallorca, Madagascar, Angola, Mexico, and California. Family Gaudryceratidae Spath, 1927 Subfamily Gaudryceratinae Spath, 1927 Genus Eogaudryceras Spath, 1927 Type species Ammonites numidus Coquand, 1880 Discussion Eotetragonites was originally separated (Breistroffer 1947) from Eogaudry- ceras by the possession of strong constrictions throughout ontogeny, and a suture with irregularly bifid saddles. However, Wiedmann (19625: 35) has noted the occurrence of species which show combinations of these characters, conse- quently treating Eotetragonites as a subgenus of Eogaudryceras. As defined by him (19625), E. (Eogaudryceras) has initially trapezoidal whorls which become rounded in maturity and are ornamented by fine, flexuous lirae, whereas E. (Eotetragonites) has quadrate juvenile whorls and lacks liration. Eogaudryceras (Eogaudryceras) italicum Wiedmann & Dieni, 1968 Figs 7-11 Eogaudryceras (Eogaudryceras) italicum Wiedmann & Dieni, 1968: 34, pl. 1 (fig. 8), fig. 6. Material Ten specimens, USNMNH 236905-236914, all with recrystallized shell preserved, and all from Porto Amboim. Description Up to 23 mm diameter: shell evolute, compressed, with a wide, shallow umbilicus (30-41 °% of the diameter) and steep umbilical walls which merge imperceptibly with the strongly convex flanks. The whorl section is oval, depressed (W/H = 1,00-1,36), with a broadly rounded venter. The outer whorl conceals about 55 per cent of the preceding whorl. The earliest whorls are very finely lirate, almost smooth to the naked eye, with sporadic, rather distinct, prorsiradiate collars. 190 ANNALS OF THE SOUTH AFRICAN MUSEUM 24-45 mm diameter: the shell form is much as at the earlier growth stages, except that the flanks flatten slightly and the whorl section changes from slightly depressed to almost quadrate. At this stage the lirae are visible to the naked eye; they arise at the umbilical seam and pass forwards (prorsiradiate) on the lower third of the flanks. Thereafter they recurve slightly, only to flex forwards again before crossing the venter. At intervals one or two adjacent lirae are strengthened, presumably corresponding to the collars of the earliest whorls. The lirae become flat-topped, band-like, much broader than the interspaces, and of variable thickness at this stage. Greater than 46 mm diameter: in maturity the whorls become flat-sided and compressed (W/H = 0,66—-0,80), with an elliptical whorl section and narrowly rounded venter. The umbilicus is moderately wide (26-33% of the diameter), shallow, with steep umbilical walls and evenly rounded umbilical shoulders. Maximum width is at about mid-flank. At this growth stage the irregular band- like ribs are split by fine, threadlike grooves. In the largest specimen (still septate at 82 mm diameter) there are still occasional strengthened ribs, followed by a slightly deeper intercostal groove. These may correspond to the collars of the earliest whorls. Fig. 7. Eogaudryceras (Eogaudryceras) italicum Wiedmann & Dieni. Reconstruction of juvenile and adult shells. x1. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 191 C Fig. 8. Eogaudryceras (Eogaudryceras) italicum Wiedmann & Dieni. A~-B. USNMNH 236905. C-E. USNM 236908. x1. 192 ANNALS OF THE SOUTH AFRICAN MUSEUM Measurements No. D H W Wily U USNMNH 236905 81 37,5(46) 25(31) 0,66 21(26) 5s 56 25(45) 19,5(35) 0,78 17,561) USNMNH 236906 34 13(38) 15,5(45) 1,19 +10(29) USNMNH 236907 +47 +21(45) +18(38) 0,86 IS,363% USNMNH 236908 v1 33,5(47) +27(38) 0,80 19(27) USNMNH 236909 26 11(42) 11(42) 1,00 8(31) fe 19,5 8(41) 8(41) 1,00 +5,5(28) USNMNH 236910 24 9(38) 8(38) 1,00 8,5(35) . 18 6,5(36) 7(39) 1,08 7(39) USNMNH 236911 15 5,5(37) 6(40) 1,09 5(30) USNMNH 236912 22 8(36) 9(41) 1,13 8(36) USNMNH 236913 17 5,5(32) +7,5(44) 1,36 7(41) USNMNH 236914 25 9(36) +11(44) 122 +8,5(34) E. (E. italicum* +50 25(50) 19(38) 0,76 +14(28) E. (E.)aenigmum* 58,3 28,3(48,5) 24(41,5) 0,86 14(24) E. (E.)b. bourritianum* 33 14(42) 20(60) 1,43 10(30) E. (E.)b. hispanicum* 47 20(42) 22(47) 1,10 13(28) Gaudryceras aff. madraspatanum Spath (non Stoliczka) 28 11,8(42) 11,8(42) 1,00 10(36) * Asterisks mark holotypes or neotypes. Intraspecific variation The most noteworthy feature of the present material is the great change in whorl section with growth. The earliest whorls show a wide range in inflation (W/a = 1,00-1,36) but, with continued growth, the shell becomes increasingly high-whorled and the whorl section becomes strongly compressed (Fig. 11). At the same time there is a slight, but distinct, tendency for the umbilical ratio to decrease (Fig. 11). Discussion E. (Eogaudryceras) italicum Wiedmann & Dieni is based upon a smooth, fragmentary internal mould from the Upper Albian of Sardinia. It agrees precisely with the present material in whorl section and relative proportions; lack of liration may simply be a reflection of different preservation. Gaudryceras aff. madraspatanum (Stoliczka) (Spath 1923: 22, pl. 1 (fig. 4)) is based upon a smooth phosphatic internal mould from the Cambridge Greensand of southern England. In relative proportions it falls well within the range of variation of the present Angolan material, and the writers would tentatively refer it, therefore, to E. italicum. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 193 K Fig. 9. Eogaudryceras (Eogaudryceras) italicum Wiedmann & Dieni. A-C. USNMNH 236905. D-F. USNMNH 236906. G, K-L. USNMNH 236910. H-J. USNMNH 236909. M-O. USNMNH 236912. x1. 194 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 10. Eogaudryceras (Eogaudryceras) italicum Wiedmann & Dieni. A-B. USNMNH 236907. C-E. 236911. A-D x1, E x2. The present sample is the largest single population of an E. (Eogaudryceras) species yet described, and shows a wide range of ontogenetic and intraspecific variation. In view of this wide range of variation, it seems probable that population studies will reduce in number the twelve species currently assigned to this subgenus, viz. E. (E.) numidum (Coquand), E. (E.) intermedium (Fallot), E. (E.) vocontianum (Fallot), E. (E.) elegans Basse, E. (E.) shimizui Breistroffer, E. (E.) llosetae Breistroffer, E. (E.) turgidum Breistroffer, E. (E.) skoenbergense Collignon, E. (£.) bourritianum (Pictet), E. (E£.) aenigmum (Haas) and E. (£.) muntaneri Wiedmann. Of other Upper Albian species of Eogaudryceras (Eogaudryceras), Wiedmann (1962a: 154) selected a neotype for E. (£.) bourritianum (Pictet) (in Pictet & Roux 1848: 298, pl. 4 (fig. 1)) and divided it into two chronological subspecies: E. (Eogaudryceras) bourritianum bourritianum, from the uppermost Albian (dispar zone) of south-west France is characterized by its strongly depressed whorl section (W/H = 1,43), involute form (U = 30% of the diameter) and smooth whorls (the neotype is preserved as an internal mould and hence the lack of ornament may not be a diagnostic character). It differs from the Angolan material in its much more depressed whorl section and narrower umbilicus; E. (Eogaudryceras) bourritianum hispanicum Wiedmann (1962a: 155, pl. 12 (fig. 6), fig. 15) is a low Upper Albian form (associated with Hysteroceras and Mortoniceras) which was said to differ from the typical form in its less depressed, trapezoidal whorl section (W/H = 1,10), flattened venter, and greater involution (U = 28% of the diameter). It differs from E. (E.) italicum in being more involute, having a slightly more depressed whorl section and a flattened venter. E. (Eogaudryceras) aenigmum (Haas) (1942: 167, pl. 42 (fig. 3), pl. 44 (fig. 2), fig. 24) is from the Upper Albian of Angola. Crushed material assigned to this species, in the British Museum (Natural History), comes from Praia do Jombe UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 195 1,40 we eG —$_—_—____—_ 0,50 0 30 60 90 DIAMETER (mm) => UMBILICUS ( %) —————»> 0 100 DIAMETER (mm) ——— Fig. 11. Plot of inflation and umbilical ratio against diameter for Eogaudryceras italicum Wiedmann & Dieni. Dots = present Angolan material; square = holotype (after Wiedmann & Dieni 1968). 196 ANNALS OF THE SOUTH AFRICAN MUSEUM where it is associated with a low Upper Albian fauna which includes Prohystero- ceras wordiei Spath, Beudanticeras beudanti (Brongniart) and Mortoniceras cf. inflatum (J. Sowerby). It is thus a contemporary of E. (E.) bourritianum hispanicum. The style of ornament is difficult to judge from Haas’s (1942) illustration, although the specimen is clearly lirate (it has shell preserved). Although its whorl width/height ratio is closely comparable to E. italicum (? due to crushing), it is much more involute (U = 24% of the diameter). Haas’s (1942) whorl section of the species shows inner whorls with a strongly fastigiate to keeled venter which suggest his material to be crushed. If true, this species may prove to have priority over E. (E.) bourritianum hispanicum. Occurrence E. (Eogaudryceras) italicum is currently known from the uppermost Albian of Sardinia, Angola, and possibly southern England. Suborder ANCYLOCERATINA Wiedmann, 1966 Superfamily ANCYLOCERATACEAE Meek, 1896 Family Ancyloceratidae Meek, 1876 Subfamily Anisoceratinae Hyatt, 1900 Genus Anisoceras Pictet, 1854 Type species Hamites saussureanus Pictet, 1847 Anisoceras (Anisoceras) perarmatum Pictet & Campiche, 1861 Figs 12A—H, 13C—D, 14A-C, 15C-—F, 16B Hamites armatus J. de C. Sowerby (non J. Sowerby), 1850: pl. 29 (fig. 13). Anisoceras perarmatum Pictet & Campiche, 1861: 65, pl. 48 (figs 7-8), pl. 49 (figs 1-3, 5-7). Pictet, 1861: 21. Ooster, 1863: 19. Pictet & Renevier, 1866: 103. Jukes-Browne, 1875: 288. Barrois, 1878: 271. Renevier, 1890: 340. Boule, Lemoine & Thévenin, 1907: 35. Ganz, 1912: 121. Spath, 1921: 289. Bése, 1923: 144. Diener, 1925: 73. Spath, 19255: 191. Barbu, 1932: 16. Roman, 1938: 52. Spath, 1939: 548, pl. 59 (figs 1-3), pl. 61 (figs 3-7), fig. 192. Breistroffer, 1947: 62. Reyment, 1955: 12, pl. 1 (fig. 1). Collignon, 1963: 50, pl. 259 (figs 1126-1127). Swensen, 1963: 67, pl. 4 (figs 1, 3, 7). Dieni & Massari, 1963: 798. Clark, 1965: 25, pl. 6 (figs 1, 3, 7). Wiedmann & Dieni, 1968: 65, pl. 6 (fig. 4), pl. 7 (fig. 9), figs 38-39. Renz, 1968: 74, pl. 13 (fig. Sa—c), pl. 14 (figs 1, 3, 5), figs 27a, 28g. Hamites (Anisoceras) perarmatus (Pictet & Campiche) von Hauer, 1861: 644, pl. 2 (figs 2-4). Pervinquiére, 1907: 85. Hamites (Anisoceras) saussureanus von Hauer (non Pictet), 1861: 644, pl. 2 (fig. 1). Anisoceras armatus Stoliczka (non J. Sowerby), 1866: 174, pl. 81 (figs 8-10), pl. 82 (fig. 1). Hamites perarmatus (Pictet & Campiche) Neumayr, 1875a: 30; 18755: 898. ? Anisoceras vraconense Renz, 1968: 75, pl. 16 (fig. 2), fig. 27b. Anisoceras perarmatum simplex Renz, 1968: 75, pl. 13 (fig. 7), pl. 14 (fig. 4), fig. 27k. ? Anisoceras pseudopunctatum Pictet & Campiche, 1861: 74, pl. 52 (figs 1-3). Breistroffer, 1947: 62. Renz, 1968: 79, pl. 16 (figs 1, 3, 5). Wiedmann & Dieni, 1968: 72. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 197 Fig. 12. A-H. Anisoceras perarmatum Pictet & Campiche. A-B. USNMNH 236928. C-D. USNMNH 236929. E-F. USNMNH 236944. G-H. USNMNH 236940, I-J. Anisoceras sp. juv., USNM 236745. x1. 198 ANNALS OF THE SOUTH AFRICAN MUSEUM A B C D Fig. 13. A-B. Anisoceras armatum (J. Sowerby), USNMNH 236934. C-D. Anisoceras perarmatum Pictet & Campiche, USNMNH 236931. x1. Material Twenty-seven specimens, SAM-PCA2940, 2950, 2956, 3115, 3122, 3143, 3148, 3152, 3193, 3214, 3243, 3283, 3285, 3299, 3339, 3352, 3355, 3358, 3361 and 3391, all from Egito, SAM-—PCA4801, 4804 from Cabo Ledo and USNMNH 236928-29, 236931, 236933, 236940, and 236944 from Porto Amboim, either retaining recrystallized shell or preserved as composite internal moulds. Description Whorl section varies from almost circular in specimens replaced by calcite to slightly elliptical in those crushed individuals preserved as composite internal moulds. Ornament comprises small dorsolateral to lateral tubercles and the septate bases of prominent ventrolateral spines, between which strong ribs are looped in pairs. There are sporadic non-tuberculate intercalatories between looped ribs. The main ribs are also looped across the venter, and split into fine riblets, thirteen per three dorsolateral tubercles, on the dorsum. On the body chamber there may be one to two simple ribs (SAM-PCA3143, 3148) between looped ribs, whilst on the final shaft of USNMNH 236940, the button-and-loop ribbing is lost, and all ribs become single. Discussion Amongst contemporaneous species of Anisoceras, only A. saussureanum and A. armatum closely approach this species. Differences are noted under the discussion of A. armatum. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 199 kat Fig. 14. A-C. Anisoceras perarmatum Pictet & Campiche. Dorsal, lateral and ventral views of SAM-PCA3115. D-E. Anisoceras cf. armatum (J. Sowerby). A fragment of a body chamber, SAM-PCA3287, which may belong here. F. Anisoceras sp. Oblique lateral view of SAM- PCA3220. x0,75S. 200 ANNALS OF THE SOUTH AFRICAN MUSEUM Anisoceras vraconense Renz (1968: 75, pl. 16 (fig. 2), fig. 27b) differs from A. perarmatum in having most ribs simple and much weaker dorsolateral tubercles. This species may be based upon a fragment of the final shaft of A. perarmatum, since the ribbing simplifies on the body chamber. Anisoceras charlottense Anderson (1958: 209, pl. 11 (fig. 3)) has a circular whorl section and very prominent looped ribs separated by three to four simple, finer intercalatories. It may be conspecific with the low Upper Albian A. salei Clark (1958: 1079, pl. 140 (fig. 3)). Anisoceras perarmatum simplex Renz (1968: 75, pl. 13 (fig. 7), pl. 14 (fig. 4), fig. 27k) is simply a variant based on a small body chamber fragment. Anisoceras pseudopunctatum Pictet & Campiche (1861: 74, pl. 52 (figs 1-3)) is based on material which, like A. perarmatum simplex, has also lost button-and- loop ribbing, and may equally be an intraspecific variant. Occurrence Anisoceras perarmatum is a typical dispar Zone species known from England, France, Switzerland, Sardinia, Tunisia, Nigeria, Madagascar, southern India, Texas, and Angola. Anisoceras (Anisoceras) armatum (J. Sowerby, 1817) Figs 13A—B, 14D-E, 16A, C, E, I, 17-19 Hamites armatus J. Sowerby, 1817: 153, pl. 168. De Haan, 1825: 152, no. 2. Buckland, 1837: 65, pl. 44 (figs 9-10). Brown, 1837: 2, pl. 2 (fig. 6). Romer, 1840: 94, pl. 15 (fig. 2). D’Orbigny, 1842: 547, pl. 135. Non Hamites armatus Mantell (non J. Sowerby), 1822: 121, pl. 23 (figs 3-4) (= A. plicatile). Baculina armata (J. Sowerby) Fleming, 1828: 250. ? Hamites undulatus Brown (non Forbes), 1837: pl. 2 (fig. 11). Non Hamites armatus J. de C. Sowerby (non J. Sowerby), 1850: pl. 29 (fig. 13) (= A. perarmatum). ? Hamites armatus J. Sowerby, Dixon, 1851: pl. 29 (fig. 13). Anisoceras armatum (J. Sowerby) Pictet & Campiche, 1861: 62, pl. 48 (figs 1-2, 4, 6). Spath, 1939: 543, pl. 59 (fig. 6), pl. 60 (fig. 1), pl. 61 (figs 9-11), pl. 62 ( ?fig. 5), fig. 191. Swensen, 1963: 66, pl. 3 (fig. 4), pl. 4 (?fig. 6). Clark, 1965: 25, pl. 5 (fig. 4), pl. 6 ( ?fig. 6), fig. 7a. Renz, 1968: 75, pl. 15 (figs 1, 3), figs 27d, 28a. ? Hamites (Anisoceras) armatus J. Sowerby, von Hauer, 1861: 644, pl. 1 (figs 9-10). Non Anisoceras armatum Stoliczka (non J. Sowerby), 1866: 174, pl. 81 (figs 8-10), pl. 82 (fig. 1) (= A. perarmatum). Hamites (Anisoceras) armatus J. Sowerby, Kossmat, 1895: 149. ? Anisoceras armatum (J. Sowerby) Choffat, 1905: 41, pl. 1 (fig. 6). Non Hamites (Anisoceras ?) armatus Pervinquiére (non J. Sowerby), 1907: 84, pl. 4 (figs 2-3) (= ? A. exoticum). Anisoceras aff. armatum (J. Sowerby) Adkins, 1920: 69. ? Hamites cf. armatus J. Sowerby, Passendorfer, 1921: 237. Anisoceras cf. armatum (J. Sowerby) Bose, 1923: 143, pl. 10 (figs 22-24). Haughton, 1924: 94. Clark, 1958: 1080, pl. 139 (fig. 2). Non Anisoceras aff. armatum (J. Sowerby) Spath, 19256: 190 (= A. raynaudi). Anisoceras picteti Spath (non Matheron), 1926a: 432. Spath, 1939: 554, pl. 59 (fig. 4), pl. 61 (fig. 8), pl. 63 (figs 3, 8), fig. 194. Renz, 1968: 76, pl. 13 (figs 8-9), pl. 14 (figs 6-9), pl. 15 (fig. 4), figs 27c, 28f. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 201 Fig. 15. A-B. Anisoceras sp. Oblique dorsolateral and lateral views of SAM-PCA3301 from Egito. C-F. Anisoceras perarmatum Pictet & Campiche. C-D. Ventral and lateral views of SAM-PCA3154. E-F. Lateral and ventral views of SAM-—PCA4801. G-—H. Anisoceras sp. An indeterminate body chamber fragment from Egito, SAM-PCA2942. I-J. Mortoniceras (Angolaites) simplex (Choffat). Lateral and ventral views of SAM-—PCA3249. A-D, G-H x 0,75, E-F, I-J x1. 202 UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL ? Hamites aff. armatus J. Sowerby, Scott, 1926: 80. Bose, 1928: 146. Adkins, 1928: 24. ? Anisoceras cf. armatum (J. Sowerby) Passendorfer, 1930: 667. ? Non Hamites (Anisoceras) armatus Collignon (non J. Sowerby), 1932: 20, pl. 4 (fig. 1). Anisoceras Saussureanum var. spinosa Haas, 1942: 192, pl. 46 (figs 1-3), fig. 30. Non Anisoceras armatum Haas (non J. Sowerby), 1942: 189, fig. 29 (= A. tropicale). Anisoceras jacobi Breistroffer, 1946: 310; 1947: 62. Wiedmann & Dieni, 1968: 67, pl. 6 (fig. 13), fig. 41. ? Anisoceras aff. picteti Spath, Kennedy, 1971: 13, pl. 3 (fig. 6), pl. 7 (fig. 10). ? Non Anisoceras armatum (J. Sowerby) Kennedy, 1971: 14, pl. 5 (fig. 11). Material Fourteen specimens, SAM-—PCA2932, 2947, 2953, 3174, 3364, and 3398 from Egito, USNMNH 236930, 236934, 236936-37, 236939, and 9236952 from Porto Amboim, and SAM-—PCA4606 and 4610 from Cabo Ledo. Description Shell form suggests initial coiling in an open planispiral, straightening in maturity. The whorl section is approximately circular. The ornament is rather variable, comprising slightly rursiradiate to slightly prorsiradiate looped ribs, tetween which are generally one, sometimes two, simple intercalatories. Looped ribs are ornamented with small pointed dorso- lateral tutercles and the septate bases of prominent ventrolateral spines. All rits are of more or less equal strength. Across the dorsum the ribs divide into fine riblets and are accompanied by intercalatories. In USNMNH 236939, which is taken to represent an early portion of the spire of the species, main ribs are weakly looped and are separated by two to three intercalatories. In USNMNH 236952, the largest specimen (which is, however, still septate), the whorl section is slightly compressed, oval, and there is generally only one intercalatory between looped ribs, although even these are sometimes absent. Discussion Anisoceras armatum is a widely-cited but poorly understood species, and in consequence the holotype is refigured here (Fig. 17). As noted by Spath (1939: 546), this is a crushed, composite internal mould which shows the following features: the penultimate shaft has distinct looped ribs separated by only one non-tuberculate intercalatory which is of approximately the same strength as the looped ribs. At this stage there are about five ribs in a distance equal to the whorl height (allowing for post mortem crushing). On the hook, the preservation is poor, but the looped ribs clearly break up so as all to become single on the final non-septate shaft. At this stage there are still only about five ribs in a distance equal to the whorl height, and most of the ribs are of approximately the same strength. The dorsolateral tubercles are still present in maturity, although weak and irregularly developed. It is clear from the above description, and comparison with Sowerby’s original illustration (cf. Spath 1939, fig. 191), that this species has been mis- UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 203 Fig. 16. A, C, E, 1. Anisoceras armatum (J. Sowerby). A. SAM-—PCA5470. C. SAM-PCAS5471. E. SAM-PCAS5473. I. SAM-PCAS5472. B. Anisoceras perarmatum Pictet & Campiche. Lateral view of SAM-PCA3339, showing change of ornament on the body chamber. D. Hamites duplicatus Pictet & Campiche. Lateral view of SAM-PCA2955. F. Hamites virgulatus Brongniart. Lateral view of SAM-PCA3158, showing venefzianus-type ribbing. G. Mariella (Mariella) gresslyi (Pictet & Campiche). SAM-PCA3133. H. Hamites virgulatus Brongniart. Lateral view of SAM-PCA2959, showing rather dense subvirgulatus-type ribbing. J-K. Anisoceras phillipsi sp. noy. Lateral and dorsal views of SAM-PCA 4799. B 0,75, other x1. 204 ANNALS OF THE SOUTH AFRICAN MUSEUM interpreted. Indeed, Sowerby’s figure bears no great resemblance to the original and shows too many intercalated ribs both on the penultimate and final shafts, while the differentiation of the ribs on the final shaft is less prominent than is shown in Sowerby’s drawing. Anisoceras jacobi Breistroffer (nom. nov. pro Anisoceras picteti Spath (non Matheron) 1939: 554, pl. 59 (fig. 4), pl. 61 (fig. 8), pl. 63 (figs 3, 8), fig. 194) was diagnosed as follows: ‘Like A. armatum, but more coarsely ornamented, with Fig. 17. Anisoceras armatum (J. Sowerby). Sowerby’s (1817, pl. 178) original figured specimen from the Upper Greensand of Roak, near Benson, Oxfordshire. Oxford University Museum K675a. x 1. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 205 A B C Fig. 18. A-C. Anisoceras armatum (J. Sowerby), USNMNH 236952. x1. the intermediate ribs (one, rarely two) as prominent as the main ribs that meet at the strong tupercles. Suture-line similar to that of A. armatum.’ These are the exact features which characterize the penultimate shaft of the holotype of the contemporaneous A. armatum, and the authors regard A. jacobi as a junior subjective synonym of A. armatum. Anisoceras armatum and A. perarmatum are contemporaneous species which differ in that the latter generally lacks intercalatories between the 206 ANNALS OF THE SOUTH AFRICAN MUSEUM looped ribs in maturity, although there are intermediates between the two species, e.g. USNMNH 236952 (Fig. 18). Spath (1939) noted the very close resemblance between A. saussureanum (Pictet) and A. armatum and considered immature growth stages indistinguish- able. In maturity, A. saussureanum was said to differ by its distant tuberculation, with commonly two to three intercalatories between looped ribs, whilst coming from a lower level in the Albian. Spath (1939) appeared to place much weight, in his separation of the above two species, on the supposedly lower horizon of t \ v h itis ail 77) / £. som aka F G H Fig. 19. Anisoceras armatum (J. Sowerby). A-B. USNMNH 236930. C-D. USNMNH 236936. E-F. USNMNH 236937. G-H. Doubtful juvenile, USNMNH 236939. x1. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 207 A. saussureanum, although Breistroffer (1940, 1947) and Renz (1968) record it from the uppermost Albian of France and Switzerland respectively. Moreover, A. saussureanum spinosum (Haas) is recorded from the uppermost Albian of Angola and Mexico (fide Haas 1942). The holotype of A. saussureanum spinosum comes from the same locality as the Washburn collection and the authors have no hesitation in regarding it as a junior subjective synonym of A. armatum. However, Haas’s identification clearly focuses on the difficulties of separating A. armatum and A. saussureanum and, in view of the fact that they are contem- porary species in the uppermost Albian, they probably do not bear specific separation. However, until the type and topotype material of A. saussureanum are restudied with regard to their intraspecific variation, it seems preferable to retain these two well-known species separate. Anisoceras tropicale (Meunier) (1887: 62, pl. 1 (fig. 5)) is based upon a very poorly preserved internal mould which was tentatively referred to A. armatum by Choffat (1905: 41) and Haas (1942: 191), although Spath (1939: 558) considered it closer to A. pseudoelegans. Meunier’s specimen shows three fine intercalatories between looped ribs and, judging from the locality, a valley to the north of Lobito, is somewhat older than A. armatum. This is supported by the example of A. armatum figured by Haas (1942, fig. 29), which the authors would assign to A. tropicale; it occurs on the reverse side of the holotype of Mortoniceras vokesi (Haas), topotype material of which comes from the mid-Upper Albian zone of Elobiceras elobiense (Cooper 1978) at Lobito, in association with Puzosia cuvervillei (Meunier). Anisoceras exoticum Spath (1939: 555, pl. 59 (fig. 7), pl. 60 (fig. 4), pl. 63 (fig. 2), fig. 195) differs from A. armatum in having four to six intercalatories between main ribs, with very feebly-developed dorsolateral tubercles. The ribbing of this species is also attenuated across the siphonal line. In Anisoceras oldhamianum (Stoliczka) (1865: 135, pl. 83 (figs 1-4), pl. 92 (fig. 1)) all the ribs of the early growth stages are tuberculate and presumably looped, whilst on the straight shaft there is a fine, non-tuberculate rib separating looped ribs. The early whorls are coiled in a shallow, open helix. Occurrence Anisoceras armatum is at present known with certainty only from the uppermost Albian of southern England, France, Switzerland, Sardinia, Angola, Texas, and Mexico. Anisoceras haasi sp. nov. Figs 20-22 Idiohamites (?) indet. sp., Haas, 1942: 195, pl. 45 (fig. 6). Idiohamites spiniger Haas (non J. Sowerby), 1942: 195, pl. 46 (fig. 4), fig. 31a—b. Idiohamites indet. sp., Haas, 1942: 197, pl. 45 (fig. 7), fig. 31c. Idiohamites aff. subspinigero Haas (non Spath), 1942: 197, pl. 46 (fig. 5), fig. 31d-e. ANNALS OF THE SOUTH AFRICAN MUSEUM 208 ‘TX ‘sode}S YIMOIS I[PPIUI puv ATIvSd JO UOTJONI}SUODEY “AOU ‘ds IspDY SDAQIOSIUP “OZ “BIZ inh ee oe, fo we eo mmctcqea UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 209 Fig. 21. Anisoceras haasi sp. nov. A-C. USNMNH 236923. D, F, H. USNMNH 236924, E, G. USNMNH 236922. I-J. USNMNH 236925. x1. 210 ANNALS OF THE SOUTH AFRICAN MUSEUM Material Seven specimens, USNMNH 236922-7, all retaining recrystallized shell and all from Porto Amboim, together with a single crushed internal mould, SAM-PCA2952 from Praia-Egito. Type material USNMNH 236922 is designated holotype; the remaining specimens from Porto Amboim are paratypes. Etymology Named for Otto Haas, pioneer of ammonite studies on the Angola Cretaceous. Diagnosis A densely ribbed species of Anisoceras with 8-15 ribs in a distance equal to the whorl height; 2 or 3 ribs are looped from small, acute dorsolateral tubercles to prominent rounded bases of septate ventrolateral spines, commonly with 3-4 looped between tubercles across the venter; there are between 2 and 9 fine intercalatories separating looped ribs. Whorl section circular to elliptical, compressed. Description Up to 13 mm whorl-width: at this stage, the shell is coiled in a loose planispire, and has a slightly depressed to circular cross-section. Ornament comprises fine, slightly rursiradiate ribs, about as wide as the interspaces and effaced across the dorsum. There are periodic broad bulges which follow the course of the ribbing and are ornamented with very weak, pointed dorsolateral tubercles and the septate bases of large, prominent, ventrolateral spines. The fine ribs commonly arise in pairs from the dorsolateral tubercles and are joined by a third rib to meet the ventrolateral spine-base. Three to four ribs are looped across the venter connecting the ventrolateral spines. There are four to nine fine inter- calated ribs between adjacent bulges. 14-16 mm whorl-width: beyond 13 mm whorl width, the shell begins to uncoil, developing an almost straight shaft. At the same time, the whorl section becomes slightly compressed (“/H = 0,83-0,86) and elliptical. The bulges become more regular with generally three in a distance equal to the whorl height, and commonly with two to three fine ribs separating adjacent bulges. Ribbing remains rursiradiate at this stage and the dorsolateral tubercles have migrated up flank to a low lateral position. Discussion The lectotype of A. pseudoelegans (Renz 1968, pl. 14 (fig. 12)) differs from the Angolan material in having a strongly compressed whorl section with flattened flanks and maximum width near the dorsolateral tubercles, whilst it is UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 211 Fig. 22. Anisoceras haasi sp. nov. Dorsal and lateral views of a slightly crushed composite internal mould, SAM-—PCA2952, from Egito. x1. more coarsely ribbed, with ribs arising in bundles from distinctly bullate dorso- lateral tubercles. Juveniles of A. pseudoelegans also appear to be much more coarsely ribbed than the present material (compare Renz 1968, pl. 14 (figs 10-11)). According to Kennedy (1971: 12, pl. 3 (figs 12-13), pl. 4 (figs 1-3)), Anisoceras plicatile is a Middle Cenomanian species with 12 fine ribs in a distance equal to the whorl height, and a circular whorl section. 2 or 3 ribs are looped between the midlateral and ventrolateral tubercles and are separated by 1-3 non-tuberculate intercalatories. Anisoceras plicatile differs from the present material, therefore, in having much more swollen main ribs and a much more prominent lateral tubercle which is at the middle of the flank, not dorso- lateral as in A. haasi. The Angolan species also has more numerous, fine intercalatories, whilst the ribbing is not as coarse in maturity as it is in A. plicatile (compare Kennedy 1971, pl. 2 (fig. 12)). Anisoceras bendirei (Adkins) (1920: 8, pl. 11 (fig. 1)) from the late Albian of Texas differs from the Angolan material in being much more coarsely ribbed, with the lateral tubercle higher on the flanks. Anisoceras raynaudi (Boule, Lemoine & Thévenin 1907: 170, pl. 4 (figs 7-8), fig. 38) is a finely ribbed species in maturity, which differs from A. Aaasi in having fewer intercalatories (only 2-3) between looped ribs in the early growth stages, a more prominent dorsolateral tubercle and, judging from material from Catuane, southern Mozambique, in the South African Museum, Cape Town, in 212 ANNALS OF THE SOUTH AFRICAN MUSEUM having the early whorls coiled in an open helical spire up to 20 cm in height. It is also an earlier species (orbignyi-varicosum Subzones). Anisoceras nanaense (von Hauer) (1861, pl. 1 (figs 11-12)) differs from the present species in lacking dorsolateral tubercles, and in having prorsiradiate ribs with only two to three intercalatories between tuberculate ribs. Anisoceras arrogans (Giebel) (1852: 305) (nom. nov. pro Hamites elegans d’Orbigny (non Parkinson) 1842: 542, pl. 133 (figs 1-5)) can be distinguished from A. haasi in its lack of dorsolateral tubercles, with only two to four non- tuberculate intercalatories, and in the coarse, distant ribbing of the body chamber. Occurrence Anisoceras haasi sp. nov. is at present known only from the uppermost Albian of Angola. Anisoceras phillipsi sp. nov. Figs 16J-K, 23-24 Material Eight fragments, SAM—PCA2974, 3179, 3183, 3211-3212, 3220 and 3222, all preserved as composite internal moulds, from Praia-Egito, and SAM-— PCA4799 from the Quissama Ridge at Cabo Ledo. Type material The specimen illustrated as Figure 23, SAM-—PCA3183 from Praia-Egito is designated as holotype. All other specimens cited above are paratypes. Etymology The species is named for Denis Phillips of the British Museum (Natural History) who, during many years, has given both authors invaluable assistance and advice in connection with their researches. Diagnosis A large species of Anisoceras with subrectangular, depressed to ovate whorl section in maturity. Prominent dorsolateral clavae give rise to two, occasionally only one, rursiradiate ribs which meet the well-developed ventrolateral clavae singly or in pairs. There are no intercalated ribs. Across the venter the ribs are looped or single. Description In the smallest example, SAM-—PCA3179, the whorl section is elliptical, compressed, although it may have suffered lateral compaction. All the other larger fragments show a strongly depressed, subrectangular whorl section (Fig. 24) prior to the final hook, and an ovate whorl section afterwards. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 213 Fig. 23. Anisoceras phillipsi sp. nov. Dorsal, ventral and lateral views of the holotype, SAM-PCA3183, preserved as an internal mould. 0,75. 214 ANNALS OF THE SOUTH AFRICAN MUSEUM (\O) Seq Fig. 24. Anisoceras phillipsi sp. nov. Whorl sections. A-B. SAM-—PCA2974. C. SAM- PCA3183. D. SAM-—PCA3222. x1. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 215 On the dorsolateral shoulder, prominent clavae give rise to commonly two, sometimes only one, coarse, rursiradiate flank ribs which, on curved fragments, are strongly convex adorally. The flank ribs meet the well-developed ventro- lateral clavi either singly or in pairs, with six to eight ribs in a distance equal to the whorl height. Across the venter, ribs are single or looped. Discussion The only species with which the present form may usefully be compared is Anisoceras arrogans (Giebel) (= Anisoceras campichei Spath 1942: 559, pl. 63 (figs 6-7), fig. 197) which differs in lacking the dorsolateral clavae and in the regular nature of its ribbing across the venter. Occurrence Anisoceras phillipsi sp. nov. is known only from the uppermost Albian of Angola. Anisoceras cf. arrogans (Giebel, 1852) Figs 25-26 Compare Hamites elegans d’Orbigny (non Parkinson), 1842: 542, pl. 133 (figs 1-5). Hamites arrogans Giebel, 1852: 305. Anisoceras pseudoelegans Pictet & Campiche, 1861: pl. 50 (figs 6-7 only). Anisoceras campichei Spath, 1926: 432; 1939: 559, fig. 197 only. Metahamites (?) arrogans (Giebel) Spath, 1939: 559. Anisoceras aff. campichei Spath, 1939: pl. 63 (figs 6-7). Anisoceras arrogans (Giebel) Wiedmann & Dieni, 1968: 69, pl. 7 (fig. 10), pl. 8 (figs 5, 7, 11), figs 46-50. Description This species is known only from large fragments. In SAM-—PCA3205, the whorl section is slightly compressed, subquadrate, but the remaining material shows some variation in this character. Ornament comprises prominent, single flank ribs arising from weak umbilical tubercles and passing slightly prorsiradiate across the flanks to the bases of large, septate ventrolateral spines. Ribs are either single or looped across the venter, and the rib pattern is shown in Figure 26. There are six to seven flank ribs in a distance equal to the whorl height. Across the dorsum, fine ribs are looped between the weak dorsolateral tubercles, with generally an intercalated rib between pairs, so that there are about fifteen ribs across the dorsum per five dorsolateral tubercles. Discussion The present specimen is closest to Anisoceras arrogans (Giebel) (= A. campichei Spath, 1939: 559, fig. 197 only) from which it differs in its subquadrate whorl section, slightly prorsiradiate flank ribs, and the fact that all 216 ANNALS OF THE SOUTH AFRICAN MUSEUM ribs across the venter arise from ventrolateral tubercles. Anisoceras arrogans also does not show the peculiar zigzagging of the ribs across the venter (see Fig. 26) seen in the Angolan material. However, since A. arrogans and the present material represent different ontogenetic stages, the differences may not be as great as they first appear, and consequently the present material is identified as A. cf. arrogans (Giebel). Fig. 25. Anisoceras cf. arrogans (Giebel). Dorsal, ventral and lateral views of SAM—PCA3205. x 0,75. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 217 Fig. 26. Anisoceras cf. arrogans (Giebel). Schematic rib pattern across the venter of SAM-—PCA3205. x 0,75. Anisoceras phillipsi sp. nov. differs from the present material in the posses- sion of distinctly clavate dorsolateral tubercles, from which ribs frequently arise in pairs. Occurrence Anisoceras arrogans is known from the uppermost Albian of Switzerland, France and Sardinia, and may also be present in southern England and Angola. Anisoceras aff. exoticum Spath, 1939 Fig. 27 Compare Anisoceras saussureanum Pictet & Campiche (non Pictet), 1861: 118, pl. 1 (fig. 2). Anisoceras exoticum Spath, 1939: 555, pl. 59 (fig. 7), pl. 60 (fig. 4), pl. 63 (fig. 2), fig. 195. Material A single specimen, SAM-PCA3174, preserved as a composite internal mould, from Praia-Egito. 218 ANNALS OF THE SOUTH AFRICAN MUSEUM Description The specimen represents a slightly crushed fragment of a straight shaft in which the whorl section was originally probably almost circular. Ornament comprises slightly rursiradiate main ribs between which are three to seven fine, secondary ribs, the density of which increases adorally. The fragment lacks obvious tuberculation. Fig. 27. Anisoceras aff. exoticum Spath. A body chamber fragment, SAM-PCA3174. 0,75. Discussion The specimen is a body chamber fragment and, since ornament frequently changes drastically on the body chamber of Anisoceras, reference to Spath’s species is somewhat tentative. Occurrence Anisoceras exoticum is known from England, Switzerland, Sardinia, and possibly Angola. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 219 Anisoceras aff. subarcuatum Spath, 1939 Fig. 28 Compare Anisoceras subarcuatum Spath, 1939: 560, pl. 65 (fig. 1), pl. 66 (fig. 1), fig. 198. Material A single specimen, SAM-—PCA3143, from the dispar Zone at Praia-Egito. Description A short fragment shows affinities with Spath’s species and may belong here. The whorl section is slightly compressed, oval. Ornament comprises slightly tursiradiate, strengthened, simple main ribs which bifurcate or trifurcate across the dorsum and are ornamented with dorsolateral and ventrolateral tubercles. Between main ribs are two to three, slightly weaker, somewhat irregular secondary ribs, which occasionally bifurcate across the venter. All the secondary ribs lack tubercles. Discussion Body chamber fragments of Anisoceras are notoriously difficult to identify and hence full determination of the present specimen must await further material. Fig. 28. Anisoceras aff. subarcuatum Spath. Ventral, lateral and dorsal views of SAM-PCA3143. x 0,75. 220 ANNALS OF THE SOUTH AFRICAN MUSEUM Occurrence Anisoceras subarcuatum is known with certainty only from the low Upper Albian (varicosum and auritus subzones) of southern England, but may also be present in the uppermost Albian of Angola. Anisoceras aff. spathi (Wiedmann, 1962) Fig. 29 Compare Anisoceras subarcuatum Spath, 1939: 560, pl. 63 (fig. 5 only). Idiohamites spathi Wiedmann, 1962a: 188. Material A single fragment, SAM-—PCA3223, from the upper part of the dispar Zone at Praia-Egito, preserved as a composite internal mould. Fig. 29. Anisoceras aff. spathi (Wiedmann). Ventral and lateral views of SAM—PCA3223. 0315: UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 221 Description The single fragment available represents part of the crozier. It has a compressed, elliptical whorl section with simple rursiradiate ribs. On the adoral portion of the fragment all ribs are relatively fine, of even strength, and lack tuberculation. On the adapical portion, every second rib is strengthened slightly and bears weak ventrolateral tubercles. Discussion The absence of lateral or dorsolateral tubercles led Wiedmann (1962a) to assign the present species to Idiohamites. However, the holotype is a body- chamber fragment which merely shows the radical change in ornament on the body chamber displayed by many Anisoceras species. Occurrence Anisoceras spathi is known from the low Upper Albian of southern England and Spain, and may be present in the dispar Zone of Angola. Anisoceras sp. indet. Fig. 14F Material A single, crushed fragment, SAM-—PCA3220, from Praia-Egito, preserved as a composite internal mould. Description Only the greater part of one flank, one row of ventrolateral tubercles and much of the venter of the specimen concerned is preserved. Rather fine (?looped) ribs on the flanks are slightly narrower than the interspaces, with at least 10 in a distance equal to the whorl height. The flank ribs join the ventrolateral clavi in pairs whereas, across the venter, the ventro- lateral tubercles are connected by 2-3 ribs so that for 16 flank ribs, there are 20 ribs across the venter. The whorl section gives the appearance of having been strongly depressed. Discussion The specimen under discussion is too poorly preserved for specific identifi- cation, but may be regarded as belonging to the plicatile-haasi plexus. Occurrence Upper Albian, dispar Zone, Praia-Egito. 222 ANNALS OF THE SOUTH AFRICAN MUSEUM Genus Idiohamites Spath, 1925 Type species Hamites tuberculatus J. Sowerby, 1818 Discussion Wiedmann (1962a) separated Idiohamites from Anisoceras because the former was considered to have the early whorls coiled in a helix, whereas in Anisoceras the juvenile whorls were considered to be planispirally coiled. The differences are not, however, so clear cut, since Anisoceras of the type referred to A. raynaudi (Boule, Lemoine & Thévenin) by Forster (1975) (= Anisoceras saussureanum quadrifasciatum Klinger) from the low Upper Albian of Catuane, Mozambique, and now in the South African Museum, show the early whorls to be coiled in an open helix up to 20 cm high. The straight final shafts of this species are figured by Forster (1975, pl. 4 (fig. 8)). Moreover, Matsumoto (1959, pl. 28 (fig. 1), pl. 29 (fig. 2)) has also figured an Anisoceras in which the early growth stages are helically coiled. Nor is the presence or absence of lateral tubercles a diagnostic feature since some Anisoceras, e.g. A. auberti (Pervinquiére) and A. gracile Renz, lack them. Moreover, Renz (1968) has recently figured a number of specimens which he considers transitional between Jdiohamites and Anisoceras in this respect (cf. pl. 12 (figs 7-8), pl. 13 (figs 3, 6)). In addition both genera show a suture line with a fairly shallow, bifid external lobe (E), an asymmetrically bifid first lateral saddle (E/L), a bifid first lateral lobe (L) which is deeper than the external lobe, and a second lateral saddle (L/U) which is bifid, and a bifid second lateral lobe (U) which is almost as deep as the first. Clearly the two genera are very closely related although, at present, the typically smaller size, almost complete absence at any growth stage of looped flank ribs, and the helically coiled juvenile whorls, are considered to be sufficient for the generic separation of Idiohamites from Anisoceras. Idiohamites dorsetensis Spath, 1926 Figs 30, 31G Anisoceras alternatus Pictet & Campiche (non Mantell), 1861: 71, pl. 51 (figs 1, 3-4). Idiohamites dorsetensis Spath, 1926b: 432; 1939: 596, pl. 62 (figs 2-3), pl. 63 (figs 1, 9, 15), pl. 65 (fig. 2), fig. 215. Renz, 1968: 70, pl. 11 (figs 39-40), pl. 12 (figs 3-4), figs 25a-d, f, 26a-d. ? Idiohamites aff. turgidus robustus Spath, Renz, 1968: 72, pl. 11 (figs 33-37), figs 25k—I, 26f-h. ? Idiohamites elegantulus laticostatus Renz, 1968: 73, pl. 11 (figs 38, 41-42), pl. 12 (figs 1-2), figs 25m, 26i-m. Material Three specimens in the South African Museum, SAM-—PCA4803 and two unnumbered fragments, retaining recrystallized test, together with USNMNH 236951, preserved as a composite internal mould from Porto Amboim, and a single specimen from Cabo Ledo, SAM-PCA5469. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 223 Fig. 30. A-—D. Idiohamites dorsetensis Spath. A-B. Ventral and lateral views of SAM-PCA4803. C-—D. Ventral and lateral views of SAM-PCAS5S469. x1. Description All the fragments show the shell to be loosely coiled in a single plane, with an elliptical, compressed whorl section. Ornament comprises simple, strong, slightly prorsiradiate ribs, of which there are four to five in a distance equal to the whorl height. Every second or third (or occasionally adjacent) rib bears very small, sharp, pointed lateral tubercles and more prominent ventrolateral spines. At small growth stages, tubercles are connected across the venter by simple ribs but, on the mature body chamber, they may be weakly looped. 224 ANNALS OF THE SOUTH AFRICAN MUSEUM Discussion Until populations of individuals are available for study, the differences between several named species of /diohamites appear suspiciously small. The authors assign their material to J. dorsetensis because of the very close similarities to the types, and because it is the oldest available name for material of this age. Occurrence Idiohamites dorsetensis is known from the uppermost Albian of southern England, France, Switzerland and Angola. Idiohamites cf. elegantulus Spath, 1939 Fig. 31A—D Compare Idiohamites elegantulus Spath, 1939: 599, fig. 216. Material One specimen, USNMNH 236950, a composite internal mould from Porto Amboim. Discussion The specimen consists of a body chamber hook with a maximum whorl height of 12 mm. There are six fine prorsiradiate ribs in a distance equal to the whorl height, the majority bearing ventral tubercles, suggesting reference to Spath’s species. Occurrence Stoliczkaia dispar Zone of England, and possibly Angola. Idiohamites pygmaeus sp. nov. Fig. 31J-N Material Eight specimens, USNMNH 236942-49, all retaining recrystallized shell and all from Porto Amboim. Type material USNMNH 236942, a complete adult, is designated holotype; the remaining specimens are paratypes. Etymology From the Latin adjective pygmaeus, pygmy-like, dwarfish; derived from the Greek pygmaios. The pygmaioi were a fabulous dwarfish race of antiquity, especially in Africa; at war with the cranes, they were constantly defeated (Pliny). UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 225 L Fig. 31. A-D. Idiohamites cf. elegantulus Spath. A-B. USNMNH 236948. C-D. USNMNH 236938. E-F. Mariella (Mariella) cirumtaeniatus (Kossmat), USNMNH 236958. G. Idiohamites cf. dorsetensis Spath,h USNMNH 236951. H-I. Borissiakoceras sp. noy.? aff. reymenti (Brunnschweiler), USNMNH 236980. J-N. Jdiohamites pygmaeus sp. nov. J-L. USNMNH 236942. M-N. USNMNH 236943, x1. 226 ANNALS OF THE SOUTH AFRICAN MUSEUM Diagnosis A very small Jdiohamites with a known maximum length of only 52 mm, characterized by helicoid early whorls with three quadrituberculate rursiradiate ribs in a distance equal to the whorl height, a straight shaft with four prorsi- radiate ribs in a distance equal to the whorl height and ribbing tending to be rectiradiate at the aperture. Description Shell small, initially coiled in a low, open helical spire, straightening in maturity before recurving on the body chamber to form a crozier. Earliest whorls ornamented with rather robust ribs, narrower than the interspaces and with three in a distance equal to the whorl height, which cross the venter obliquely so that the four rows of tubercles on the main ribs are asymmetrically arranged. There is a regular alternation of tuberculate and non- tuberculate ribs, which, at this stage, are slightly rursiradiate. On the first straight shaft, which commences at a whorl height of 6 mm, the ribbing becomes prorsiradiate, with still only three ribs in a distance equal to the intercostal whorl height. With the uncoiling of the shell, the tuberculation is no longer asymmetrical and main ribs are ornamented with small pointed dorsolateral and ventrolateral tubercles. The rib direction changes to rectiradiate on the final shaft. In maturity, the intercostal whorl section is oval, depressed, although it is almost circular at small diameters. Discussion This species is characterized by its small size at maturity. Idiohamites dorsetensis 1s adult at much larger diameters, with a slightly compressed whorl section, and closer ribbing (four to six in a distance equal to the whorl height). Idiohamites elegantulus is even more densely ribbed (seven in a distance equal to the whorl height), also with a compressed whorl section. /diohamites recticostatus Renz is a much larger species with an almost circular whorl section and four rectiradiate ribs in a distance equal to the whorl height. It closely approaches the present material, but its very much larger size (septate at 23 mm whorl height) is distinctive. Occurrence Uppermost Albian of Angola only. Family Hamitidae Hyatt, 1900 Genus Hamites Parkinson, 1811 Type species Hamites attenuatus J. Sowerby, 1814 Discussion Recent discussions of the family are to be found in Wiedmann & Dieni (1968) and Klinger (1976), and further discussion on the material available here is unnecessary. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 227 Hamites virgulatus Brongniart, 1822 Figs 16F, H, 32B—C Hamites virgulatus Brongniart (in Cuvier & Brongniart), 1822: pl. 0 (fig. 6). Hamites (Stomohamites) virgulatus (Brongniart ?) Pictet & Campiche, Spath, 1941: 635, pl. 71 (figs 7-10), pl. 72 (fig. 11), fig. 230 (with synonymy). Renz, 1968: 65, pl. 11 (figs 9-11), fig. 23b-d (with synonymy). Hamites (Stomohamites) subvirgulatus Spath, 1941: 645, fig. 234. Renz, 1968: 66, pl. 11 (figs 13-14), figs 23e, 24a (with synonymy). Stomohamites brongniarti Breistroffer, 1947: 77. Hamites (Hamites) virgulatus Brongniart, Wiedmann & Dieni, 1968: 53, pl. 5 (figs 1-2, 10), pl. 7 (figs 1-2), figs 21-27. Klinger, 1976: 60, pl. 23 (figs 4-5, 8), figs 8 I,n—o, 11i (with synonymy). Hamites venetzianus Pictet (in Pictet & Roux), 1847: 134, pl. 14 (fig. 6). Hamites (Stomohamites) venetzianus Pictet, Spath, 1941: 638, pl. 71 (figs 11-13), fig. 231. Renz, 1968: 67, pl. 11 (figs 15-16), figs 23f, 24b (with synonymy). Material Ten composite internal moulds, SAM-—PCA2959, 2963-64, 2966, 2971, 3118, 3157-58, and 3371 from the upper part of the dispar Zone at Praia-Egito, three fragments retaining recrystallized tests from Porto Amboim, USNMNH 236955-7, and SAM-PCA4603 from Cabo Ledo. Description The whorl section is elliptical compressed, with three to six prorsiradiate ribs in a distance equal to the whorl height. The ribs are effaced across the dorsum. Discussion The writers follow Wiedmann & Dieni (1968) in regarding H. brongniarti, H. venetzianus, and H. subvirgulatus as synonyms of H. virgulatus, although there may be some justification for retaining venetzianus at the varietal level for those variants in which the ribs are as thick as, or thicker than, the interspaces. Hamites duplicatus Pictet & Campiche, 1861 Figs 16D, 32A Hamites virgulatus Pictet (non Brongniart) (in Pictet & Roux), 1847: 391, pl. 14 (figs 7, 9 only). Hamites duplicatus Pictet & Campiche, 1861: 98. Hamites (Stomohamites) duplicatus Pictet & Campiche, Spath, 1941: 640, pl. 72 (figs 12-16), fig. 232. Renz, 1968: 68, pl. 11 (figs 19-21), fig. 23h-k. Material A single composite internal mould from Praia-Egito, SAM-—PCA2955. Description The whorl section is oval, compressed, with a somewhat flattened dorsum. Ornament comprises fine, rursiradiate ribs, seven in a distance equal to the whorl height, which are very weak across the dorsum. 228 ANNALS OF THE SOUTH AFRICAN MUSEUM Discussion The writers are not convinced by Wiedmann & Dieni’s (1968) argument for the inclusion of H. duplicatus in the synonymy of H. virgulatus and, for the present, maintain it as distinct. Occurrence Hamites duplicatus is known from the Upper Albian of England, France, Switzerland, and Angola. Superfamily TURRILITACEAE Meek, 1876 Family Turrilitidae Meek, 1876 Subfamily Turrilitinae Meek, 1876 Genus Mariella Nowak, 1916 Type species Turrilites bergeri Brongniart, 1822 Discussion Kennedy (1971) and Klinger & Kennedy (1978) have recently provided a comprehensive discussion of the taxonomic standing of Mariella, including in it three subgenera, viz. M. (Mariella), M. (Plesioturrilites) and M. (Wintonia). As noted by Clark (1965: 49), however, M. (Wintonia) and M. (Plesio- turrilites) differ only in that the former has an early, straight shaft which then passes into the helical coiling of M. (Plesioturrilites), although specimens of M. (Wintonia) graysonensis (Adkins), the only known species, cannot be distinguished from M. (Plesioturrilites) bosquensis (Adkins) in the absence of this straight shaft. Klinger & Kennedy (1978) suggest the ‘shaft’ is an artefact of preservation and treat Plesioturrilites as a synonym of Wintonia. Mariella (Mariella) circumtaeniatus (Kossmat, 1895) Figs 31E-F, 39H Turrilites gresslyi Stoliczka (non Pictet & Campiche), 1865: 186, pl. 87 (figs 1-5, ? non 2). Turrilites circumtaeniatus Kossmat, 1895: 141, pl. 18 (figs 4-5); Boule, Lemoine & Thévenin, 1907: 57, pl. 13 (fig. 4). Non Turrilites circumtaeniatus Scott (non Kossmat), 1926: 145, pl. 1 (figs 10-11) (= M. worthensis). Paraturrilites aff. circumtaeniatus (Kossmat) Collignon, 1963: 46, pl. 258 (fig. 1120). Non Turrilites circumtaeniatus Kossmat, Woods, 1917: 11, pl. 5 (figs 2-3) (= M. thomsoni Henderson). Material A single specimen USNMNH 236958, with recrystallized shell preserved, from Porto Amboim. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 229 @yonc Fig. 32. A. Hamites duplicatus Pictet & Campiche. Whorl section of SAM-PCA2955. B-C. Hamites virgulatus Brongniart. Whorl sections. B. SAM- PCA2959. C. SAM-PCA3158. x1. Description Coiling sinistral, with a very acute apical angle (approximately 20°). Flanks rounded intercostally, with two rows of prominent, rounded tubercles and a third hidden in the spiral seam. There are nine to eleven tubercles per half-whorl situated on very weak, slightly oblique ribs. The upper row of tubercles is situated slightly above mid-flank, and the central row midway between the upper row and the lower spiral suture. Ribbing is conspicuous only on the area between the upper row of tubercles and the upper spiral suture, where rather fine ribs commonly connect in pairs to the upper row of tubercles, frequently with an intercalated rib between tubercles. Discussion Amongst contemporaneous species of Mariella, M. (M.) cantabrigiensis (Jukes-Browne) (Spath 1937: 518, pl. 57 (fig. 36), pl. 58 (figs 1-2), figs 18la—b, 182d-e) approaches the present species most closely. It differs in its much larger spiral angle (30° or more) and in lacking the fine looped ribs on the upper, outer face of the whorls. M. (Mariella) nobilis (Jukes-Browne) (Spath 1937: 520, pl. 58 (figs 10-11), fig. 182) differs in having more prominent single ribs and subdued tuberculation. M. (Mariella) bergeri (Brongniart) (1822: 395, pl. 7 (fig. 3)), of which M. (M.) miliaris (Pictet & Campiche) (1861: 136, pl. 51 (fig. 5)) may be only an extreme variant, has four rows of tubercles exposed on the outer face of the whorls. M. (Mariella) dubourdieui (Collignon) (1963: 46, pl. 285 (fig. 1121)) resembles M. (M.) circumtaeniatus in the possession of looped ribs, but also has four rows of tubercles exposed on the outer face of the whorls. M. (Mariella) worthensis (Adkins & Winton) (1920: 44, pl. 7 (figs 10-11, 13)) from the uppermost Albian Pawpaw Formation of Texas, closely resembles M. (M.) circumtaeniatus, but appears to lack the fine looped ribs of Kossmat’s species, as does M. (M.) hillyi (Dubourdieu) (1953: 46, pl. 4 (figs 1-3)). Occurrence This species is known from the uppermost Albian of India, Madagascar, Zululand, Angola, and New Zealand. 230 ANNALS OF THE SOUTH AFRICAN MUSEUM Mariella (Mariella) nobilis (Jukes-Browne, 1877) Fig. 33A-B Turrilites intermedius Pictet & Campiche, 1861: 127, pl. 57 (fig. 15 only). Turrilites escherianus Pictet & Campiche (von Pictet), 1861: 130, pl. 56 (figs 6-8 only). Turrilites nobilis Jukes-Browne, 1877: 493, pl. 21 (fig. 1). Mariella nobilis (Sukes-Browne) Breistroffer (in Besairie), 1936: 147. Spath, 1937: 520, pl. 58 (figs 10-11), fig. 182a—c. Breistroffer, 1940: 147. Clark, 1965: 40, pl. 10 (figs 2-4), pl. 11 (figs 4-5). Renz, 1968: 89, pl. 17 (figs 28, 40), figs 311, 32k. Mariella nobilis var. cruciana Breistroffer (in Besairie), 1936: 148. Spath, 1937: 521. Breistroffer, 1940: 148. Renz, 1968: 90, pl. 17 (fig. 34), figs 31b, 32d. Mariella aff. nobilis (Jukes-Browne) Spath, 1937: 520, pl. 58 (fig. 21). Paraturrilites nobilis (Sukes-Browne) Breistroffer, 1947: 60; 1953: 1350. Paraturrilites nobilis var. cruciana (Breistroffer) Breistroffer, 1947: 60. Paraturrilites nobilis var. brownei Breistroffer, 1947: 96. Mariella nobilis brownei (Breistroffer) Renz, 1968: 90, pl. 17 (figs 35-36, 38-39), figs 31i, 32e. Material Two specimens, SAM-—6531 (from an unknown location on the Angolan littoral), and a single fragment in the Washburn collection, USNMNH 237019 from Porto Amboim. Description These two fragments of M. (Mariella) bear prominent oblique ribs, narrower than the interspaces, estimated at totalling twenty-six per whorl. They are ornamented by three rows of weak tubercles; the tubercles of the adoral row are weakly clavate, whereas those of the other two rows are weakly rounded. Discussion M. (Mariella) nobilis var. cruciana Breistroffer (in Besairie 1936: 148) was separated from the typical form by its denser ribbing (30 ribs per whorl, as against 26-28) and more prominent tubercles. M. (Mariella) nobilis var. brownei Breistroffer (1947: 96) was distinguished by its sparser ribbing (24-26 per whorl) and weaker tuberculation. As noted by Spath (1937: 521), M. (Mariella) escheriana (Pictet) (in Pictet & Roux 1847: 154, pl. 15 (fig. 11)) closely resembles the present species and there even appear to be intermediates. Pictet’s species was distinguished by its denser ribbing (35-40 ribs per whorl), and flattened flanks with only two rows of tubercles. The closest species to M. (M.) nobilis is M. (M.) cantabrigiensis (Jukes- Browne) (Spath 1937: 518, pl. 57 (fig. 36), pl. 58 (figs 1-2), figs 18la—b, 182d-e) from the late Albian of western Europe. Not only are they strictly contem- poraneous, but Spath (1937: 519) also admits to the presence of intermediates. Typically, however, it differs in its sparser ribbing and more prominent tubercles. M. (Mariella) worthensis (Adkins & Winton) (1920: 44, pl. 7 (figs 10-11, 13)) is very close to M. (M.) nobilis but is generally more coarsely ribbed (14-28 ribs per whorl) with four rows of tubercles per whorl. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 231 M. (Mariella) gresslyi, M. (M.) cantabrigiensis, M. (M.) nobilis, M. (M.) escheriana, M. (M.) worthensis and M. (M.) hillyi are a contemporaneous group of very closely allied species whose intraspecific variation clearly needs documenting before the true taxonomic status of the species involved can be resolved. Occurrence M. (Mariella) nobilis is known only from the upper Upper Albian of southern England, Texas, and Angola. Fig. 33. A-B. Mariella (Mariella) nobilis (Jukes-Browne), SAM-6531. C-D. Puzosia (Puzosia) sp., SAM-6407. x1. Mariella gresslyi (Pictet & Campiche, 1861) Fig. 16G Turrilites gresslyi Pictet & Campiche, 1861: 132, pl. 57 (figs 11-13). Neumayr, 1875a: 901. Renevier, 1890: 339. Pervinquiére, 1910: 54. Bose, 1923: 147. Diener, 1925: 83. Collignon, 1929: 65, pl. 1 (fig. 15). Turrilitoides (2) gresslyi (Pictet & Campiche) Breistroffer, 1936: 65. Mariella gresslyi (Pictet & Campiche) Spath, 1937: 516, pl. 58 (figs 3-4), fig. 180. Breistroffer, 1940: 149. Renz, 1968: 89, pl. 17 (figs 30a—b, 32-33), figs 31c, 32a—b. Paraturrilites gresslyi (Pictet & Campiche) Breistroffer, 1947: 60. Collignon, 1963: 47, pl. 258 (fig. 1122). ? Turrilites gresslyi Pictet & Campiche, Boule, Lemoine & Thévenin, 1907: 39, pl. 6 (fig. 2). Mariella gresslyi bifurcata Renz, 1968: 89, pl. 17 (fig. 42a—b), figs 3le, 32c. ? Turrilites cantabrigiensis Jukes-Browne, 1877: 493. ? Mariella cantabrigiensis (Jukes-Browne) Spath, 1937: 518, pl. 57 (fig. 36), pl. 58 (figs 1-2), figs 18la—b, 182d-e. 232 ANNALS OF THE SOUTH AFRICAN MUSEUM Material A single specimen, SAM-—PCA3133, from low down in the coastal cliffs immediately north of the estuary at Praia-Egito, and thus from a level below the main Stoliczkaia occurrence. Description The specimen is rather poorly preserved, but retains recrystallized test. The shell is sinistrally coiled, with a moderately large apical angle. The outer faces of the whorls are gently convex intercostally, and angular, polygonal costally. Three rows of tubercles are visible on rather oblique ribs. The upper row of tubercles are conical whereas the middle row is distinctly clavate. The nature of the tubercles of the lower row was difficult to discern, but they, too, appear to be clavate. Discussion The features of the present specimen are those of M. gresslyi and the authors have no hesitation in assigning the Angolan example to this species. Mariella cantabrigiensis (Jukes-Browne) (Spath, 1937: 518, pl. 57 (fig. 36), pl. 58 (figs 1-2), figs 180h, 18la—b, 182d-e) closely approaches M. gresslyi, but was said to differ in having all tubercles rounded, whereas the lower two rows are clavate in M. gresslyi. However, the two species are strictly contemporaneous and occur side by side, whilst Spath (1937) admits to transitions between the two. Population studies may show M. cantabrigiensis to be an intraspecific variant of M. gresslyi. Mariella gresslyi bifurcata Renz (1968: 89, pl. 17 (fig. 42), figs 3le, 32c) is based on a single specimen which shows the ribs on the base of the spire bifurcating from the lowest (adoral) row of tubercles. Mariella nobilis (Jukes-Browne) (Spath 1937: 520, pl. 58 (figs 10-11), figs 18lc, 182a—c) differs from the present species in having very subdued tubercles, with ribs dominant. Occurrence Mariella gresslyi is currently known from southern England, France, Switzerland, Algeria, Angola, Zululand, and Madagascar. Mariella cf. oehlerti (Pervinquiére, 1910) Figs 3H, 34 Compare Turrilites gresslyi Boule, Lemoine & Thévenin (non Pictet & Campiche), 1907: 57, pl. 13 (fig. 2). Collignon, 1929: 65, pl. 6 (fig. 15). ? Turrilites cf. gresslyi Spath (non Pictet & Campiche), 1921: 289. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 233 Turrilites oehlerti Pervinquiére, 1910: 53, pl. 5 (figs 14-17). Collignon, 1929: 65, pl. 6 (figs 16-17); 1964: 15, pl. 320 (figs 1398-1399). ? Mariella malgachensis Breistroffer, 1940: 79. ? Turrilites bergeri Choffat (non Brongniart), 1903: 15, pl. 1 (figs 4-6). ? Mariella bergeri var. conduciensis Breistroffer, 1940: 149. Mariella (Mariella) oehlerti (Pervinquiére), Férster 1975: 190, pl. 7 (figs 7-8), fig. 52. Klinger & Kennedy 1978: 31, pl. 3 (fig. E), pl. 4 (fig. E), pl. 6 (figs H, N), pl. 7 (fig. G), pl. 8 (figs G-H), figs la—b, 7b, d, 8g. Material A single composite internal mould, SAM-—PCA4798, from the Quissama Ridge at Cabo Ledo. Fig. 34. Mariella (Mariella) cf. oehlerti (Pervinquiére), SAM-PCA4798. x1. Description The shell is a high-spired, sinistrally-coiled turriliticone, with a spiral angle of about 22°. The whorls are just touching with the outer face gently convex, and somewhat flattened. The adapical shoulder is abrupt and subangular (Fig. 3H) whilst the adoral shoulder is evenly rounded. The outer face is ornamented with 234 ANNALS OF THE SOUTH AFRICAN MUSEUM four rows of prominent tubercles arranged on weak, oblique ribs, of which there are about thirty per whorl. The upper row of tubercles is the most prominent, and they are slightly bullate and situated slightly above mid-whorl. The tubercles of the two middle rows are conical and are separated by a weak spiral groove. The tubercles of the lower row are the smallest and are situated in the whorl seam. On the adoral face of the final whorl there are prominent ribs. The spacing of the tubercles is subequal on the penultimate whorl of the present specimen, but on the final whorl the middle two rows are distinctly closer together than the others. Discussion At present the species M. gresslyi—M. cantabrigiensis—M. oehlerti— M. circumtaeniatus are not well differentiated. Mariella gresslyi (Pictet & Campiche) (1861: 132, pl. 57 (figs 11-13)) is typically an uppermost Albian species which differs from the present form and M. oehlerti in that the rows of tubercles are typically clavate. However, Spath (1937: 519) records passage form between M. gresslyi and the contemporaneous M. cantabrigiensis (Jukes-Browne) (Spath 1937: 518, pl. 51 (fig. 36), pl. 58 (figs 1-2)), the latter distinguished from Pictet & Campiche’s species in having rounded tubercles of equal size, and thus very close to M. oehlerti and the Angolan material. Since, however, the present specimen is much larger than known material of M. gresslyi and M. cantabrigiensis, differences may be due to a comparison of different ontogenetic stages. Clearly, however, this problem cannot be resolved in the present paper. Mariella circumtaeniatus has only three rows of tubercles, generally with conspicuous looping of the ribs on the adapical shoulder of the whorls. Mariella dorsetensis (Spath) (= Turrilites bergeri Sharpe (non Brongniart) 1857: 65, pl. 26 (fig. 11 only)) differs from the present material in having fewer ribs per whorl and in having the third and fourth (adoral) rows of tubercles approximated, whilst ribs are absent on the base of the spire. Mariella gallienii (Boule, Lemoine & Thévenin) (1906: 60, pl. 14 (figs 5—6)) differs from the present specimen in that the tubercles of the four rows are distinctly bullate. Occurrence Mariella oehlerti is known from the Lower Cenomanian of Algeria, Madagascar, Mozambique, Zululand, and possibly Japan, and possibly the uppermost Albian of Angola and Switzerland. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 235 Suborder AMMONITINA Hyatt, 1889 Superfamily HOPLITACEAE Douvillé, 1890 Family Desmoceratidae Zittel, 1895 Subfamily Puzosiinae Spath, 1922 Genus Puzosia Bayle, 1878 Subgenus Puzosia Bayle, 1878 Type species Ammonites planulatus J. de C. Sowerby, 1827 Puzosia (Puzosia) sp. Fig. 33C-D Material A single crushed specimen, SAM-6407, retaining recrystallized shell, from Porto Amboim. Description The shell is compressed, moderately involute, with about 60 per cent of the preceding whorl covered by the outer whorl. The umbilicus is moderately narrow (22% of the diameter), with steep umbilical walls and evenly rounded umbilical shoulders. The whorl section is oval, compressed (W/H = 0,83) with an evenly rounded venter. There are an estimated five constrictions per half- whorl, preceded by a strong ventral rib. Flank and venter also bear faint, ill-preserved, fine ribs. Measurements No. D H W W/H U SAM-6407 a7 18(49) 15(41) 0,83 +8(22) Discussion The specimen cannot be usefully compared with the large number of late Albian puzosiids known, although if the estimated number of constrictions is correct, it must approach forms such as P. crebrisulcata Kossmat (1898: 116, pl. 17 (fig. 4), pl. 18 (fig. 2) and P. malandiandrensis Collignon (1963: 66, pl. 265 (fig. 1156)). Puzosia (Puzosia) cf. sharpei Spath, 1923 Fig. 35 Compare Ammonites planulatus Sharpe (non Sowerby), 1854: 29, pl. 12 (fig. 4 only). Puzosia sharpei Spath, 1923: 46, pl. 1 (figs 11-12), fig. 11b. Renz, 1968: 21, pl. 1 (figs 4, 8), figs 6b, 7e (with synonymy). Material A single fragment, SAM-PCA3141, preserved as an internal mould, from the dispar Zone of Praia-Egito. 236 ANNALS OF THE SOUTH AFRICAN MUSEUM Description The single specimen is a fragment of about one-quarter whorl. The shell was moderately evolute, with a fairly wide, shallow umbilicus and evenly rounded umbilical shoulders. The flanks are broad, subparallel and only slightly convex, with a broadly rounded venter. There are two deep constrictions on the fragment, separated along the venter by a distance slightly greater than the whorl height. The constrictions are initially prorsiradiate, but soon recurve before flexing strongly forwards to form a chevron across the venter. Discussion The Angolan material differs from P. sharpei in that the constrictions do not show as strong a geniculation, and are therefore not as strongly falcate. Wiedmann & Dieni (1968) included this species in the synonymy of Puzosia provincialis (Parona & Bonarelli), a lower Middle Albian species. The material figured by these authors (1968) as P. provincialis shows relict lappet structures and are, therefore, microconchs. The holotype of P. sharpei, on the other hand, is still septate at 83 mm diameter and appears to be a macroconch. Consequently, the authors regard the inclusion of P. sharpei into the synonymy of P. provincialis as premature, and prefer to maintain Spath’s species as distinct, for the time being. Occurrence Puzosia sharpei is known with certainty only from southern England, France, Switzerland, and possibly Angola, where it is typical of the dispar Zone. Fig. 35. Puzosia (Puzosia) cf. sharpei Spath. Lateral and ventral views of SAM-PCA3141. x1. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 237 Fig. 36. Desmoceras (Desmoceras) latidorsatum (D’Orbigny). The original of D’Orbigny’s (1841, pl. 80 (fig. 5)) figured specimen from the Middle Albian of France, in the D’Orbigny collection, Natural History Museum, Paris. x1. Subfamily Desmoceratinae Zittel, 1895 Genus Desmoceras Zittel, 1884 Subgenus Desmoceras Zittel, 1884 Type species Ammonites latidorsatus Michelin, 1838 Desmoceras (Desmoceras) latidorsatum perinflatum subsp. nov. Figs 37-38, 39D-F Desmoceras latidorsatum var. inflata Breistroffer, Haas, 1952: 2, figs 1, 3-10. Desmoceras reynesianum Haas, 1952: 4, figs 2, 11-13. Material Twenty-one specimens, SAM-6414, SAM-PCA2931, 2934, 2968, 3170 and 3172 from Egito, preserved as composite internal moulds, and USNMNH 236961-75 from Porto Amboim with recrystallized shell generally preserved. Type material USNMNH 236970 from Porto Amboim is designated holotype; the remaining specimens are paratypes. 238 ANNALS OF THE SOUTH AFRICAN MUSEUM Etymology From the Latin, per—exceedingly, very much, inflatus—swollen; applying to the strongly inflated shell form. Diagnosis A rather small, late Albian (dispar Zone) subspecies of D. latidorsatum in which the majority of the population comprises strongly inflated individuals (“/H = 1,10-1,50) which correspond to the D. latidorsatum var. inflatum of previous workers. Description Very involute, cadicone, with a narrow, crater-like umbilicus (16-20% of diameter). Umbilical walls steep, with gently rounded shoulders. Flanks flattened (USNMNH 236972) to strongly convex (USNMNH 236964), with maximum width just below mid-flank. Venter broadly rounded. Shell smooth, K Fig. 37. Desmoceras (Desmoceras) latidorsatum perinflatum subsp. nov. A-B. USNMNH 236966. C-E. The holotype USNMNH 236970. F-G. USNMNH 236973. H-J. USNMNH 236768. K-L. USNMNH 236967. M-O. USNMNH 236964. x1. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 239 except for weak constrictions on some specimens (USNMNH 236970). Where present, constrictions begin at the umbilical seam and pass strongly forwards to the umbilical shoulder where they recurve strongly, flexing forwards again just above mid-flank to form a prominent U-shaped tongue across the venter. Measurements No. D H W W/H U USNMNH 236961 21 +9(43) +12(57) 1,33 +4(19) USNMNH 236962 14 7(50) S561) . “1,23 2,3(16) USNMNH 236964 27 13(48) 18,3(68) 1,41 +5(18) USNMNH 236965 21 E5(S4) 14(67) 122 2 USNMNH 236966 23 9,5(41) 13(56) 137 +4,5(20) USNMNH 236967 20 10,2(51) 12(60) 1,18 +4(20) USNMNH 236968 12,3 6,2(50) 8,3(67) 1,34 +2(16) USNMNH 236969 42 22(52) +24(57) 1,09 +8(9) USNMNH 236970 24 11(46) 15(63) 1,36 4,5(19) USNMNH 236971 14,6 7,5(51) 9(62) E20 rs USNMNH 236972 15 6(40) 8,5(57) 1,42 2,7(18) = 10,5 5(48) 6,5(62) 1,30 2,0(19) USNMNH 236973 13 +6(46) 8,1(62) 1,35 ? USNMNH 236975 +14 6,5(46) 8,2(59) 1,26 2,2(16) Intraspecific variation The wide range of intraspecific variation shown by D. (Desmoceras) latidorsatum has long been recognized (D’Orbigny 1941, Pictet (in Pictet & Roux) 1848, Kossmat 1897, Jacob 1908, Fallot 1910, Spath 1923, Wiedmann & Dieni 1968) and it would appear that species separation is possible only at the population level since gross intraspecific variation exceeds subtle differences between successive faunas. Consequently, the following names, which are used both at the varietal and subspecific levels by various authors, are based merely upon individuals within these populations and are of no taxonomic value: media (Jacob), complanata (Jacob), inflata (Breistroffer), obesa (Reynés), petkovici Breistroffer, Jongesulcata (Collignon) and angusteumbilicata Haas. Jacob (1908) studied the intraspecific variation within Desmoceras latidor- satum from the Balme de Rencurel, a fauna contemporaneous with D’Orbigny’s (1841) material (Fig. 36) from the ‘Argile 4 Hoplites dentatus Sow.’ of Aube, and of early Middle Albian age. Within this assemblage, Jacob (1908) recognized the typical form, as well as his varieties media and complanata which are more compressed than typical D. Jatidorsatum. In contrast, the present collection, together with that described by Haas (1952) shows that inflated variants, normally assigned to the variety inflata Breistroffer, form the vast majority of the populations from the S. dispar Zone of Angola (Fig. 38), whereas compressed 240 ANNALS OF THE SOUTH AFRICAN MUSEUM 0 ater a — 0 40 SME = a Fig. 38. Plot of inflation and umbilical ratio against diameter for Desmoceras latidorsatum perinflatum subsp. nov. Dots = present Angolan material; open triangles = material described by Haas (1952), and Wiedmann & Dieni (1968); black triangle = Desmoceras reynesianum (after Haas 1952); square = Desmoceras latidorsatum latidorsatum (Michelin), (after D’Orbigny 1841). UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 241 forms are entirely absent. It would seem, therefore, that there has been a genuine shift within the population structure of D. latidorsatum through time, from predominantly compressed individuals in the Middle Albian to predominantly inflated variants in the uppermost Albian. The observation that certain individuals throughout this range can be assigned to Desmoceras latidorsatum s.s. suggests that the differences are not of specific importance. Since the Angolan population comprises mainly strongly inflated individuals (W/H = 1,10-1,50), the differences are sufficient for subspecific separation and the entire Angolan population (it is the characters of the population which define the subspecies) is assigned to D. latidorsatum perinflatum subsp. nov. Desmoceras reynesianum Haas (1952: 4, figs 2, 11-13) was proposed to replace Ammonites obesus Reynés (non Stoliczka) and is characterized by its extreme inflation. The Angolan material assigned to this species, however, differs in being of uppermost Albian age and, as can be seen from Figure 38, merely represents extreme variants within the present population, and consequently Haas’s (1952) material is included in the synonymy of D. latidorsatum perinflatum subsp. nov. Discussion Population studies will probably show that D. collignoni Breistroffer (in Besairie 1936: 170, pl. 16 (fig. 2), fig. 10d), D. inane (Stoliczka) (1865: 121, pl. 59 (figs 13-14)) and D. chirichense (Pervinquiére) (1907: 152, pl. 6 (figs 17-20)) do not bear separation from D. /atidorsatum. Desmoceras barryae Anderson (1958: 214, pl. 12 (fig. 2)) and D. merriami Anderson (1902: 103, pl. 6 (figs 135-138)), which was treated as a variety of D. latidorsatum by Breistroffer (1947: 61), are probably better referred to the subgenus Pseudouhligella. Occurrence Desmoceras latidorsatum perinflatum subsp. nov. is currently known with certainty only from Angola. Superfamily ACANTHOCERATACEAE Hyatt, 1900 Family Lyelliceratidae Spath, 1921 Discussion Wright (in Arkell et al. 1957) included the following genera within the Lyelliceratidae—Prolyelliceras, Lyelliceras, Tegoceras, Neophlycticeras, Stolicz- kaia (with Faraudiella as a subgenus), Budaiceras and Salaziceras. More recently, Casey (1965) introduced the subgenus Stoliczkaia (Villoutreysia), and proposed the new genus Paradolphia for forms from the S. dispar Zone of southern England said to te transitional between Stoliczkaia and Forbesiceras. Matsumoto & Inoma (1975: 277) have also introduced the subgenus Stoliczkaia (Shumarinaia). 242 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 39. A. Stoliczkaia (Stoliczkaia) tenuis Renz. Lateral view of SAM—PCAS5478. B-C. Stoliczkaia (Stoliczkaia) sp. A fragmentary internal mould, SAM—PCA4805, from Cabo Ledo. D-F. Desmoceras latidorsatum perinflatum subsp. noy. D-E. Ventral and lateral views of SAM-PCA2934. F. Lateral view of SAM—PCA2931. G. Mortoniceras (Angolaites) gregoryi (Spath). Lateral view of a crushed fragment, SAM—PCA3168. H. Mariella aff. circumtaeniatus (Kossmat). SAM—PCA3130, from low in the sea cliffs (lower part of dispar Zone) at Praia-Egito. eile UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 243 Amedro (1976) has recently suggested that Paradolphia is better regarded as a subgenus of Neophylicticeras (along with Protissotia and Eotropitoides), whilst there is a case for regarding both Paracalycoceras and Cottreauites, both of which derive from Stoliczkaia, as Lyelliceratidae rather than Acanthoceratidae. The collection of Stoliczkaia to be discussed below shows a remarkable similarity to the earliest Forbesiceras, i.e. the beaumontianum-largilliertianum group (compare Juignet & Kennedy 1977, pl. 6 (fig. 1)), and both from a stratigraphic and morphological point of view is most likely to have provided the ancestor to Forbesiceras. The close similarity between Forbesiceras and Stoliczkaia of the dispar—clavigera group suggests that the monogeneric sub- family Forbesiceratinae is superfluous, and that Forbesiceras should be trans- ferred to the Lyelliceratidae. A study of the early ontogenetic stages of Forbesiceras has led Casey (1965) to suggest that Neopulchellia (Collignon 1929) was based upon pyritic nuclei of Forbesiceras. Genus Stoliczkaia Neumayr, 1875 Type species Ammonites dispar d’Orbigny, 1841 Discussion Stoliczkaia occupies a key position in the evolution of the mid-Cretaceous Acanthocerataceae, as it appears to be the origin of both the Mantelliceratinae and Acanthoceratinae which in turn gave rise to the remaining Upper Cretaceous acanthoceratids. More than a score of specific names have been applied to the genus, but there has been no sound account of intraspecific variation, nor of the apparent dimorphism present, some subgenera and species reaching a large size and becoming feebly ribbed at maturity (e.g. S. (S.) dispar), others remaining small with strong ribs throughout (S. (Shumarinaia)). Furthermore, because of their transitional position between Lyelliceratidae, Acanthoceratinae and Mantelli- ceratinae, there are a number of forms whose position is equivocal. Indeed, the authors find themselves in disagreement over the precise position of some of these passage forms. The subgenera of Stoliczkaia are as follows: 1. Stoliczkaia (Stoliczkaia) (type species Ammonites dispar d’Orbigny). Typically rather involute, compressed ammonites with straight or slightly curved primary ribs, with shorter intercalated ribs between. Primary ribs may be weakly bullate, and during early growth stages bear ventrolateral tubercles or clavi, whilst the venter may be flat or slightly raised. In middle growth tuberculation disappears and ribs extend across a rounded venter; at maturity ribs broaden, become irregular, and may become effaced on the adult body chamber. Typical repre- sentatives of the type species S. (S.) dispar are shown as Figures 40 and 41. 2. Stoliczkaia (Faraudiella) (type species Ammonites blancheti Pictet & Campiche). Small Stoliczkaia in which distinct siphonal, and sometimes ventro- 244 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 40. Stoliczkaia (Stoliczkaia) dispar (d’Orbigny). The holotype, Renaux collection, Faculté des Sciences, Montpellier, from the Upper Albian of Ventoux, Vauceuse, France. Slightly reduced. lateral tubercles persist on to the body chamber. Typical representatives are shown in Figure 42. 3. Stoliczkaia (Shumarinaia) (type species S. (Shumarinaia) hashimotoi Matsumoto & Inoma). Small, with simple suture line and coarse ribbing throughout. A fourth subgenus, Villoutreysia was proposed by Casey (1965: 435, fig. 161; type species S. (V.) villoutreysi Casey) (Fig. 43) for what he described as Hypacanthoplites homoeomorphs diagnosed as ‘Stoliczkaia with broad, square venter and strong persistent ribbing, differing from Mantelliceras in much UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 245 Fig. 41. Stoliczkaia (Stoliczkaia) dispar (d’Orbigny). Juvenile specimen in C. W. Wright collection WW 72344, from the dispar Zone Ammonite Bed of the Dorset Coast. x1. A B c D Fig. 42. Stoliczkaia (Faraudiella) sexangulata (Seeley). Seeley’s original specimen, Sedgwick Museum, Cambridge, B53, from the Cambridge Greensand, Cambridge. 1. 246 ANNALS OF THE SOUTH AFRICAN MUSEUM earlier loss of ventral tubercles, squarer whorls and, generally, narrow and shallower umbilicus’. The holotype of S. (V.) villoutreysi is illustrated here as Figure 43; after an examination of the large collections of Stoliczkaia in the Paris Museums the authors have concluded that it is not separable subgenerically from Stoliczkaia sensu stricto. The genus Paradolphia Casey was proposed (Casey 1965: 461, pl. 77 (figs 5—6)) for the type species P. prisca Casey (illustrated here as Fig. 44) for intermediates between Stoliczkaia and Forbesiceras but, as noted above, is possibly a subgenus of Neophlycticeras. Paracalycoceras (type species Ammonites wiestii Sharpe 1857: 47, pl. 21 (fig. 3)) is an enigmatic genus from the Lower Cenomanian of southern England, known with certainty only from two specimens—the lost holotype (fide Kennedy 1971) and an extant specimen in the collections of C. W. and E. V. Wright (Fig. 45). Kennedy (1971: 79) diagnosed the genus as follows: ‘Medium-sized, somewhat involute ammonites. Inner whorls slightly compressed, with long ribs bearing umbilical bullae, and lower and upper ventrolateral tubercles separated by 1, 2, or sometimes more shorter ribs. There is a distinctly raised siphonal area, and an incipient siphonal tubercle on all ribs. Outer whorl with a broad venter, and broad, distant, flexuous rursiradiate ribs, irregularly long and short.’ Both morphologically and in the observed ontogenetic changes, Paracaly- A B c Fig. 43. Stoliczkaia (Stoliczkaia) villoutreysi Casey. Holotype, O. de Villoutreys collection, Uppermost Albian, Monte Carlo Tunnel, Monte Carlo. x1. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 247 Fig. 44. Neophylicticeras (Paradolphia) prisca (Casey). A—C. Holotype, Norwich Castle Museum 61.18(1679). D-E. Paratype, Sedgwick Museum, Cambridge, B93303. Both from the Cambridge Greensand, Cambridge. A-C x1, D-E x2. coceras is virtually indistinguishable from certain species of Stoliczkaia, from which it is obviously descended. Little more can be said until topotype material is studied, but it may prove more satisfactory to regard it as a subgenus of Stoliczkaia. Kennedy (1971: 80) considered that the genus Coftreauites (Collignon 1929), based upon pyritic nuclei, ‘may be wholly or partly a synonym of Paracalycoceras’. Again, this question cannot be resolved until Paracalycoceras is better known or mature Cottreauites are described. It seems very likely, however, that some Coftreauites are juveniles of either Stoliczkaia or Paracaly- coceras, and that it should be considered a nomen dubium. 248 ANNALS OF THE SOUTH AFRICAN MUSEUM C D Fig. 45. Stoliczkaia (Paracalycoceras) wiestii (Sharpe). C. W. Wright collection 3556, Lower Cenomanian, Cenomanian Limestone, Bed A2, White Cliff, Seaton, Devon. x1. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 249 Subgenus Stoliczkaia (Stoliczkaia) Neumayr, 1875 Renz (1968: 46) recognized four species groups within Stoliczkaia sensu stricto: 1. The group of S. dispar with ventrolateral tubercles only on the inner whorls and weak ribbing on the body chamber, comprising S. dispar (d’Orbigny), S. dorsetensis Spath and S. tenuis Renz. 2. The group of S. africana which retains ventrolateral tubercles on to the body chamber, with well-developed, broad, falcate ribs, and includes S. africana (Pervinquiére) and S. flexicostata Breistroffer. Matsumoto & Inoma (1975) have proposed the subgenus Shumarinaia for this group. 3. The group of S. notha, which lacks ventrolateral tubercles whilst retaining strong ribbing on to the body chamber, and comprises S. notha (Seeley) and S. clavigera Neumayr. 4. The group of S. /evis which lacks ventrolateral tubercles and is virtually without ornament. Only S. /evis Renz, based on a unique holotype, is assigned to this group. Stoliczkaia (Stoliczkaia) tenuis Renz, 1968 Figs 46-53, 54A-F, 55, 68E Stoliczkaia tenuis Renz 1968: 48, pl. 6 (figs 6, 12), fig. 16b, f. Material 37 specimens, USNMNH 236981-237012a—b, 237014-5, together with 3 specimens in the collections of the South African Museum, SAM-—PCA5477-78 and 6811, all retaining part or all of their recrystallized shell, from Porto Amboim, and 7 specimens, SAM-—PCA2938-39, 2944, 3169, 3373 and 5475-5476, preserved as composite internal moulds from Egito. Description Up to 20 mm diameter: shell compressed (“/H = 0,50-0,63), very involute with a deep, narrow umbilicus (11-17% of diameter). Umbilical walls steep, almost vertical, with evenly rounded umbilical shoulders. Flanks broad, slightly convex to flat, with maximum width below mid-flank, converging towards the narrow venter. Venter slightly convex to almost tabulate, and weakly raised along the siphonal line. Ornament comprises thirteen to sixteen prorsiradiate ribs per half-whorl, generally alternating long and short. The ribs are more or less strongly flexed and bear distinct ventrolateral tubercles. The latter are joined across the venter by convex ribs. Up to a diameter of 12 mm in USNMNH 237005 (10 mm in USNMNH 237012) ribs appear to be absent, although there are weak tubercles possibly marking their position along the ventrolateral shoulders. In USNMNH 237010, the main ribs are ornamented by rather distinct umbilical bullae. 250 ANNALS OF THE SOUTH AFRICAN MUSEUM 21-40 mm diameter: shell compressed (W/H = 0,53-0,68), very involute (umbilicus 15-17% of diameter), with a high rectangular whorl section. Ribbing denser (twenty to twenty-four ribs per half-whorl), not infrequently with two intercalatories between main ribs. The ventrolateral tubercles are generally still prominent at this stage. Fig. 46. Stoliczkaia (Stoliczkaia) tenuis Renz. Reconstructed juvenile and middle growth stages. Xl. 41-70 mm diameter: the ventrolateral tubercles are commonly lost between 40-45 mm diameter (37 mm in USNMNH 236996) and the ribs pass uninter- ruped across the venter, sometimes with a slight thickening in the ventrolateral position. The shell becomes slightly more inflated (W/~ = 0,58-0,69), with distinctly convex flanks and a rounded venter. The flank ribs coarsen consider- ably and there are commonly one or two intercalatories between long ribs, although in USNMNH 236994 and USNMNH 236987 there are probably more long ribs than intercalatories. Where there are two intercalatories between long ribs they may be of markedly different lengths (USNMNH 236984). Immediately prior to the aperture, all ornament is lost and the body chamber becomes smooth (USNMNH 236981). UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 251 V Ww X Fig. 47. Stoliczkaia (Stoliczkaia) tenuis Renz. A-C. USNMNH 236999. D-F. USNMNH 237009. G-I. USNMNH 237006. J-—L. USNMNH 237010. M-O. USNMNH 237015. P-R. USNMNH 237014. S—U. 236997. V-X. USNMNH 237003. x1. 252 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 48. Stoliczkaia (Stoliczkaia) tenuis Renz. A-C. USNMNH 236988. D-F. USNMNH 237003. G-I. USNMNH 236990. x1. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 253 Fig. 49. Stoliczkaia (Stoliczkaia) tenuis Renz. A-C. USNMNH 236945. D-F. USNMNH 237004. G-I. USNMNH 237000. J-—L. USNMNH 237001. M-N. USNMNH 237007. x1. ANNALS OF THE SOUTH AFRICAN MUSEUM 254 Fig. 50. Stoliczkaia (Stoliczkaia) tenuis Renz. A-C. USNMNH 236985. D-F. USNMNH 236984. x1. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL Measurements No. USNMNH USNMNH USNMNH USNMNH USNMNH USNMNH USNMNH USNMNH USNMNH USNMNH USNMNH USNMNH USNMNH USNMNH USNMNH USNMNH USNMNH USNMNH USNMNH USNMNH USNMNH USNMNH USNMNH USNMNH Intraspecific variation 236983 236985 236986 236987 236988 236989 236990 236992 236994 236995 236996 236997 236998 236999 237000 237001 237002 237003 237004 237005 237006 237008 237010 237012 38 H 32(48) 36(55) 29(52) 35(52) 33(48) 26(52) 27,5(50) 13,5(52) +19(50) 26(52) 30(54) 17,5(47) 18(54) 19(56) 19(56) 14(54) 15(50) 15,7(52) 10,8(53) +16,5(55) 17(54) 15(54) 9,5(54) 9(52) 9(51) 6(49) 11(52) 7(47) 8(47) 4(42) Ww +22(33) 26(40) 17,6(31) +23,5(35) ? 18(36) +19(34) 7,2(28) +10(26) 15,2(30) 18,7(34) +12(32) 9(26) +10(29) 11(32) 7,5(29) 9(30) 8,3(28) 5,5(27) 8,5(28) 10(32) 8(29) 6(34) +4,5(26) 5(28) 3,4(28) 6(29) 4(27) 4,5(26) 3(32) W/y 0,69 0,72 0,61 0,67 9 0,69 0,69 0,53 0,53 0,58 0,62 0,68 0,50 0,52 0,58 0,54 0,60 0,53 0,51 0,51 0,59 0,53 0,63 0,50 0,56 0,57 0,54 0,57 0,56 0,75 255 U 10(15) +13(20) +9(16) +9,7(14) 11,1(16) +9(18) +10(18) ? 7,7(15) 7,9(14) ? 6(17) 5,5(16) 5(15) ? 2 4,7(16) 2,5(12) ; +4,5(14) 4(14) 2,7(15) +3(17) 2(11) ? 3(14) 2,2(15) 2,5(15) +1,5(16) The large number of well-preserved specimens available to the authors permits a better understanding of the intraspecific and ontogenetic variation in this species. The marked ontogenetic change in ornament shown by this species has been outlined above. However, Figure 55 also shows that there is considerable variation in the degree of inflation of the whorls (W/H = 0,50-0,72), with a distinct tendency for the whorls to become more inflated at large diameters. Futhermore, Figure 55 shows that not only is there some variation in the width of the umbilicus, but there is also a slight tendency for the shell to become more evolute with growth. 256 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 51. Stoliczkaia (Stoliczkaia) tenuis Renz. A-D. USNMNH 236994. x1. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 257 Discussion As shown above, the present material is rather variable, and the type of S. (S.) tenuis and the specimen referred to as S. (S.) aff. tenuis by Renz (1968, pl. 6 (fig. 12)) fall within this range. When mature, S. tenuis closely resembles S. (S.) clavigera Neumayr (= Ammonites dispar Stoliczka (non d’Orbigny) 1865: 45, pl. 85 (figs 1-3 only)) (see Fig. 56), from which it appears to differ only in being consistently more compressed. Further work may show that the two merit only subspecific separation. S. (Stoliczkaia) dispar (d’Orbigny) (1841: 143, pl. 45 (figs 1-2)) is a widely- cited but much misinterpreted species. Consequently, the holotype is here photographically figured for the first time (Fig. 40), as well as a typical specimen from the dispar Zone of Dorsetshire, England (Fig. 41). S. (Stoliczkaia) dispar differs from the present species in having far more (up to nine) intercalatories between long ribs, whilst the latter are ornamented with weak umbilical bullae. The venter loses its ventrolateral tubercles and becomes rounded at smaller diameters and the body chamber ornament is also different; in S. (S.) dispar, ribbing rapidly weakens in maturity (at least on the internal mould) and all that Fig. 52. Stoliczkaia (Stoliczkaia) tenuis Renz. A-C. USNMNH 236987. x1. 258 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 53. Stoliczkaia (Stoliczkaia) tenuis Renz. A-B. Lateral and ventral views of SAM-— PCA3169. C—D. Ventral and lateral views of SAM—PCA5475. E. Lateral view of SAM-— PCAS5476. F. Lateral view of SAM-PCA2939. G-H. Lateral and ventral views of SAM-— PCA2938. I. Lateral view of SAM-—PCA3373. A-B, I x0,75, C-H x1. 259 UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL A-B. Lateral and ventral views of SAM-PCA6811. C—D. Ventral and lateral views of SAM-PCA2944. E. Lateral view of SAM-PCAS5477. F. Ventral view of SAM-—PCA5478. G-H. Mortoniceras (Angolaites) simplex (Choffat). Ventral and lateral views of SAM—PCA3107. A-B, E-H x1, C-D 0,75. Fig. 54. A-F. Stoliczkaia (Stoliczkaia) tenuis Renz. 260 ANNALS OF THE SOUTH AFRICAN MUSEUM ! " OTH EIGHT —______._g a a oes ™ UMBILICUS (%) —___g 0 70 DIAMETER (am) == Fig. 55. Plots of inflation and umbilical ratio against diameter for S. tenuis. Dots = present material; squares = holotype and paratype (after Renz 1968). remains are the weak umbilical bulges of the long ribs. Furthermore, the upper half of the flanks are concave, forming a broad, spiral depression. S. (Stoliczkaia) notha (Seeley) (1865: 232; Spath 1929: 335, fig. 110) (Fig. 57) differs from S. tenuis in having a rounded venter at all growth stages. Stoliczkaia dorsetensis Spath (1929: 337, pl. 31 (fig. 2), pl. 33 (fig. 1)) is more inflated and more densely ribbed than the Angolan material, as well as having more (three to nine) intercalatories between long ribs on the inner whorls. S. (Stoliczkaia) villoutreysi Casey (Fig. 43) has more intercalatories between long ribs (up to seven), whilst the venter is tabulate in maturity, when it homoeomorphs Hypacanthoplites. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 261 S. (Stoliczkaia) argonautiformis (Stoliczka) (1864: 87, pl. 46 (figs 1-2)) has strongly converging flanks, subdued ribbing and a more narrowly arched venter than the present species. Stoliczkaia tetragona Neumayr (nom. nov. pro Ammonites dispar Stoliczka (non d’Orbigny), 1864: 85, pl. 45 (fig. 2)) (Fig. 58) differs from the Angolan material in being much more inflated, although the style of ornament is similar in adults. Stoliczkaia grandidieri Boule, Lemoine & Thévenin (1907: 34, pl. 8 (fig. 8)), S. gardonica (Herbert & Meunier-Chalmas) (1875: 116, pl. 4 (figs 1-2)); S. rhamnonota (Seeley) (1865: 233, pl. 11 (fig. 7); Spath 1929: 333, fig. 109)) and S. blancheti (Pictet & Campiche) (1859: 188, pl. 23 (figs 2, 6); Renz 1968: 46, pl. 5 (fig. 21)) are all referable to the subgenus Faraudiella, whilst S. africana Pervinquiére (1907: 388, pl. 12 (fig. 10)), S. flexicostata Breistroffer (nom. nov. pro A. dispar Pictet & Campiche (non d’Orbigny) 1860: 264, pl. 38 (fig. 4); Renz 1968: 49, pl. 6 (fig. 9)), S. hashimotoi Matsumoto & Inoma, and S. asiatica Matsumoto & Inoma (1975) are all Shumarinaia. As suggested by Matsumoto & Inoma (1975), Stoliczkaia uddeni Bése (1927: 211, pl. 4 (figs 12-15)), is probably a synonym of S. texana Cragin (1893: 235, pl. 44 (fig. 1)) which itself is a close relative of S. crotaloides (Stoliczka) (1864: 88, pl. 46 (fig. 3)) (Fig. 59). The latter species differs from the Angolan material in maintaining single prorsiradiate ribs to the peristome, with only rare intercalatories, and in the flattened venter of the inner whorls; they may represent a distinct Cenomanian offshoot. ‘Stoliczkaia’ razafimbeloi Collignon (1968: 29, pl. 6 (fig. 7), pl. 7 (fig. 4)) and ‘S.’ vendegiesi Collignon (1968: 31, pl. 7 (fig. 5)) both differ from the present material in the development of distinct upper and lower ventrolateral tubercles on the body chamber and are thus transitional to Graysonites. They do not appear to be referable to the genus Stoliczkaia. Stoliczkaia patagonica Stoyanow (1949: 128, pl. 26 (figs 3-4)), S. excentrum- bilicata Stoyanow (1949: 129, pl. 26 (figs 5-6)) and S. scotti Stoyanow (1949: 129, pl. 26 (figs 7-8)) are all from the same stratigraphic level and locality and it is doubtful whether more than one species is represented. In this material, flexuous main ribs are ornamented with umbilical bullae and separated by two to four intercalatories. The ribs pass strongly across the venter and appear to lack ventrolateral tubercles. These ‘species’ are very close to S. dorsetensis Spath. Stoliczkaia adkinsi Bose (1927: 193, pl. 18 (figs 9-17)) differs from the present material in having more strongly differentiated long and short ribs, the former with distinct umbilical tubercles. ‘Submantelliceras’ worthense (Adkins) (1920: 93, pl. 1 (figs 11—13)) from the Pawpaw Formation of Tarrant County, Texas, may be based upon juveniles of Stoliczkaia. It differs from the present material in the possession of umbilical tubercles and in the (?) earlier loss of ventrolateral tubercles. Some of the Mantelliceras (Submantelliceras) saxbii (Sharpe) figured by Thomel (1972: 16-17, pl. 1 (figs 8-12 only)) are clearly based upon Stoliczkaia 262 ANNALS OF THE SOUTH AFRICAN MUSEUM A B Fig. 56. Stoliczkaia (Stoliczkaia) clavigera Neumayr. Copy of Stoliczka 1864, pl. 45 (fig. 1-1a). xi of the dispar—dorsetensis group and differ from the Angolan material in having more intercalatories between long ribs. Stoliczkaia praecursor Anderson (1958: 246, pl. 12 (fig. 1)) is too poorly figured and described for proper comment, but appears to differ from the present species in being more inflated, more coarsely ribbed, and in the presence of umbilical bullae at large diameters. It seems to be allied to S. tetragona Neumayr. In his original account of this species, Renz compared S. tenuis with Mantelliceras martimpreyi (Coquand), and, indeed, as demonstrated by the present population S. (S.) tenuis confirms that some Stoliczkaia have sub- mantellicerine nuclei, as suggested by Kennedy (1971). In particular, the authors are impressed by their close similarity to pyritic nuclei such as those figured by Collignon (1929, pl. 3 (figs 4-5)) from Diego-Suarez, Madagascar, as Acantho- ceras (Mantelliceras) martimpreyi Coquand. The latter differ only in having UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 263 C D Fig. 57. Stoliczkaia (Stoliczkaia) notha (Seeley). A-B. Holotype, Sedgwick Museum, Cambridge, B40. C-D. BMNH C4811, type of the variety u/tima Spath. Both from the Cambridge Greensand, Cambridge. 1. 264 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 58. Stoliczkaia tetragona Neumayr. Copy of Stoliczkaia 1864, pl. 45 (fig. 2-2a). x1. distinctly differentiated lower ventrolateral tubercles. However, there can be little doubt that records of Mantelliceras (Thomel 1972) and Submantelliceras (Adkins 1920) from the Upper Albian are based upon juveniles of Stoliczkaia. Kennedy & Hancock (1971) have shown Submantelliceras martimpreyi (Coquand) to be a junior subjective synonym of M. saxbii (Sharpe), from which the authors’ material differs in being more compressed, lacking distinct lower ventrolateral tubercles and in showing a marked, and characteristic, change of ornament on the body chamber. However, the fact that Thomel (1972) assigned late Albian species of Stoliczkaia to M. saxbii merely serves to emphasize the close relationship between these two genera and suggests that the origin of (at least) compressed Mantelliceras and Utaturiceras lies close to Stoliczkaia of tenuis type. The writers would also point to the close similarity of compressed variants to juvenile Forbesiceras (see Juignet & Kennedy 1977), generally described as Neopulchellia (a subjective synonym), and evidence for the descent of Forbesiceras from the S. (S.) tenuis group is to be published elsewhere. Occurrence S. (Stoliczkaia) tenuis is so far known only from the Upper Albian of Switzerland and Angola. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 265 Fig. 59. Stoliczkaia crotaloides (Stoliczka). Copy of Stoliczka 1864, pl. 46 (fig. 3-3a). xT Family Brancoceratidae Spath, 1934 Subfamily Brancoceratinae Spath, 1934 Genus Hysteroceras Hyatt, 1900 Type species Ammonites varicosus J. de C. Sowerby, 1824 Discussion Hysteroceras is typically a low Upper Albian micromorph genus, clearly descended from the earlier Brancoceras, and has generally been regarded as characteristic of, and largely confined to, the Mortoniceras inflatum Zone. However, micromorph brancoceratids persist into the highest levels of Albian where they have generally been referred to as Spathiceras or Mortoniceras (Cantabrigites). Whitehouse (1927: 110) introduced Spathiceras without formal diagnosis, merely naming as type of the genus Hystrichoceras antipodeum Etheridge (1902: 47, pl. 7 (figs 6—-7)). The holotype of S. antipodeum is from the Upper Albian of Point Charles, near Darwin, Northern Australia, where it occurs associated with Desmoceras latidorsatum (Michelin) (= D. carolensis Etheridge 1902, pl. 7 (figs 2-5)), Idiohamites cf. spinulosus (J. Sowerby) (= Ancyloceras (?) sp. ind., Etheridge 1902, pl. 7 (figs 14-15)), Scaphites eruciformis Etheridge (very close to S. simplex Jukes-Browne), Hamites cf. virgulatus (Brongniart) (= Hamites (?) sp. ind., Etheridge 1902, pl. 7 (figs 12-13)), Aucellina gryphaeoides (J. de C. Sowerby) (= A. incurva Etheridge), together with the genera Beudanticeras, Labeceras, Myloceras, Anisoceras and Ptychoceras (Whitehouse 1928). Because 266 ANNALS OF THE SOUTH AFRICAN MUSEUM V W X Y z Fig. 60. Hysteroceras? spp. juv. A-B. USNMNH 237018. C-E. USNMNH 237017. F. Hysteroceras? cf. ootatoorense (Stoliczka), USNMNN 237016. G—H, K-Z. Aysteroceras antipodeum (Etheridge), a series of limonitic specimens from northern Australia. G—H. BMNH C26539. K-L. BMNH C26551. M-N. BMNH C26523. O-Q. BMNH (C26552. R-T. BMNH C26548. U-V. BMNH C26518. W-X. BMNH C26546. Y-Z. BMNH C35269. I_J. Hysteroceras ? ootatoorense (Stoliczka). Copy of Stoliczka 1865, pl. 32 (fig. 2). All x1. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 267 Whitehouse (1928: 279) considered the fauna to be ‘. . . typical of the substuderi Zone of the Upper Albian’, the name Spathiceras has been applied to all flat-sided, weakly tuberculate micromorph mortoniceratinids of uppermost Albian age. However, the genera Labeceras, Myloceras and Beudanticeras are nowhere known to range above the Mortoniceras inflatum Zone, and the faunas recorded by Etheridge (1902) and Whitehouse (1927) seem to be typical low Upper Albian (approximately varicosum Subzone) assemblages. It is, perhaps, not coincidental, therefore, that S. antipodeum (Fig. 60K-—Z) bears a remarkable resemblance to Hysteroceras of the binum-subbinum group, from which it differs only in being consistently more evolute (umbilicus 40-45 % of the diameter). The differences are not sufficient for generic separation, and Spathiceras is considered a junior subjective synonym of Hysteroceras. There is no good evidence for Cenomanian occurrences of the genus. We are left, therefore, only with Hysteroceras for those uppermost Albian mortoniceratinid micromorphs with an evolute shell, and undivided ventro- lateral tubercles. Either these represent a continuation of the Hysteroceras stock, or they represent an unnamed homoeomorphic development. Until better and larger collections are known, the authors prefer to follow Renz (1968) and refer their material to Hysteroceras. Hysteroceras ? cf. ootaturense (Stoliczka, 1865) Fig. 60E-F, I-J Compare Ammonites ootaturensis Stoliczka, 1865: 56, pl. 32 (fig. 2). Mortoniceras ootaturense (Stoliczka) Pervinquiére, 1910: 64, pl. 6 (figs 2-5). Spathiceras ootaturense (Stoliczka) Spath, 1934: 444, 445, fig. 160h. Breistroffer, 1940: 75. ? Pervinquieria (2) sp. nov. Breistroffer, 1940: 75. Material A single specimen, USNMNH 237016, with recrystallized shell preserved, although somewhat corroded, from Porto Amboim. Description Shell small, evolute, with a wide, shallow umbilicus, steep umbilical walls and evenly rounded umbilical shoulder. The whorl section is subquadrate, compressed with flattened flanks. Ribbing is very weak on the inner flank, but strengthens markedly over the ventrolateral shoulder, whilst there is a prominent keel. Discussion Stoliczka’s (1865) type (Fig. 60I—J) comes from a locality ‘near Odium’, from where he also records other typical uppermost Albian species such as M. (Mariella) bergeri (Brongniart), M. (M.) circumtaeniatus (Kossmat), 268 ANNALS OF THE SOUTH AFRICAN MUSEUM Anisoceras perarmatum Pictet & Campiche, D. (Desmoceras) latidorsatum (Michelin), and Lechites gaudini (Pictet). Ammonites ootaturense is probably, therefore, of latest Albian age, although Stoliczka’s record of Turrilites costatus Lamarck and Neoptychites xetra (Stoliczka) from the same locality suggests the presence of beds as high as the Lower Turonian in the vicinity. Hysteroceras antipodeum (Etheridge) (Fig. 60G-—H, K-—Z) differs from the Angolan specimen in being somewhat older, with a wider umbilicus, flatter flanks and more prominent umbilical tubercles. The specimen of Schloenbachia rostratus var. antipodeus (Etheridge) figured by Etheridge (1909, pl. 67 (figs 3—-4)) (non Etheridge 1902) was referred to Dipoloceras bouchardianum (d’Orbigny) by Stieler (1920), but was renamed Prohysteroceras richardsi var. nitidum by Whitehouse (1926). However, Etheridge’s specimen appears indistinguishable from Dipoloceras quadratum Spath (1921: 278, pl. 25 (fig. 3)). Hysteroceras ? nanum Renz (1968: 63, pl. 11 (fig. 4), fig. 22c—d) differs from H? ootaturense in having prominent umbilical tubercles at an early stage. The specimen of H. semileve Haas recorded by Renz (1968: 63, pl. 11 (fig. 6), fig. 22g-h) from the ‘Unteren Vraconnien’ appears to be based upon a larger fragment of his H.? nanum. Hysteroceras ? tunisiense (Spath) (nom. nov. pro Mortoniceras inflatum var. orientalis (?) Pervinquiére (non Kossmat) 1907: 229, pl. 11 (fig. 2)) differs from H ? ootaturense and the present specimen in having sharp, flexuous prorsiradiate ribs which bifurcate from distinct umbilical bullae at only 17 mm diameter. Hysteroceras ? wenoense (Adkins) (1928: 229, pl. 20 (fig. 13)) differs from the authors’ material in its sharp ribbing and distinct umbilical tubercles. The specimen figured by Renz (1968: 62, pl. 11 (fig. 5)) as Hysteroceras cf. subbinum Spath may possibly belong here. ‘Algericeras’ boghariense (Coquand) (Pervinquiere 1907: 240, pl. 11 (fig. 16)) differs from the present specimen in having a quadrate whorl section (W/H = 1,00) with dense, fine, straight ribs, about thirty-two per half whorl, which arise in pairs from umbilical bullae and terminate in ventrolateral tubercles. It is said to be of Cenomanian age but appears merely to be based upon pyritic nuclei of Mortoniceras. Occurrence Hysteroceras ? ootaturense is known from the Upper (? uppermost) Albian of southern India, and possibly the uppermost Albian of France, Algeria and Angola. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 269 Subfamily Mortoniceratinae Spath, 1925 Genus Mortoniceras Meek, 1876 Subgenus Durnovarites Spath, 1932 Type species Subschloenbachia perinflata Spath, 1921 Discussion Wiedmann & Dieni (1968: 142) have divided the subgenus Durnovarites into two species groups: 1. The group of M. (D.) subquadratum, characterized by four rows of tubercles on the ribs. To this group may be assigned M. (D.) subquadratum Spath, M. (D.) quadratum Spath, M. (D.) perinflatum (Spath), M. (D.) postinflatum Spath, M. (D.) depressum (Spath), M. (D.) adkinsi (Young), M. (D.) vraconense Renz, M. (D.) spinosum (van Hoepen non Pervinquiére), M. (D.) subnanum (Breistroffer), M. (D.) ishiguaense Reyment, M. (D.) levecostatum Reyment, M. (?D.) crassicornutum (Reyment), M. (D.) lowrii McLearn, M. (D.) downii McLearn, M. (D.) rerati Collignon and M. (D.) haueri Collignon. 2. The group of M. (D.) spinosum, with only three rows of tubercles and sub- ordinate ribbing (appears merely to be based upon juveniles which have still to develop the fourth row of tubercles). To this group belong M. (D.) spinosum (Pervinquiére), M. (D.) kentronotum Spath, M. (D.) lemoinei (Spath), M. (D.) neokentroides Wiedmann & Dieni and M. (D.) aubersonense Renz. Mortoniceras (Durnovarites) perinflatum (Spath, 1922) Figs 3G, 61, 62D-I, 63-64 Ammonites inflatus Pictet & Campiche (non J. Sowerby), 1860: 178, pl. 21 (fig. 5), pl. 22 (fig. 3). Inflaticeras (‘Subschloenbachia’) perinflatum Spath, 1922: 113. ? Inflaticeras (Subschloenbachia) depressum Spath, 1922: 114, figs B, 2a—d. Inflaticeras (Subschloenbachia) quadratum Spath, 1922: 115. Pervinquieria quadrata (Spath) Spath, 19265: 423. ? Pervinquieria depressa (Spath) Spath, 1928: 51. Mortoniceras (Durnovarites) perinflatum (Spath) Spath, 1933: 430, pl. 40 (fig. 2), fig. 150. Wiedmann & Dieni, 1968: 143, pl. 14 (figs 3-4), fig. 92. Renz, 1968: 51, pl. 8 (figs 3, 5, 8), pl. 9 (figs 1-2), figs 17a, 18c, 19c, f. Marcinowski & Naidin, 1976: 109, pl. 6 (figs 1-2). Mortoniceras (Durnovarites) quadratum (Spath) Spath, 1933: 432, pl. 45 (fig. 3), pl. 46 (fig. 6), pl. 49 (fig. 12). Reyment, 1955: 38, pl. 6 (figs 4-5), pl. 7 (fig. 3), fig. 15. Wiedmann & Dieni, 1968: 145, fig. 93. Mortoniceras (Durnovarites) postinflatum Spath, 1933: 433, pl. 40 (figs 3-5), pl. 46 (figs 3, 7), pl. 47 (fig. 6). Renz, 1968: 53, pl. 8 (figs 1-2, 6), figs 17b, d, 18b, 19a—b, d. Marcinowski & Naidin, 1976: 109, pl. 7 (fig. 1), pl. 8 (fig. 1). ? Mortoniceras (Durnovarites) depressum (Spath) Collignon (in Besairie), 1936: 195. Reyment, 1955: 37, pl. 7 (fig. 4), fig. 14. Durnovarites adkinsi Young, 1957: 6, pl. 1 (figs 3, 6). Mortoniceras (Durnovarites) vraconense Renz, 1968 : 54, pl. 7 (figs 6-7, 11), fig. 19e. Marcinowski & Naidin, 1976: 111, pl. 9 (fig. 1). Material Two specimens, USNMNH 237021-22, both with recrystallized shell preserved from Porto Amboim, together with three specimens, SAM—PCA4802, 4576 and 4587, from Cabo Ledo. 270 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 61. Mortoniceras (Durnovarites) perinflatum (Spath). The holotype, Pictet collection, Natural History Museum, Geneva. From the Upper Albian of Vraconne, Switzerland. x1. Description The shell is moderately inflated and rather evolute (umbilicus 32-36% of the diameter), with a wide, shallow umbilicus and steep umbilical walls. The whorl section is almost quadrate (W/H = 0,93-1,05), with subparallel flanks. The umbilical shoulder is evenly rounded intercostally. Ribs begin at the umbilical seam and are rectiradiate to the umbilical shoulder where they termi- nate in weak bullae. The latter give rise to 1-2 prorsiradiate flank ribs, 38-43 per whorl and broader than the interspaces. All ribs are ornamented by a midlateral tubercle and closely spaced upper and lower ventrolateral tubercles. On the venter, the ribs pass forwards, finally becoming effaced in the sulci bordering the siphonal keel. The ribs show spiral ornament which is especially prominent on the ventrolateral tubercles. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 271 Fig. 62. A-C. Mortoniceras (Durnovarites) subquadratum Spath, USNMNH_ 237023. D-I. Mortoniceras (Durnovarites) perinflatum (Spath). D-F. USNMNH 237021. G-I. USNMNH 237022. x1. 272 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 63. Mortoniceras (Durnovarites) perinflatum (Spath). Ventral and lateral views of SAM-PCA4802. x1. Measurements ; No. D H WwW W/a U USNMNH 237021 ao 22,5(42) ? 2 17(32) ue 44 +19(43) +20(45) 1,05 ? USNMNH 237022 35 +16(46) +15(43) 0,93 12,5(36) Discussion Renz (1968) showed M. (D.) quadratum to be based upon juveniles of M. (D.) perinflatum, whilst M. (D.) vraconense appears to comprise hyponodose adults which the authors do not consider to bear specific separation from M. (D.) perinflatum. Similarly, M. (D.) postinflatum differs from the strictly contemporaneous M. (D.) perinflatum only by its more prominent ventrolateral tubercles and more inflated whorls (W/H = 1,20—-1,65). The authors do not regard the differences as sufficient for specific separation and place M. (D.) postinflatum within the synonomy of M. (D.) perinflatum, although the name might usefully be retained at the varietal level. Durnovarites adkinsi Young was separated from M. (D.) perinflatum on the basis of its slightly rounder ribs, less tumid flanks, and denser-ribbed inner UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 273 Fig. 64. Mortoniceras (Durnovarites) perinflatum (Spath). The holotype of Durnovarites spinosus van Hoepen, SAM—D3154, from the Upper Albian of the Mzinene Formation, Zululand. x1. whorls. The differences appear to be those between individuals, and the Texas species is considered to be referable to M. (D.) perinflatum. Mortoniceras (Durnovarites) depressum (Spath) was said to differ from M. (D.) quadratum in its far more depressed whorl section, with rounded instead of parallel flanks. It closely approaches M. (D.) postinflatum and may, therefore, be no more than an extreme variant of M. (D.) perinflatum. Occurrence Mortoniceras (Durnovarites) perinflatum is known from the uppermost Albian of Switzerland, Poland, southern England, Sardinia, Texas, Nigeria and Angola. The authors have also seen comparable material from Zululand. Mortoniceras (Durnovarites) subquadratum Spath, 1933 Figs 62A-C, 65C-D ? Subschloenbachia meunieri Spath, 1922: 115. ? Mortoniceras (Durnovarites) meunieri (Spath), Spath 1932: 399. Mortoniceras (Durnovarites) subquadratum Spath, 1933: 435, pl. 42 (fig. 5), pl. 43 (fig. 1), pl. 44 (fig. 6), pl. 45 (fig. 5), pl. 47 (figs 2-4), pl. 48 (fig. 2). Reyment, 1955: 38. Dieni & Massari, 1963: 798. Wiedmann & Dieni, 1968: 142, pl. 13 (fig. 9), fig. 90. Renz, 1968: 55, pl. 7 (figs 8, 10), pl. 10 (figs 1-4, 7-8), fig. 17f,-f.. Marcinowski & Naidin, 1976: 110, pl. 6 (fig. 3). Mortoniceras (Durnovarites) subquadratum var. tumida Spath, 1933: 435, pl. 48 (fig. 4). Wiedmann & Dieni, 1968: 143, pl. 13 (fig. 10), fig. 91. Mortoniceras (Durnovarites) subquadratum var. crassicostata Spath, 1933: 432, pl. 42 (fig. 9). ? Mortoniceras (Pervinquieria) sp. juv., Spath, 1933: 412, pl. 41 (fig. 7). ? Pervinquieria (Cantabrigites ?) subnana Breistroffer, 1947: 91. Pervinquieria (Durnovarites) subquadrata (Spath) Breistroffer, 1947: 61. Durnovarites spinosum van Hoepen, 1951: 324, figs 380-383. ? Mortoniceras (Durnovarites) levecostatum Reyment, 1955: 38, pl. 7 (fig. 2). ? Mortoniceras (Durnovarites) subnanum (Breistroffer) Renz, 1968: 56, pl. 10 (figs 5—6), fig. 17g. Material A single specimen, USNMNH 237023, with recrystallized shell preserved from Porto Amboim, and one, SAM-PCA3235, from Praia-Egito, preserved as an internal mould. ANNALS OF THE SOUTH AFRICAN MUSEUM 274 1% 60UttVJd-WVSs yUuA}eAeU E JO” SE EE OF TED — SMOIA [BIOIL] pue [eIUDA “AOU “ds JUOUsI]]09 (SajtADAOUANG) SD41BIIUOLAOW “J-A “SETEVOI-WYVS JO SMOIA [RIVA Pure [PINE T “yyedg winjoaponbqns (Sajtapaouing) spsaUojsoBM “G-D “OSTEVIdM-WYS JO SMOIA JerJoyeT pue [eIUSA “QeyoyD) xajdiuis (sajipjosup) svda1uoJAOw “G-VY °S9 “BI UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 275 Description The shell is very evolute (umbilicus 50° of diameter), compressed, with a slightly depressed, subquadrate whorl section (W/H = 1,11). The umbilicus is wide, shallow, with steeply inclined walls and evenly rounded umbilical shoulders. The flanks are flattened, with maximum width close to the umbilical shoulder, and converge slightly to the broad venter. There are 10 prominent, somewhat bullate umbilical tubercles per half- whorl, from which arise 1-2 rectiradiate to slightly rursiradiate ribs. Where single there is frequently an adjacent intercalated rib, so that there are 19 ribs per half-whorl. The ribs are thick, robust, about as wide as the interspaces, and are ornamented by closely spaced double ventrolateral tubercles. The lower ventrolateral tubercle is sharp and prominent, whereas the upper ventrolateral tubercle is more weakly developed and clavate. On the final third of the outer whorl (which is entirely septate), there is a weakly developed midlateral tubercle. The well-developed siphonal keel is separated from the upper ventrolateral clavi by prominent sulci. The ribs on the adoral portion of the outer whorl show weakly developed spiral ornament. Measurements No. D H W W/a U USNMNH 237023 46 13,5(29) +15(33) 1,11 23(50) Discussion Mortoniceras (Durnovarites) meunieri (Spath) (1922: 115; 1932: 399) (nom. nov. pro Ammonites inflatus Meunier (non J. Sowerby) 1887: 61, pl. 1 (fig. 2)) is an evolute species with about 30 coarse, rectiradiate to slightly rursiradiate ribs arising singly or in pairs from umbilical tubercles, each ornamented with a prominent midlateral tubercle and a (?) double ventrolateral tubercle. Meunier (1888) figured his specimen only in lateral view, without description, and hence comparison is difficult. If it is, indeed, a M. (Durnovarites), then not only might it be a synonym of M. (D.) subquadratum, but it also has priority over that name. This species differs from M. (D.) perinflatum (Spath) in its much wider umbilicus (43-50% as against 28-36%) and its typically less depressed whorl section (W/H = 1,03-1,16). However, adults of M. (D.) subquadratum are unknown and until population studies are undertaken it is not known whether the differences are truly of specific importance. Renz (1968) included Durnovarites spinosum van Hoepen (non Pervinquiére) in the synonomy of M. (D.) subquadratum, an assignment with which the authors concur; the type is re-illustrated here as Figure 64. Mortoniceras (Durnovarites) ishiaguense Reyment (1955: 38, pl. 7 (fig. 1)) is very close to M. (D.) subquadratum but is apparently much more densely ribbed. Since the Nigerian species is based upon a mature individual, it is not 276 ANNALS OF THE SOUTH AFRICAN MUSEUM directly comparable with M. (D.) subquadratum at the present time. Mortoniceras (Durnovarites) levecostatum Reyment (1955: 38, pl. 7 (fig. 2)) is based upon a somewhat distorted composite internal mould showing about twenty-six coarse, robust ribs per whorl and with a wide umbilicus: Judging from Reyment’s (1955) description, it may not bear separation from M. (D.) subquadratum. Occurrence Mortoniceras (Durnovarites) subquadratum is known from southern England, Poland, Switzerland, Sardinia, Zululand, and Angola. Mortoniceras (Durnovarites) collignoni sp. nov. Figs 65E-F, 66-67, 68B—D, 69 Material About a hundred specimens, in the South African Museum, Cape Town, from Praia-Egito. All are preserved as composite internal moulds. Type material SAM-PCA3227 is designated as holotype; paratypes are SAM-—PCA2975, 3189, 3199, 3202, 3217, 3259, 3277, 3294, 3309, 3317 and 3407. Etymology For the late General Maurice Collignon who contributed so much to our knowledge of ammonite systematics, and who helped the authors with their studies in many ways. Diagnosis A densely ribbed species of M. (Durnovarites) characterized by a dramatic change in shell morphology on the adult body chamber. The phragmocone has a subrectangular, compressed whorl section and is ornamented with 36-42 rectiradiate to prorsiradiate ribs per whorl which frequently bifurcate from umbilical bullae and are all ornamented with midlateral and double ventro- lateral tubercles. On the adult body chamber, however, all tuberculation is rapidly lost and the simple ribs develop a strong convex-adoral curvature, while the whorl section now becomes strongly compressed and lanceolate. Description Almost all the material has been crushed to varying degrees. The coiling is evolute (umbilicus about 24-30% of diameter), with a more or less compressed whorl section from an early growth stage (Fig. 69). Up to the body chamber, the intercostal whorl section is generally subrectangular, compressed. On the body chamber, the flanks become strongly convergent and the whorl section eventually becomes lanceolate. The umbilicus is wide, shallow, with a steep umbilical wall on the inner whorls, and an evenly rounded umbilical shoulder. On the outer whorl, the umbilical wall becomes sloping and the umbilical shoulder is gently rounded. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 277 Fig. 66. Mortoniceras (Durnovarites) collignoni sp. nov. A reconstruction based on the holotype and paratype material. 0,66. —eghap = aly mamas @ eal ANNALS OF THE SOUTH AFRICAN MUSEUM 278 ‘TX 3 ‘990 E-W “8LZTEVOd-WYS ‘odAjered & JO MOIA [BINT “OD SLZZTEVOd-WYVS ‘edAi0[0Y 9Y} JO SMOIA [e.I]UBA PUR [eJaIeT “g—YW “AOU ‘ds 1u0USI//09 (Sa1ADAOUANG ) SDADIIUOJAOJY *“D-Y “L9 “SIF UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 279 Fig. 68. A. Mortoniceras (Angolaites) simplex (Choffat). Ventral view of SAM-PCA3142. B-D. Mortoniceras (Durnovarites) collignoni sp. nov. B. Lateral view of a paratype, SAM-PCA3269. C-D. Lateral and ventral views of a paratype, SAM-PCA3182. E. Stoliczkaia tenuis Renz. Lateral view of SAM-—PCA3313. A-B x0,66, C-E x1. 280 ANNALS OF THE SOUTH AFRICAN MUSEUM In the earliest observed growth stages, the ribbing is generally simple, slightly prorsiradiate, with frequent intercalatories. All ribs are ornamented by distinct lateral and upper and lower ventrolateral tubercles, while long ribs arise from fairly prominent bullae. Even at this stage, spiral notching is evident on the tubercles. In the middle growth stages, the connections between the intercalated ribs and the umbilical bullae strengthen, and many ribs are seen to bifurcate from the latter, while the lateral and upper and lower ventrolateral tubercles become more swollen. At large growth stages, the ribbing again becomes simple but very subordinate to the now very swollen and prominent lateral and ventrolateral tubercles. At this stage, the lower lateral tubercle is somewhat clavate and the upper lateral and ventrolateral tubercles strongly so. The ventrolateral tubercles are now prominently raised above the narrow, sunken, keeled venter. On the last portion of the body chamber there is a great change in ornament. The whorl section changes from subrectangular to lanceolate, with the disappearance of all tubercles, and the ribs become strongly convex. There are generally 36—42 ribs per whorl in the middle growth stages, about as wide as the interspaces, with somewhat fewer in juveniles and on the outer whorl. Discussion The body chamber ornament of this species is characteristic. Mortoniceras (Durnovarites) perinflatum (Spath) (Renz 1968: 51, pl. 9 (fig. 1)) differs from M. (D.) collignoni sp. nov. in having a strongly depressed whorl section in maturity, whilst M. (D.) subquadratum Spath (1933: 435, pl. 37 (fig. 6)) differs from the Angolan species in being more evolute (umbilicus 40-48 % of the diameter) and in apparently lacking the dramatic change in body chamber ornament shown by M. (D.) collignoni. Mortoniceras rostratum (J. Sowerby) (Fig. 70) differs from the present species in having sparser, more distant ribbing, whilst the ribs of the body chamber retain four rows of tubercles almost to the peristome. Mortoniceras (Durnovarites) ishiaguense Reyment (1955: 38, pl. 7 (fig. 1)) differs from the present species in being more evolute, with less compressed whorls, and in apparently lacking the characteristic change in the body ornament seen in M. (D.) collignoni. Mortoniceras levecostatum Reyment (1955: 38, pl. 7 (fig. 2)) is from the same locality and horizon as M. (S.) ishiaguense but was said to differ in being more distantly ribbed, with more irregular ornament. The differences may not be of specific significance. Howarth (1965) considered Neokentroceras curvicornu crassicornutum Reyment (1955: 41, pl. 4 (figs 7-8)) a species of Durnovarites, but Reyment (1955) records this form in association with a typical N. curvicornu Spath, and hence it is much older than typical M. (Durnovarites). Mortoniceras (Styphloceras) lowrii McLearn (1972: 72, pl. 30 (figs 1-3), pl. 39 (figs 3-4)) and M. (S.) downii McLearn (1972: 73, pl. 31 (figs 1-3)) are UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 281 Fig. 69. Mortoniceras (Durnovarites) collignoni sp. nov. Whorl sections. A. SAM—PCA3202. B. SAM-PCA3309. C. SAM-—PCA3278. D. SAM-—PCA3257. E. SAM-PCA3309. x1. D both species of Durnovarites, thus indicating the presence of uppermost Albian strata at the Skidegate Inlet, British Columbia. Mortoniceras (Durnovarites) lowrii differs from the present species in having very depressed inner whorls, somewhat more distant ribbing, and in lacking the modifications of the body chamber ornament seen in the Angolan species. Mortoniceras (Durnovarites) downii is based upon body chamber fragments. In its swollen, clavate tuber- culation it approaches M. (D.) collignoni, but it appears to be more inflated, and shows no sign of the body chamber becoming lanceolate. Mortoniceras (Durnovarites) depressum (Spath) (1922: 114, figs B, 2a—d) is based upon a body chamber fragment of a specimen about 50 mm in diameter. At this stage, the whorls are very depressed (W/y = 1,41) and the fragment appears to have been very evolute. Slightly prorsiradiate ribs arise from prominent umbilical bullae and are indistinctly bifurcating, so as to appear alternating long and short. There are four rows of tubercles, with maximum width at mid-flank. This species would seem to be closely allied to M. (D.) subquadratum var. tumidum Spath; it differs from the Angolan material in being 282 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 70. The holotype of J. Sowerby’s Ammonites rostratus, from the Upper Greensand of Roak, near Benson, Oxfordshire. Oxford University Museum K835. 0,75. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 283 more coarsely ribbed and (?) more evolute, with a strongly depressed whorl section. Mortoniceras ( Durnovarites) baueri Collignon (1963: 159, pl. 305 (fig. 1311)) differs from M. (D.) collignoni sp. nov. in its coarse ribbing, depressed whorls, wide umbilicus and prominent umbilical tubercles. Its body chamber ornament is not known. Mortoniceras (Durnovarites) rerati Collignon (1963: 162, pl. 307 (figs 1312—1313)) differs from the Angolan material in much the same respects, but does not have the swollen umbilical tubercles of M. (D.) baueri. It very closely approaches M. (D.) ishiaguense. Mortoniceras (Durnovarites) subdepressum Collignon (in Besairie 1936: 196, pl. 21 (figs 4-5)) is based upon a unique fragment with a very depressed whorl section. It was said to differ from M. (D.) depressum in having straighter ribs which are not projected forwards on the venter. It is more coarsely ribbed with a more depressed whorl section than M. (D.) collignoni. Occurrence Mortoniceras (Durnovarites) collignoni is known only from the uppermost Albian of Angola. Subgenus Angolaites Spath, 1932 Type species Subschloenbachia gregoryi Spath, 1922 Discussion Angolaites was separated as a subgenus of Mortoniceras (Spath 1932: 380) for “. . . serpenticones, with single costation from a very early stage, and two peripheral tubercles, close together’. The characters of the subgenus are con- sistent, making it an easily recognized and useful taxon. Amongst mortoniceratinids, only Drakeoceras Young, 1957, and Canta- brigites Spath, 1933, have the same closely spaced ventrolateral tubercles whilst also lacking flank tubercles. Drakeoceras appears, however, to be a Goodhallites derivative characterized by its much narrower umbilicus and high whorls. The micromorph Cantabrigites is a contemporaneous form, abundant in western Europe where Angolaites is unknown, while the extreme rarity of Cantabrigites in Angola makes it unlikely that they represent sexual dimorphs. Mortoniceras (Angolaites) gregoryi (Spath, 1922) Figs 39G, 71, 72C, 73D Subschloenbachia gregoryi Spath, 1922: 127, pl. 3 (fig. 1). Mortoniceras (Angolaites) gregoryi (Spath) Reyment, 1955: 37, pl. 4 (fig. 13), pl. 6 (fig. 3). Material Three specimens, SAM-PCA3110, 3145 and 3235, from Praia-Egito, together with seven specimens from the Quissama Ridge of Cabo Ledo, SAM-— PCA4601, 4608, 4611, 4626, 4685, 4712, and 4813, all preserved as composite internal moulds. 284 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 71. Mortoniceras (Angolaites) gregoryi (Spath). The holotype, from the uppermost Albian at Catumbella, BMNH C20066. x1. Description The shell is very evolute, compressed, with a wide, shallow umbilicus (42-47% of the diameter). The umbilical walls are gently rounded and the flanks are slightly convex intercostally, converging somewhat towards the venter, with greatest width slightly above the umbilical shoulder. Ribs begin very weakly on the umbilical wall and pass radially outwards to the umbilical shoulder where they terminate in fairly prominent bullae, about seventeen per whorl. Each bulla gives rise to one to two flank ribs, with frequent intercalatories which become more abundant in maturity. Across the flanks the ribs are prorsiradiate and all are ornamented with closely spaced double ventro- lateral tubercles. There are thirty-nine ventrolateral tubercles on the outer whorl, and the upper ventrolateral tubercles are spirally notched. The venter is moderately narrow, with shallow sulci on either side of the siphonal keel. ‘TX “SPIEWOd-WVS JO MaIA [eI9}e'T “(YyredS) NN 1408048 (Sajwjosup) svdauojsop *D ‘OOTEWOd-WYS JO SMOIA eNUDA pue [e1eT “eYOyD) xajduus (sayivjosuy) svsaquojsop ‘g-y “ZL “B14 85 UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 286 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 73. A—C. Mortoniceras (Angolaites) simplex (Choffat). A. Lateral view of SAM-—PCA3142. B. Lateral view of SAM-—PCA3153. C. Lateral view of SAM-—PCA3114. D. Mortoniceras (Angolaites) gregoryi (Spath). Lateral view of SAM—PCA3147. A x0,66, B-D x1. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 287 Measurements No. D H W W/a U SAM-PCA3145 85 25(29) 20(24) 0,8 40(47) e 58 18(31) —(—) — 27(47) SAM-PCA3235 67 22(32) 20(30) 0,9 28(42) Discussion Mortoniceras (Angolaites) gregoryi differs from M. (A.) simplex (Choffat) and M. (A.) vicina (Haas) in the common occurrence of bifurcating and inter- calated ribs, and in having far fewer umbilical bullae. Occurrence Mortoniceras (Angolaites) gregoryi is known with certainty only from the Upper Albian of Angola and Nigeria. Mortoniceras (Angolaites) simplex (Choffat, 1905) Figs 3A—D, 54G-H, 72A-B, 73A-C, 74-77 Schloenbachia simplex Choffat, 1905: 35, pl. 4 (fig. 3). Pervinquieria simplex var. tenuis Haas, 1942: 81, pl. 16 (fig. 1), figs 7e, 8a. Inflaticeras sp. n. aff. gregoryi Spath, 1922: 127, pl. 3 (fig. 2). Pervinquieria vicina Haas, 1942: 82, pl. 16 (fig. 2), fig. 8b. Pervinquieria vicina var. evoluta Haas, 1942: 83, pl. 16 (fig. 3), fig. 8c. Material 9 specimens, SAM—PCA3107, 3116, 3142, 3146, 3150, 3153, 3166, 3200 and 3249, from Praia-Egito, together with 30 specimens from the Quissama Ridge at Cabo Ledo, SAM-PCA4575, 4578-79, 4581-82, 4584-85, 4588, 4590, 4593-94, 4596, 4605, 4609, 4613, 4615-16, 4618, 4628, 4631, 4640, 4718, 4756, 4770, 4774, 4863, 4867-69, and 4874, all preserved as composite internal moulds. Description This species is abundant at Egito, with adult specimens attaining a diameter of 170 mm (SAM-PCA3142). The shell is evolute, compressed, with the outer whorls only covering the preceding whorls to the top of the lower ventrolateral tubercles. The umbilicus is shallow and wide (41-47% of the diameter), with steep umbilical walls and well-rounded umbilical shoulders. Ribbing begins at the umbilical seam, is rather faint at first, strengthening as it passes radially outwards to the umbilical shoulder. The ribs may strengthen slightly on the umbilical shoulder, but true umbilical tubercles are absent. In the immature growth stages, the flank ribs are slightly sinuous but on the final whorl they become adorally concave. Rare intercalated ribs occur only during the very early ontogenetic stages, and there are about twenty ribs per whorl. Lateral tubercles are lacking, but all ribs are ornamented with closely spaced double 288 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 74. A-B. Mortoniceras (Angolaites) simplex (Choffat). Ventral and lateral views of SAM-PCA3116. C—D. Mortoniceras (Angolaites) cf. simplex (Choffat). Lateral and ventral views of SAM-PCA3179. x1. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 289 Fig. 75. Mortoniceras (Angolaites) vicina (Haas). The holotype BMNH-C20067. x1. ventrolateral tubercles which are spirally notched, especially the upper ventro- lateral tubercle. The narrow, rounded venter is keeled, with smooth sulci on either side. At large growth stages, the ventrolateral tubercles stand somewhat above the level of the keel. Measurements No. D H W W/H U SAM-PCA3142 173 48(28) 41(24) 0,85 81(47) si 135 45(33) 35(26) 0,77 56(41) SAM-PCA3107 91 31(34) 24(26) 0,77 37(41) SAM-PCA3200 — 43(—) 33(—) 0,76 — SAM-PCA3249 — 33(—) 26(—) 0,78 — SAM-PCA3150 — 39(—) 35(—) 0,89 — SAM-PCA3153 — 22(—) 19(—) 0,86 — SAM-PCA3146 iss si) 26(—) 0,84 a 290 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 76. Mortoniceras (Angolaites) simplex (Choffat). Whorl sections. A. SAM-—PCA3166. B. SAM-PCA3116. x1. Discussion Spath (1922) briefly discussed and figured a specimen which he considered to differ from M. gregoryi in having coarser, more distantly ribbed inner whorls (Fig. 75). This specimen was renamed Pervinquieria vicina by Haas (1942); it is in the British Museum (BMNH C20067) and, so far as the writers are able to judge, differs from M. (A.) simplex only in its coarser, more distant ribbing with five ribs in a distance equal to the whorl height, whereas in M. (A.) simplex there are eight to nine. The differences are slight, and within the range of variation seen in M. (A.) simplex from Egito; they are not regarded as of specific significance. Occurrence Mortoniceras (Angolaites) simplex is known only from Angola. Genus Cantabrigites Spath, 1933 Type species Mortoniceras (Cantabrigites) cantabrigense Spath; by original designation Discussion The first appearance of the name Cantabrigites Spath (1932: 380) was as a nomen nudum, the diagnosis and description of the type species appearing only a year later (Spath 1933: 436). Consequently, the valid date of introduction for Cantabrigites is 1933. Spath (1933) proposed Cantabrigites as a subgenus of Mortoniceras for ‘dwarf-forms with reduced, generally single and almost untuberculate costation and greatly simplified suture-line’. In maturity, many typical Mortoniceras commonly exceed 200-300 mm in diameter, whereas Cantabrigites is mature at diameters of less than 80 mm. It would appear, therefore, to be a genuine micromorph taxon. The fact that Cantabrigites is restricted to one level in the Upper Albian, viz. the dispar Zone, suggests that it is not the microconch of Mortoniceras. Cantabrigites differs so greatly from Mortoniceras vespertinum (Morton), the type of the genus, that the authors consider the differences sufficient for generic separation. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 291 Fig. 77. Mortoniceras (Angolaites) simplex (Choffat). Whorl sections. A. SAM-—PCA3249. B. SAM-PCA3147. C. SAM-PCA3150. D. SAM-PCA3144. E. SAM-PCA3200. F. SAM-PCA3153. G. SAM-PCA3111. H. SAM-PCA3179. I. SAM-PCA3200. x1. 292 ANNALS OF THE SOUTH AFRICAN MUSEUM Cantabrigites ? curvatum Renz, 1968 Figs 78-79 Cantabrigites curvatum Renz, 1968: 61, pl. 11 (figs la—b, 2a-c), figs 20g, 21f. Material A single specimen, SAM-PCA3177, from a horizon some way below the main occurrence of Stoliczkaia at Egito, in road gravels. Description The shell is evolute, with a wide, moderately deep umbilicus and a slightly compressed, subquadrate intercostal whorl section. The umbilical shoulders are well rounded and the flanks convex, with maximum width at about midflank. Ribs begin at the umbilical seam and pass radially outwards to small, but distinct, bullae on the umbilical shoulder. The ribs are rather thick, robust, and vary from slightly rursiradiate to slightly prorsiradiate across the flanks. Where no umbilical bullae are present, some ribs are intercalated at the level of the umbilical shoulder, so that there are eleven ribs per half whorl, of which six arise from umbilical bullae. Each rib is ornamented with a prominent, obliquely clavate, ventrolateral horn which projects backwards. There appears to be the faintest swelling just below the ventrolateral horns which may represent a very weakly developed lower ventrolateral tubercle. There is a distinct siphonal keel. Measurements No. D H W W/H U SAM-PCA3177 29 11(38) 9(31) 0,82 13(41) Discussion The Angolan specimen is indistinguishable from the holotype from Ste Croix, Switzerland, although the writers are in some doubt as to the generic identity of this species, since it closely resembles some of the earlier Neokentro- ceras spp. known from Angola. As, however, this species is known only from three small specimens and Neokentroceras is typical of the low Upper Albian of Angola and Nigeria, tentative assignment to Cantabrigites seems preferable, it being an homoeomorphous development only. Fig. 78. Cantabrigites? curvatum Renz. Lateral and ventral views of SAM-—PCA3177. Sill UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 293 Fig. 79. Cantabrigites? curvatum Renz. Whorl section of SAM-PCA3177. x2. Mortoniceras (Durnovarites) neokentroides Wiedmann & Dieni (1968: 146, pl. 13 (fig. 5)) was, as its specific name implies, considered to resemble Neokentroceras. It differs from the present species in having well-developed upper and lower ventrolateral tubercles which are elongated at right angles to the keel. Mortoniceras ? nanum Spath (1933: 411, pl. 43 (fig. 6), pl. 46 (figs 4-5), fig. 141) (Fig. 80) resembles the present species, but has double ventrolateral tubercles and lacks the posteriorly directed horns of C? curvatum. Occurrence Cantabrigites ? curvatum Renz is known only from Switzerland and Angola. Fig. 80. Mortoniceras? nanum Spath. The holotype, BMNH-C72726, from the Cambridge Greensand. x1. Genus Drakeoceras Young, 1957 Type species Drakeoceras drakei Young, 1957 Discussion Young (1957) erected the new genus Drakeoceras for Goodhallites-like forms in which the ventrolateral tubercles are doubled. As such, therefore, Drakeoceras bears the same relationship to Goodhallites that Angolaites does to 294 ANNALS OF THE SOUTH AFRICAN MUSEUM Mortoniceras, and further work may show that it is best treated as a subgenus of Goodhallites. Drakeoceras differs from M. (Angolaites) in being higher whorled, with Goodhallites-like :nner whorls and a more quadrate whorl section in maturity, as well as retaining prominent umbilical bullae and bifurcating ribs onto the body chamber. Drakeoceras cf. dellense Young, 1957 Figs 3E-F, 81-82 Compare Drakeoceras dellense Young 1957: 25, pl. 7 (fig. 1), pl. 10 (figs 4-5, 8-10), figs 2j, 3a, h-j. Material Six specimens, SAM-—PCA4662, 4673, 4705, 4733, 4786, and 4800, from the Quissama Ridge at Cabo Ledo, all poorly preserved limestone steinkerns, appear to belong here. Description The shell is strongly compressed and moderately involute (umbilicus 23-25 % of the diameter). The umbilicus is fairly narrow and shallow, with steep umbilical walls and evenly rounded umbilical shoulder. The broad flanks are slightly convex and converge towards the narrowly arched venter. The whorl section is compressed, elliptical (¥/H = 0,70-0,94). Fig. 81. Drakeoceras cf. dellense Young. Ventral and lateral views of SAM-PCA4800. x1. UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 295 Ornament comprises about twelve weak umbilical bullae per half-whorl, from which ribs commonly arise in pairs, or singly with an intercalated rib between long ribs. The ribs are initially strongly prorsiradiate, but recurve just below midflank. The ribs are broader than the interspaces and there are about twenty-four per half-whorl at the venter. All ribs are ornamented with very weak (? due to abrasion) lower and distinct upper ventrolateral tubercles. There is a prominent siphonal keel. Fig. 82. Drakeoceras cf. dellense Young. Eroded suture at 65 mm diameter. Approx. x 3. Discussion The Angolan specimen is closest to D. dellense from which it differs only in being somewhat more involute. It is significant that in Texas D. dellense is associated with M. (Durnovarites) perinflatum (Spath) (= D. adkinsi Young) and is probably, therefore, strictly contemporaneous with the Angolan speci- mens. Differences from other species of Drakeoceras are noted by Young (1957). Occurrence Drakeoceras dellense is currently known only from the dispar Zone of Texas and probably Angola. 296 ANNALS OF THE SOUTH AFRICAN MUSEUM Family Binneyitidae Reeside, 1927 Genus Borissiakoceras Arkhangel’skii, 1916 Type species Borissiakoceras mirabile Arkhangel’skii, 1916 Discussion As currently diagnosed (Wright in Arkell et al., 1957), the Binneyitidae is a family of micromorph ammonites characterized by their compressed, flat-sided form, narrow umbilicus and greatly simplified suture. Considered to be descended from the typically Middle Albian Falciferella, the earliest recorded binneyitids are from the Middle Cenomanian of the Western Interior (Cobban 1961), although we know of specimens of comparable age from western Europe (Kennedy & Juignet 1973) and Zululand. The time separating the last appearance of Falciferella and the first appearance of Borissiakoceras has been something of a problem in this phylogenetic scheme. However, Brunnschweiler (1959) has recorded species of ‘Falciferella’ from the late Albian of Australia which appear to bridge this gap. Our present record of Borissiakoceras from the uppermost Albian of Angola closes the gap even further, and, by extending the record of Borissiakoceras as far back as the late Albian, provides a direct morphological, chronological and phylogenetic link with the Middle Albian Falciferella. Cobban (1961: 747) diagnosed Borissiakoceras as follows: ‘This genus is characterized by the small size of the conchs which are ordinarily compressed and moderately evolute to somewhat involute. The venter is rounded to flat. Most shells are smooth but a few have raised falcoid growth lines or faint closely spaced falcoid ribs. Nodes, when present, are on the ventrolateral shoulder. The suture has a broad ventral lobe indented by a broad, shallow ventral saddle. The first lateral saddle is bifid and as wide as the ventral lobe. The first lateral lobe is narrow, bifid, and about half as wide as the ventral lobe. The second lateral saddle is trifid and higher and broader than the first. The second lateral lobe is about half as large as the first and tends to be bifid. The auxiliaries, which are small and shallow, may be divided into bifid lobes and saddles.’ Casey (1954: 27) introduced Falciferella (type species: F. milbournei Casey, 1954: 274, pl. 7 (figs 1-5), fig. 3) as follows: ‘Micromorph platycones with narrow, sharp-rimmed umbilicus and tabulate venter, feebly carinate in early youth. Test with strongly falcoid lineation or sub-costation, and a faint spiral groove at the middle of the sides. Mouth border plain. Suture-line of simplified Aconeceras pattern, with reduced auxiliary elements.’ In southern England, Falciferella is known only from the Middle Albian Euhoplites loricatus Zone (intermedius and niobe Subzones) (Owen 1971), although Brunnschweiler (1959) has recorded two species from the Upper Albian of Australia. Falciferella breadeni Brunnschweiler (1959: 15, pl. 1 (figs 5—6)) is associated with Labeceras and Myloceras and is thus of early late Albian age. In this species, the earliest whorls are smooth, but strong falcate ribs develop in maturity and form weak folds across the tabulate venter. The suture-line of UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 297 F. breadeni is simpler than in F. milbournei, in which respect it is closer to Borissiakoceras. It differs, however, in the trifid nature of the first lateral lobe, but there is a tendency towards trifurcation of this sutural element in B. orbiculatum Stephenson (cf. Cobban 1961, fig. Sa, f). Although a trifid first lateral lobe is known in Falciferella, the characters of F. breadeni are largely those of Borissiakoceras, and it is to the latter genus that we refer ‘F.’ breadeni and ‘F.” reymenti. Borissiakoceras sp. nov. ? aff. reymenti (Brunnschweiler, 1959) Fig. 31H-I Compare Falciferella reymenti Brunnschweiler 1959: 15, pl. 1 (figs 5-6). Material A single specimen, USNMNH 236980, with recrystallized shell preserved from Porto Amboim. Description Shell small, compressed (in part due to post-mortem deformation), with broad, flat flanks and a narrowly rounded venter. In places the venter appears to be fastigiate, but this is probably due to secondary crushing. The umbilicus is narrow (23% of the diameter) and the shell smooth. Measurements No. D H W W/a U USNMNH 236980 17 +7,3(43) +3(18) 0,41 4(23) Discussion The only other species of Borissiakoceras yet known from late Albian strata are B. breadeni (Brunnschweiler) (1959: 15, pl. 1 (figs 5—6)) and B. reymenti (Brunnschweiler) (1959: 16, pl. 1 (figs 7-8)) from the Oodnadatta region of South Australia. Both these species differ from the Angolan specimen in their much narrower umbilicus (13-14% of diameter) and in possessing rather prominent falcate ribs. Cobban (1961) has shown, however, that the latter feature is not consistently developed, even within a single Borissiakoceras population, with both ribbed and smooth variants occurring side by side. Consequently differences in ornament of this type may not be of specific importance. Smooth variants of Borissiakoceras orbiculatum Stephenson (1955: 64, pl. 6 (figs 1-4)) closely resemble the present specimen, but are much younger (Middle Cenomanian). Borissiakoceras compressum Cobban (1961: 747, pl. 87 (figs 19-33), pl. 89 (figs 1-9), fig. 4a-k) also resembles the present species, but is of Middle Cenomanian age and has a narrower umbilicus (14-17% of the diameter). Borissiakoceras mirabile Arkhangel’skii (1916: 55, pl. 8 (figs 2-3)) 298 ANNALS OF THE SOUTH AFRICAN MUSEUM differs from the Angolan example in its much wider umbilicus (37% of the diameter) as well as being a Lower Turonian species. Borissiakoceras reesidei Morrow (1935: 463, pl. 49 (fig. 7), pl. 50 (fig. 5), fig. 8) is an Upper Cenomanian species which, like B. orbiculatum, cannot be separated from the unique Angolan example without knowledge of the suture-line characteristics of the latter. Occurrence Upper Albian of Angola; species with which the specimen is compared came from the Upper Albian of Australia. AGE OF THE FAUNA At present there are certain problems concerning the scope and nomen- clature of the Stoliczkaia dispar Zone and its subzones. The divisions of the Albian Stage in England were erected by Spath on the basis of his understanding of the successions at Folkestone and elsewhere in England, and are based on firm stratigraphic principles. Spath (1943) gave the following zonation through the Upper Albian of southern England: Stoliczkaia dispar/ Durnovarites perinflatum Stoliczkaia dispar Zone | Subzone Arrhaphoceras substuderi Subzone Mortoniceras aequatoriale Subzone Callihoplites auritus Subzone Hysteroceras varicosum Subzone Hysteroceras orbignyi Subzone Mortoniceras inflatum Zone Whilst accepting Spath’s subdivision of the S. dispar Zone, Breistroffer (1940) showed that Arrhaphoceras substuderi (Spath) was common to both the perinflatum/dispar and substuderi Subzones and was, therefore, unsuitable for use as a subzonal index. Instead, he recognized a lower subzone of S. (Faraudiella) gardonica—S. (F.) rhamnonota— Mariella gresslyi—Turrilitoides toucasi. By 1947, however, Breistroffer had realized that S. (F.) gardonica and S. (F.) rhamnonota were synonyms of S. (F.) blancheti (Pictet & Campiche) and consequently renamed Spath’s A. substuderi Subzone the ‘Zone 4 Para- turrilites Gresslyi, Turrilitoides Hugardianus et Stoliczkaia (Faraudiella) Blancheti’. The upper horizon (Spath’s dispar/perinflatum Subzone) was referred to a ‘Zone a Pervinquieria (Durnovarites) perinflata, P. (Subschloenbachia) rostrata, Paraturrilites Bergeri et Stoliczkaia dispar’. On the basis of newly exposed sections, however, Owen (1976) has recently shown that Spath’s aequatoriale Subzone is a remanié assemblage which cannot be distinguished from the underlying subzone of Callihoplites auritus and he has, therefore, proposed its abandonment. In addition, Owen (1976) recalled Brei- stroffer’s (1940) otservation that Arrhaphoceras substuderi was unsuitable for use as a subzonal index in the S. dispar Zone and, because Owen (1976: 492) quite inexplicably considered S. (F.) blancheti ‘. . . is not sufficiently distinct to ay, UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 299 be used as a subzonal index’, he proposed to replace this subzonal index with Mortoniceras rostratum (J. Sowerby). This, however, is a very unfortunate suggestion because M. rostratum was for many years (because of mis- identification) the index species for the earlier zone now called after M. inflatum. In addition, Breistroffer (1947) has subsequently used it as one of the subzonal indices for Spath’s dispar/perinflatum Subzone. Moreover, it is abundantly clear that the true characters of M. rostratum are not known since all the material assigned by Spath (1932, pl. 38 (fig. 4), pl. 39 (fig. 4), pl. 40 (figs 1, 7), pl. 41 (fig. 7)) to this species was renamed Pervinquieria fallax by Breistroffer (1940: 67). Thus, only Sowerby’s holotype (Fig. 70), now in the Oxford University Museum, is without doubt assignable to this species. The authors’ have had the oppor- tunity of studying the holotype of M. rostratum. Its inner whorls are obscured by matrix whilst the ribs of the body chamber bear four rows of tubercles. Although a number of species of Mortoniceras s.s. have four rows of tubercles on the inner whorls, the upper and lower ventrolateral tubercles coalesce on the body chamber of M. (Mortoniceras) to produce ventrolateral horns and thus only three rows of tubercles remain on the body chamber. In the writers’ opinion, therefore, M. rostratum is a species of Durnovarites, a subgenus so far recorded only from Spath’s dispar/perinflatum Subzone, and is unsuited, therefore, for use as the subzonal index of the lower part of the S. dispar Zone. Moreover, since Owen (1976) does not state what he takes to represent M. rostratum, the species to which he is referring is totally unrecognizable. For this reason, the writers would propose a simple return to Breistroffer’s (1947) subdivision of the S. dispar Zone into a lower subzone of S. (F.) blancheti, immediately overlying the Callihoplites auritus Subzone, and an upper subzone of M. (Durnovarites) perinflatum. With the possible exception of the lower horizon at Egito, the present faunas can be referred with confidence to the M. (D.) perinflatum Subzone on the basis of the presence of S. (Stoliczkaia), M. (Durnovarites) and the hetero- morphs present. There does, however, seem to be some compositional difference in the faunas. Thus, at Praia-Egito the fauna comprises :* Anisoceras perarmatum Pictet & Campiche . armatum (J. Sowerby) . haasi sp. nov. . phillipsi sp. nov. . ef. arrogans (Giebel) . aff. exoticum Spath . aff. subarcuatum Spath . aff. spathi (Wiedmann) Hamites virgulatus Brongniart H. duplicatus Pictet & Campiche Puzosia cf. sharpei Spath Desmoceras latidorsatum perinflatum subsp. nov. MRA AAA DR * These lists on pp. 299-301 follow the order in the text. Ed. 300 ANNALS OF THE SOUTH AFRICAN MUSEUM Stoliczkaia tenuis Renz Mortoniceras (Durnovarites) collignoni sp. nov. M. (Angolaites) simplex (Choffat) M. (A.) gregoryi (Spath) A somewhat lower level at this locality has yielded: Tetragonites kitchini (Krenkel) Mariella gresslyi (Pictet & Campiche) Cantabrigites ? curvatum Renz The fauna from Cabo Ledo includes the following species: Anisoceras perarmatum Pictet & Campiche A. armatum (J. Sowerby) A. phillipsi sp. nov. Idiohamites dorsetensis Spath Hamites virgulatus Brongniart Mariella cf. oehlerti (Pervinquiére) Stoliczkaia sp. Mortoniceras (Durnovarites) perinflatum (Spath) M. (Angolaites) simplex (Choffat) M. (Angolaites) gregoryi (Spath) M. (Mortoniceras) spp. Drakeoceras cf. dellense Young This fauna differs from the higher horizon at Praia-Egito in that M. (Mortoniceras) is still fairly abundant whilst M. (Durnovarites) is rather rare. This suggests that the Cabo Ledo fauna may be somewhat older than the upper horizon at Praia-Egito, although the possibility of mixing of different horizons cannot be wholly dismissed. The fauna from Porto Amboim comprises: Phylloceras (Hypophylloceras) seresitense Pervinquiére Tetragonites collignoni Breistroffer T. jurinianus (Pictet) Eogaudryceras italicum Wiedmann & Dieni Anisoceras perarmatum Pictet & Campiche A. armatum (J. Sowerby) A. haasi sp. nov. Idiohamites dorsetensis Spath I. cf. elegantulus Spath I. pygmaeus sp. nov. Hamites virgulatus Brongniart Mariella circumtaeniatus (Kossmat) Desmoceras latidorsatum perinflatum subsp. nov. Stoliczkaia tenuis Renz UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 301 Mortoniceras (Durnovarites) perinflatum (Spath) M. (D.) subquadratum Spath M. (Angolaites) simplex (Choffat) Hysteroceras ? cf. ootaturense (Stoliczka) Borissiakoceras sp. nov? aff. reymenti (Brunnschweiler) The abundance of very compressed Stoliczkaia at Porto Amboim, together with the relative rarity of mortoniceratinids, suggests that this fauna may be somewhat younger than the upper horizon at Praia-Egito, although stratigraphic evidence for this is lacking. None the less, the Stoliczkaia fauna from here bears a very close resemblance to the ‘Submantelliceras’— Utaturiceras assemblages that characterize the basal Cenomanian of many regions, although the persistence of Mortoniceras at this level suggests that it is still best regarded as uppermost Albian. When the composition of the Angolan faunas is considered, there are marked differences from the faunas of the European perinflatum subzone in addition to the obvious absence of hoplitids. Noticeable in the Angolan assemblage is the complete absence of the widely distributed Mariella of the bergeri-miliaris group, Lechites, Stoliczkaia (Faraudiella), Turrilitoides, Scaphites, Ostlingoceras and nautiloids, together with the rarity of Hamites and Canta- brigites. Moreover, Stoliczkaia tenuis is rather different from the S. dorsetensis— notha—dispar plexus which characterizes the uppermost Albian of western Europe. Although some differences may be ecological (in particular with respect to the heteromorphs and the absence of hoplitinids), it seems possible that two slightly different levels in the uppermost Albian are represented. It is perhaps noteworthy, therefore, that in Texas, M. (D.) perinflatum is known only from the Pawpaw Formation (Young 1957), some way below the basal Cenomanian faunas of the uppermost Main Street and Grayson/Del Rio sequences. It also occurs below the main level of Stoliczkaia in Poland (Marcinowski & Naidin 1976). Notwithstanding these differences, the abundance of Stoliczkaia, together with M. (Durnovarites) perinflatum and Anisoceras of the perarmatum group, is sufficient to date the Porto Amboim fauna at M. (D.) perinflatum Subzone of the S. dispar Zone, and to point to the possibility of even further subzonal refinement of this critical interval. SUMMARY The Angolan littoral has yielded rich ammonite faunas referable to the uppermost Albian zone of Stoliczkaia dispar, and mainly to the upper subzone of Mortoniceras (Durnovarites) perinflatum. The following species are described : Phylloceras (Hypophylloceras) seresitense Pervinquiére Tetragonites (Tetragonites) collignoni Breistroffer Tetragonites (Tetragonites) kitchini (Krenkel) 302 ANNALS OF THE SOUTH AFRICAN MUSEUM Tetragonites (Tetragonites) jurinianus (Pictet) Eogaudryceras (Eogaudryceras) italicum Wiedmann & Dieni Anisoceras (Anisoceras) perarmatum Pictet & Campiche Anisoceras (Anisoceras) armatum (J. Sowerby) Anisoceras (Anisoceras) haasi sp. nov. Anisoceras (Anisoceras) phillipsi sp. nov. Anisoceras (Anisoceras) cf. arrogans (Giebel) Anisoceras (Anisoceras) aff. subarcuatum Spath Anisoceras (Anisoceras) aff. exoticum Spath Anisoceras (Anisoceras) cf. spathi (Wiedmann) Idiohamites dorsetensis Spath Idiohamites cf. elegantulus Spath Idiohamites pygmaeus sp. nov. Hamites virgulatus Brongniart Hamites duplicatus Pictet & Campiche Mariella (Mariella) circumtaeniatus (Kossmat) Mariella (Mariella) gresslyi (Pictet & Campiche) Mariella (Mariella) cf. oehlerti (Pervinquiére) Mariella (Mariella) nobilis (Sukes-Browne) Puzosia (Puzosia) cf. sharpei Spath Desmoceras (Desmoceras) latidorsatum perinflatum subsp. nov. Stoliczkaia (Stoliczkaia) tenuis Renz Hysteroceras ? cf. ootaturense (Stoliczka) Mortoniceras (Durnovarites) perinflatum (Spath) Mortoniceras ( Durnovarites) subquadratum Spath Mortoniceras (Durnovarites) collignoni sp. nov. Mortoniceras (Angolaites) simplex (Choffat) Mortoniceras (Angolaites) gregoryi (Spath) Drakeoceras cf. dellense Young Cantabrigites ? curvatum Renz Borissiakoceras sp. nov. ? aff. reymenti (Brunnschweiler) The majority of these species have not previously been described from Angola; although precisely correlated with the perinflatum Subzone, there are differences in composition when compared with European faunas. In part these reflect differences between biogeographic provinces (e.g. the sparsity or absence of some heteromorphs). Other differences suggest that it may be possible to further subdivide the dispar Zone and the fauna also permits discussion of recent reviews of Upper Albian zonation, especially by Owen (1976). A return to a twofold division of the dispar Zone into Stoliczkaia (Faraudiella) blancheti and Mortoniceras (Durnovarites) perinflatum Subzones is suggested. The chief systematic conclusions from the paper are discussions of intra- specific variation in Phylloceras (Hypophylloceras), T. (Tetragonites), D. (Desmoceras), E. (Eogaudryceras) and S. (Stoliczkaia) species. ~w UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 303 ACKNOWLEDGEMENTS Our best thanks are to Messrs F. Collier and E. G. Kauffman (U.S. National Museum) for allowing us to study the Washburn Collection. Dr M. K. Howarth, Dr H. G. Owen and Mr D. Phillips (British Museum), Dr R. Casey and Mr C. J. Wood (Geological Museum, London), Dr J. M. Hancock (London), Mr C. W. Wright (Oxford), Dr B. Rickards (Cambridge), Dr J. Sornay (Paris), and Dr J. P. Lefranc (Montpellier) assisted us in many ways, and their help is gratefully appreciated. Mr C. W. Wright critically reviewed the manuscript and saved us from a number of errors, for which we are most grateful. 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Stratigraphie der Mittleren Kreide (Gargasien und Albien) der oberen helvetischen Decken in den nordlichen Schweizeralpen. Neue Denkschr. schweiz. naturf. Ges. Abh. 1: i-vii, 1-148. GiEBEL, C. G. 1852. Die Fauna der Vorwelt mit steter Beriicksichtigung der lebende Thiere 3. Mollusken (i): 1-856. Leipzig: Brockhaus. HAAN, G. DE 1825. Monographiae Ammoniteorum et Goniatiteorum. Lugduni Batavorum. Haas, O. 1942. The Vernay Collection of Cretaceous (Albian) ammonites from Angola. Bull. Am. Mus. nat. Hist. 81: 1-224. Haas, O. 1952. Some Albian desmoceratid and lytoceratid ammonites from Angola. Am. Mus. Novit. 1561: 1-17. Hauer, F. von. 1861. Uber die Petrefacten der Kreideformation des Bakonyer Waldes. Sber. Akad. Wiss. Wien 44: 631-659. HAUGHTON, S. H. 1924. Notes sur quelques fossiles crétacés de l’Angola (céphalopodes et echinides). Comungdes Servs. geol. Port. 15: 79-106. HaAuGHTON, S. H. 1925. Note on some Cretaceous fossils from Angola (Cephalopoda and Echinoidea). Ann. S. Afr. Mus. 22: 263-288. HEBERT, E. & MUNIER-CHALMAS, E. C. P. A. 1875. Description du terrain Crétacé supérieur du Bassin d’Uchaux. Appendice paléontologique. Ann. Sci. Géol. 6: 113-122. Howarth, M. K. 1965. Cretaceous ammonites and nautiloids from Angola. Bull. Br. Mus. nat. Hist. Geol. 10: 335-412. Jaco, C. 1908. Etude sur quelques ammonites du Crétacé moyen. Mem. Soc. géol. Fr. 15 (38): 1-64. JUIGNET, P. & KENNEDY, W. J. 1977. Faunes d’ammonites et biostratigraphie comparée du Cénomanien du nord-ouest de la France (Normandie) et du sud de l’Angleterre. Bull. Soc. Géol. Normandie 63: 1-192. JuKES-BRrowneE, A. J. 1875. On the relations of the Cambridge Gault and Greensand. Q. JI. geol. Soc. Lond. 31: 256-314. JuKEs-BRowneE, A. J. 1877. Supplementary notes on the fauna of the Cambridge Greensand. Q. Jl. geol. Soc. Lond. 33: 485-504. KENNEDY, W. J. 1971. Cenomanian ammonites from southern England. Spec. Pap. Palaeont. 8: 1-133. KENNEDY, W. J. & HANCOCK, J. M. 1971. Mantelliceras saxbii (Sharpe) and the horizon of the Martimpreyi Zone in the Cenomanian of England. Palaeontology 14: 437-454. KENNEDY, W. J. & JUIGNET, P. 1973. First record of the ammonite family Binneyitidae Reeside 1927 in western Europe. J. Paleont. 47: 900-902. KENNEDY, W. J. & KLINGER, H. C. 1977a. Cretaceous faunas from Zululand and Natal, South Africa. The ammonite family Phylloceratidae. Bull. Br. Mus. nat. Hist. (Geol.) 27 (5): 347-380. KENNEDY, W. J. & KLINGER, H. C. 1977b. Cretaceous faunas from Zululand and Natal, South Africa. The ammonite family Tetragonitidae. Ann. S. Afr. Mus. 73: 149-197. K.uincer, H. C. 1976. Cretaceous heteromorph ammonites from Zululand. Mem. geol. Surv. S. Afr. 69: 1-142. 306 ANNALS OF THE SOUTH AFRICAN MUSEUM Kuincer, H. C. & KENNEDY, W. J. 1978. Turrilitidae (Cretaceous Ammonoidea) from South Africa, with a discussion of the evolution and limits of the family. J. Moll. Stud. 44: 1-48. KossMaT, F. 1895-1898. 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London: B. Meredith. Sowersy, J. DE C. 1823-1846. The mineral conchology of Great Britain. London: B. Meredith. Sowersy, J. DE C. 1850. In Dixon, F. The geology and fossils of the Cretaceous and Tertiary formations of Sussex. 1st ed. London: R. & J. Taylor. SpaTu, L. F. 1921. On Cretaceous Cephalopoda from Zululand. Ann. S. Afr. Mus. 12: 217-321. SpatH, L. F. 1922. On Cretaceous Ammonoidea from Angola collected by Professor J. W. Gregory, DSc., F.R.S. Trans. R. Soc. Edinb. 53: 91-160. SpaTu, L. F. 1923-43. A monograph of the Ammonoidea of the Gault. Palaeontogr. Soc. (Monogr.): 1-787. SpaTH, L. F. 19255. On Upper Albian Ammonoidea from Portuguese East Africa. With an appendix on Upper Cretaceous ammonites from Maputoland. Ann. Transv. Mus. 11: 179-200. SPATH, L. F. 1926. On the zones of the Cenomanian and the uppermost Albian. Proc. Geol. Ass. 37: 420-432. STEPHENSON, L. W. 1955. Basal Eagle Ford fauna (Cenomanian) in Johnson and Tarrant Counties, Texas. Prof. Pap. U.S. geol. Surv. 274-C: 53-67. STIELER, C. 1920. Uber sogenannte Mortoniceraten des Gault. Zenth/. Miner. Geol. Paldont. 1920: 345-352, 392-400. STOLICZKA, F. 1863-1866. The fossil Cephalopoda of the Cretaceous rocks of southern India. Mem. geol. Surv. India Palaeont. indica (3) 1-13: 41-216. Stoyanow, A. 1949. Lower Cretaceous stratigraphy in southeastern Arizona. Mem. geol. Soc. Am, 95: 1-99. SWENSEN, A. J. 1963. Anisoceratidae and Hamitidae (Ammonoidea) from the Cretaceous of Texas and Utah. Brigham Young Univ. Geol. Stud. 9 (2): 53-82. SZAJNOCHA, L. 1884. Zur Kenntnis der mittelcretacischen Cephalopoden-Fauna der Inseln Elobi an der westkuste Afrikas. Denkschr. Akad. Wiss. Wien 49: 231-238. 308 ANNALS OF THE SOUTH AFRICAN MUSEUM TuHomeL, G. 1972. Les Acanthoceratidae de chaines subalpines méridionales. Mem. Soc. géol. Fr. 116: 1-204. VAN Hoepen, E. C. N. 1951. Die gekielde ammoniete die Suid-Afrikaanse Gault. VII Pervinquieridae, Arestoceratidae, Cainoceratidae. Paleont. Navors. nas. Mus. Bloemfontein 1:1-38. VENZO, S. 1936. Cefalopodi del Cretacea medio-superiore dello Zululand. Palaeontogr. ital. 36: 59-133. WuitEaVEs, J. F. 1876. Invertebrates from the coalbearing rocks of the Queen Charlotte Islands. Mesozoic Fossils 1: 1-92. WHITEHOUSE, F. W. 1926. The Cretaceous Ammonoidea of Eastern Australia. Mem. Queens. Mus. 8: 195-242. WHITEHOUSE, F. W. 1927. Additions to the Cretaceous ammonite fauna of Eastern Australia. Mem. Queensl. Mus. 9: 109-119; 200-206. WHITEHOUSE, F. W. 1928. The correlation of the marine Cretaceous deposits of Australia. Rep. 18th Meeting Austr. Assoc. Advanc. Sci., 1926: 275-280. WIEDMANN, J. 1962a. Ammoniten aus der vascogotischen Kreide (Nordspanien). I. Phyllo- ceratina, Lytoceratina. Palaeontographica A118: 119-237. WIEDMANN, J. 19626. Unterkreide-Ammoniten von Mallorca. I. Lfrg: Lytoceratina, Aptychi. Abh. math -naturw. Kl. Akad. Wiss. Mainz 1962: 1-148. WIEDMANN, J. 1964. Unterkreide-Ammoniten von Mallorca. 2. Lfrg: Phylloceratina. Abh. math. -naturw. Kl. Akad. Wiss. Mainz 1963: 155-264. WIEDMANN, J. 1973. The Albian and Cenomanian Tetragonitidae (Cretaceous Ammonoidea), with special reference to the circum-Indic species. Eclog. geol. Helv. 66: 585-616. WIEDMANN, J. & DIENI, I. 1968. Die Kreide Sardiniens und ihre Cephalopoden. Palaeontogr. ital. 64: 1-171. Woops, H. 1917. The Cretaceous faunas of the north-eastern part of the South Island of New Zealand. Bull. geol. Surv. N.Z. Palaeontology 4: 1-41. WRIGHT, C. W. & WRIGHT, E. V. 1951. A survey of the fossil Cephalopoda of the Chalk of Great Britain. Palaeont. Soc. (Monogr.): 1-40. YOuNG, K. 1957. Upper Albian (Cretaceous) Ammonoidea from Texas. J. Paleont. 31: 1-33. ZwIERZYCKI, J. 1913. Zur Frage der Unteren Kreide in Portugiesisch-Mozambique. Sber. Ges. naturf. Freunde Berl. 7: 319-326. 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 14-1SA 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. 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.’ ee ere 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. 7 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. M. R. COOPER & W. J. KENNEDY UPPERMOST ALBIAN (STOLICZKAIA DISPAR ZONE) AMMONITES FROM THE ANGOLAN LITTORAL oe he Oh et j a t ; Mee wor ~ 4 Owes ; y mn wie . AF oe ae ia " “pepe 0ST A). 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