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B RARI ES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS Sa iuvugd . s Bue Se Mh aie ae = ff. | Yip, = oi a Zz = Yl yy ope: z S WW 2 2 § t YY a = o BE ENS “oe ae fe) Lf fy Ee Wey 2 E Nv 2 = z 7 a wy 6 6 = SS = = > | UTION NOILNLILSNI_NVINOSHLINS S3IYVYEIT LIBRARIES SMITHSONIAN INSTITUTION uJ n Hs 7 Yy, = = z = = Gey = < .< = < UY m= 0c 5 WS ac = a fer S 2 ober o hg o 3 2 RS — > = yug tl BRARIES SMITHSONIAN INSTITUTION NOILNLILSNIT NYINOSHLIWS $3 1uVvud z | : S r o = S ; Sc 5. a = B) - Ee: = e > ° = = ES = c 2 = - Dilantin Z m 2 TUTION, NOILNLILSNI_ NYINOSHLINS S31YVYEIT_LIBRARIES, SMITHSONIAN _INSTITUTIG So a a a ee ee ee ee ae > Smo WWWHE zy] my 5 ye Of fj" ® OAT = 7S p (ey == S NY - 45 (e — A, 45 Lend Peoliog irae we = 4 cy aoe “a S was m AS = wasn m TTS = m Sings 2 Sf wasn’ 77) F = wm = 7) a RARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS SSIYVYUEIT LIBRAR 7 w a @ ” = wn Dp = ‘9 = < = Py < = yy =| Zz = yt tp, = = Z Wag 3 S 2 Fe i 9 z S \ O x -60 fYWYw = fe) xc ua oe 2 = 2,97 ff & Z| E SNS = = SS > = > = aN a 7) ieee za ” z= 7) : —NVINOSHLINS S3 iuvug i1_ti BRARI ES SMITHSONIAN INSTITUTION pe ae = ud = wl wy = ud 4 = 4 - Yay, % = aA _< — fot a < Cc feed Cc < ” bly [ exe =~ amd ae . yds Perl aed oO re) 3 a re) ss 5 2 Zz : J 2 a Zz a RARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3INvVUdIT LIBRAR S — Kim §& is z 2 = “0 = x ° 9 =, a 5 ro) = 9) = = = a = 2 AALILSNI NVINOSHLINS S31IYVHUSEIT LIBRARIES SMITHSONIAN _ NOILOLI = w = OE 2 2) z< WM = | = = z = : g i eS g 3 | g = 2 = Z = Z , a ~ iS a . 2B = Ne SRARIES SMITHSONIAN _INSTITUTION NOILNLILSNI NVINOSHLINS S3I¥vugit e Z in ZS xx o Z 77) BN 4 ‘3 : % QQ = a < *9, of x od x \N : < = am se: (e) = ro) ae eo) ~ Zz at = ae aa z LALILSNI NVINOSHIJIWS LIBRARIES SMITHSONIAN _ INSTITUTION NOILALI a: = oe = ee oh S peu) > ER ow ctor er] ee a \ 5 Fe) i ag Ee > VA E > = a> = bee z D z on z JRARIES, SMITHSONIAN INSTITUTION _NVINOSHLINS, SAIUVUEIT LIBRAF " = a = < i) ee 4 z 4 z= = ea KA. WE Z S ae ig x os N Ne oO el e oO £ = (@) eke “ 3 oS = 2 2, = z ee ww TESS cS = BS > Pid = > = “SS ANLILSNI_NVINOSHLINS SAIYVYGIT_LIBRARIES SMITHSONIAN INSTITUTION | NOILNLI o = tw = ud a w mn a a. ~ a . < =I < Ks; par < = S S a. = Oo 3 : = . ro) Br 5 2 Zz ; J 2 pir z a 3RARIES [SMITHSONIAN INSTITUTION NOILALILSNI NVINOSHLINS Saluvugia _LIBRAF rs) ee ° ~ o o om - wo = ai = - a = a - ° cw ra a = a Z f 2 yom = i ILOLILSNI pod aVUGIT LIBRARIES) SMITHSONIAN INSTITUTION,, NOILOL! <= RON ~ fix > “s < s . / ieee w Ws = Lath ram VON — Y er Mae eax (Sty A be (SQV >» ~ _—_ - ~ — y, ~ JTIO i - Ate A th 7) Ae fe iar oy re eae te ae 507, 6% ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM VOLUME 75 BAND THE TRUSTEES OF THE DIE TRUSTEES VAN DIE SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM CAPE TOWN KAAPSTAD 1978 ® SET, PRINTED AND BOUND IN THE REPUBLIC OF SOUTH AFRICA BY THE RUSTICA PRESS (PTY.) LTD., WYNBERG, CAPE 612 ANNALS OF THE ANNALE VAN DIE SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM VOLUME 75 BAND 75 LIST OF CONTENTS BARNARD, J. L. Redescription of the amphipod, Calliopiella michaelseni Schellenberg from South Africa, with comparison to a new genus from the Pacific Ocean. (Published March 1978.).. Best, P. B. see Ross, J. B. Cooper, M. R. Uppermost Cenomanian-Basal Turonian ammonites from Salinas, Angola. (Published March 1978.) DAVISON, P. & HOSFORD, J. Lobedu Pottery. (Published June 1978.) Day, J. Southern African Cumacea. Part 2. Family Bodotriidae, subfamily Bodotriinae. (Published June 1978.) HENDEY, Q. B. The age of the fossils from Baard’s Quarry, Langebaanweg, South Africa. (Published February 1978.) HOSFORD, J. see DAVISON, P. KENSLEY, B. Redescription of Parabathynomus natalensis Barnard (Crustacea, Isopoda, Cirolanidae). (Published February 1978.) #2 es a KENSLEY, B. A new marine isopod family from the South-western Indian Ocean. (Published February 1978.) Ross, G. J. B., SHAUGHNESSY, P. D. & BEsT, P. B. New records of crabeater seals (Lobodon carcinophagus) from South Africa. (Published March 1978.) SHAUGHNESSY, P. D. see Ross, G. J. B. Page 35 51 291 159 25 Al 153 iN ¥ 7 v i = t) i 1 ’ j | . ) 7 + { ; v i u } — t - ta 1 = 1 7 i ?. . f N 5 ry yi == 4 i 1 = R r my A ef e o bi a ——-- Bey NEW GENERIC NAMES PROPOSED IN THIS VOLUME Alticuma Day, 1978 Austrocuma Day, 1978 Callaska Barnard, 1978 Mossambicuma Day, 1978 Page 195 192 38 188 ‘V 'OLUME 75 PART fr. FEBRUARY 1978 ISSN 0303-2515 p907.68 OF THE SOUTH AF RICAN MUSEUM > ‘APE 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, 1969b; Jones 1971)’ ‘As described (Haughton & Broom 1927)...’ “As described (Haughton et al. 1927)...’ Note: no comma separating name and year Dagination indicated by colon, not p. names of joint authors connected by ampersand et al. in text for more than two joint authors, but names of all authors given in list of references. (b) Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, b, etc. to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a). For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list o, scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number (only if independently paged) in parentheses, pagination (first and last pages of article). Examples (note capitalization and punctuation) BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140. FIsCHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. gén. 74: 627-634. \ Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320. Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51. THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. (continued inside back cover) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 75 Band February 1978 Februarie Part 1 Deel THE AGE OF THE FOSSILS FROM BAARD’S QUARRY, LANGEBAANWEG, SOUTH AFRICA By Q. B. HENDEY 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.), 51-3, 5, 7-9), 6(1, t.-p.i.), 711-4), 8, 9(1-2, 7), 10(1-3), 11(1-2, 5, 7, t.-p.i.), 15(4-5), 24(2), 27, 31(1-3), 32(5), 33 Price of this part/Prys van hierdie deel R2,60 Trustees of the South African Museum © Trustees van die Suid-Afrikaanse Museum 1978 ISBN 0 908407 35 1 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 AGE OF THE FOSSILS FROM BAARD’S QUARRY, LANGEBAANWEG, SOUTH AFRICA By Q. B. HENDEY South African Museum, Cape Town (With 3 figures and 5 tables) [MS. accepted 13 September 1977] ABSTRACT Fossils from the mined-out Baard’s Quarry at Langebaanweg came from at least two stratigraphic horizons, those from the older horizon comprising the main body of fossils from the site. A re-evaluation of available Baard’s Quarry material strongly suggests that, contrary to earlier opinions, it postdates the fossils from the near-by Varswater Formation in ‘E’ Quarry. The older, and predominant element in the assemblage is apparently late Pliocene or early Pleistocene in age and includes late surviving archaic species and endemic species in association with more advanced and more widespread taxa. CONTENTS PAGE MMtROducthiOne Pee eon Aloe Ves ke pease 1 Age of the fossils from ‘E’ Quarry . y) Geology of the Baard’s Quarry area ks Bee 3 Provenance of the Baard’s Quarry fossils . . . 4 The ‘upper level’ assemblage . : 5) The ‘lower levels assemblage . . . .. . 7 IDYRCUSSIOIO Eee Ee en nS ene weer medi: NORVOWMCULSINTNIS 6 56 56 6 © 6 6 6 © AF INCiCheNCCSaP ey rier hs. a) ee” ee eS INTRODUCTION Fossils from phosphatic deposits in the vicinity of Langebaanweg, Cape Province, were first reported by Singer & Hooijer (1958). The specimen described by these authors came from Baard’s Quarry, which was mined-out and back-filled some years later. Most of the fossils collected in the Langebaanweg area between 1958 and 1964 came from Baard’s Quarry, other occurrences (see Hendey $970a: 84-86, 96-98, 1974: 37) having been relatively unimportant and not relevant to the present study. After 1964 both mining and fossil collecting was continued in “E’ Quarry (the New Varswater Mine), which is situated about 2 km west of Baard’s Quarry (Hendey 1970a, fig. 2). The ‘E’ Quarry fossil assemblage is vastly superior to that from Baard’s Quarry in all respects and the latter has received little attention in recent years (see Hendey 1974, 1976). One of the results of the shift in collecting from Baard’s Quarry (and less I Ann. S. Afr. Mus. 75 (1) 1978: 1-24, 3 figs, 5 tables. 2 ANNALS OF THE SOUTH AFRICAN MUSEUM significant occurrences) to ‘E’ Quarry is that earlier reports on fossils from Langebaanweg dealt largely with Baard’s Quarry, while later ones have dealt largely with ‘E’ Quarry. If, as some believe, the fossils from these two occurrences are contemporaneous and represent a single fauna, the provenance of specimens is relatively unimportant and references to ‘the Langebaanweg fauna’ are justified. This attitude has been most firmly supported by Wolff et al. (1973: 212) who stated that, ‘It would be dangerous to infer a major difference in time of deposi- tion’ of the Baard’s and ‘E’ Quarry fossils. An alternative view expressed earlier (Hendey 1970a, 1972) and which was later maintained (Hendey 1974) is that the Baard’s Quarry fauna includes some material of Pleistocene age (i.e. younger than ~ 2 million years) and some which is contemporary with the Pliocene “E’ Quarry (or Varswater Formation) fauna. More recently it has been indicated that all the Baard’s Quarry fossils postdate those from ‘E’ Quarry (Hendey & Deacon 1977). The purpose of the present paper is to review the available evidence on the age of the Baard’s Quarry fossils. It is not intended to provide a final and definitive statement on these fossils. Some of the more significant specimens from this site are no longer in the collections of the South African Museum and could therefore not be examined. In addition, although all the most diagnostic of the available specimens were studied, the material warrants further sorting and preparation, while at least some of it merits further detailed study. AGE OF THE FOSSILS FROM ‘E’? QUARRY Before dealing with the fossils from Baard’s Quarry, the dating of those from the Varswater Formation in ‘E’ Quarry is reviewed. As with most fossil occurrences in the south-western Cape Province, only relative dating methods can be applied in the case of the fossils from “E’ Quarry, and this has been based largely on correlations with fossils and faunas from east Africa. In recent years it has become widely accepted that the “E’ Quarry fossils date back between 4 and 5 million years (m.y.). These fossils occur in several different horizons within the Varswater Formation but, since it was believed that deposition of this formation had not been prolonged, the fossils of the different horizons were regarded as broadly contemporaneous. The three main units constituting the Varswater Formation are the Gravel, Quartzose Sand and Pelletal Phosphorite Members (Hendey 1976) and it is likely that intervals of time intervened between their deposition. There is at present no way of gauging the interval between deposition of the Gravel and Quartzose Sand Members, since the largely marine fauna of the former is poorly known and has little in common with the largely terrestrial fauna of the Quartzose Sand Member. The situation in respect of those deposits overlying the Gravel Member is more promising and fossiliferous horizons within these deposits are potentially datable in relative terms. For many years the correlation of the Varswater AGE OF THE FOSSILS FROM BAARD’S QUARRY, SOUTH AFRICA 3 Formation with the east African sequence was based on certain species included in the fauna of the Quartzose Sand Member. An age for this member of between 4 and 4,5 m.y. was suggested by the elephant in its fauna (see Maglio 1973), while the higher limit (i.e. 4,5 m.y.) was suggested to be more appropriate by the Nyanzachoerus (Hendey 1973). This is still regarded as a reasonable age estimate, there being nothing in the Quartzose Sand Member fauna to contradict it. The Pelletal Phosphorite Member is undoubtedly younger than the Quartzose Sand and there are indications that the interval between their deposi- tion may have been appreciable, although an age of less than 4 m.y. for the Pelletal Phosphorite was considered unlikely (Hendey 1976: 244). There are as yet no grounds for questioning the upper age limit of 5 m.y. for the Varswater Formation, but the 4 m.y. minimum date may be less secure. One of the most recently recognized complications in dating this formation stems from the fact that there are within the Pelletal Phosphorite Member two fossili- ferous horizons which were perhaps separated by yet another interval of time. These two horizons, which were apparently river channel deposits, are informally termed bed 3aS and bed 3aN (Hendey 1976: 226-230). They have yielded large numbers of fossils, although relatively few have yet been studied and it is there- fore premature to discuss their age. Nevertheless, it does seem advisable to regard the 3aS and 3aN faunas as separate units and to note the possibility that either the 3aN fauna or both may be less than 4 m.y. old. In the discussions which follow references are made to the ‘faunas’ of °E’ Quarry. They are comprised of the assemblages from, in descending order of age, the Quartzose Sand Member and beds 3aS and 3aN of the Pelletal Phos- phorite Member. Although there are doubts about their actual age, all are likely to date from the earlier half of the Pliocene. GEOLOGY OF THE BAARD’S QUARRY AREA There are striking differences between the deposits in the Baard’s Quarry area and those constituting the Varswater Formation as it is exposed in °E’ Quarry, but since there are no records of the succession in the intervening area, unequivocal correlation of the two sequences has not been possible (Hendey 1970a; Wolff et al. 1973). The situation is complicated by the fact that whereas the ‘E’ Quarry deposits, of which there are still extensive exposures, are well known and have been studied in detail, the Baard’s Quarry succession was inadequately recorded and is, by comparison, poorly known. Brief accounts of the Baard’s Quarry succession appeared in early reports on the Langebaanweg fossils (Singer & Hooijer 1958; Boné & Singer 1965), while there are some unpublished records available. In addition, Tankard (1975: 273, fig. 12) gave an account of deposits on the farm Muishondsfontein, which are a northerly extension of those which were exposed in Baard’s Quarry. There clearly was variation in the nature and depth of individual units in the succession, but apparently the basic sequence was as follows: 4 ANNALS OF THE SOUTH AFRICAN MUSEUM 4. Surface aeolian sands. 3. Ferricrete and other duricrusts in an unconsolidated sand matrix. 2. Deposits, some in river channels, comprised of phosphatic sandstone, quartz porphyry cobbles and other elements in a clayey sand matrix. This was the unit from which the Baard’s Quarry phosphate was recovered. 1. Clays and clayey sands into which the channels of unit 2 are incised. PROVENANCE OF THE BAARD’S QUARRY FOSSILS Almost all the fossils from Baard’s Quarry were collected after the deposits in which they occurred had been moved by the mining operation and con- sequently the provenance of specimens has for the most part to be inferred. There are no definite records of fossils occurring in the surface sands (unit 4 of the succession), which are presumably of late Pleistocene/Holocene age. There are, however, a few unmineralized (sub-fossil) bones in the assemblage which probably came from this horizon. The material in question was excluded from the present study. Some fragmentary fossils were recovered from unit 3 during excavations carried out in the Baard’s Quarry area in 1965. No significant specimens are included in this material, but it is important since its very characteristic state of preservation is shared by some specimens in the original assemblage and it is presumed that they too were derived from unit 3. Significantly the only identi- fiable specimens in this part of the assemblage belong to species recorded else- Where in the south-western Cape Province from middle to late Pleistocene occurrences (vide infra). The unit 3 fossils are evidently appreciably younger than most of the material from Baard’s Quarry. The economically exploited unit 2 was apparently the source of the majority of the Baard’s Quarry fossils, including the first described fossil from this site (Singer & Hooijer 1958). This assemblage is comprised overwhelmingly of frag- mented bones and teeth, many of which show signs of rolling and which are presumably from the unit 2 river channel deposits. These fossils may have been derived from pre-existing deposits, they may have been contemporaneous with the deposition of the unit 2 sediments, or both. The mineralizing agent in these specimens is phosphate, while the unit 3 fossils are ferruginized. A few fragmentary fossils were recovered from unit | during the 1965 excavations, but it is unlikely that these deposits made a significant direct con- tribution to the original assemblage since the unit | deposits underlie the phosphatic horizon and consequently were not being mined. Nevertheless, part of the assemblage may have been from this horizon, while the unit 2 assemblage may include material derived from unit 1. It is worth noting that the 1965 excavations were sited near the southern margin of Baard’s Quarry, which had by then been back-filled, and that no phosphatic sandstone was encountered. No phosphatized fossils such as those which comprise the main body of Baard’s Quarry fossils were recovered from AGE OF THE FOSSILS FROM BAARD’S QUARRY, SOUTH AFRICA 5 the 1965 excavations, and this is taken as a further indication that the original material came largely from unit 2 rather than unit |. There are a few specimens in the original assemblage which are in an unusual state of preservation and which in appearance resemble neither the ferruginized unit 3 specimens nor the phosphatized unit 2 specimens. This material includes part of a series numbered SAM—PQ-L179, which were evidently found together. In this instance, at least, adherent matrix, which has a clayey component, sug- gests derivation from the lower levels in the succession. To sum up, the majority of specimens are here regarded as what is informally termed the ‘lower levels’ assemblage, that is, specimens from unit 2, but possibly from both | and 2, while a small number belong to the “upper level’ assemblage from unit 3. THE ‘UPPER LEVEL’ ASSEMBLAGE Only a few isolated teeth and tooth fragments belonging to the ‘upper level’ assemblage will be dealt with here, other available specimens being largely undiagnostic bone fragments. Equidae There are three teeth (L2095, L2106, L866) and three tooth fragments (L2111, L1527C and an unnumbered piece) belonging to Equus in the ‘upper level’ assemblage. Hipparion is not represented. The specimens L2106 and L2111, both M?’s, are here regarded as particularly significant since they can be com- pared with three fragmentary M?’s of Equus from the ‘lower levels’ assemblage (L2109, L1465E, L2129). Other Baard’s Quarry Equus teeth are no longer in the South African Museum and could not be examined. It must be stressed that Hooyer (1976a: 33-34, pl. 8) has already dealt with the specimens enumerated above. He found them unremarkable and all were provisionally assigned to Equus capensis. There is only one character, the size of the postfossette, which can be observed in all five M?’s comprising the two sets of specimens. The postfossettes of the “upper level’ specimens are of comparable size and larger than those of the ‘lower levels’ specimeus (Table 1). Otherwise comparisons must be confined to L2106 and L2109, which are the two most complete specimens. The former, TABLE | Dimensions of Equus third upper molars from Baard’s Quarry, Langebaanweg ‘upper level’ ‘lower levels’ L2106 L2111 L2109 L1405 12129 Ant.—post. diam. at occlusal surface. . . c. 39,0 — — — — Transv. diam. at occlusal surface . . . c. 31,0 ~ c. 29,0 _ -- Trans. diam. 20 mm from base of crown . 28,5 _ 26,5 — — Ant.—post. diam. of protocone 20 mm from base 16,5 — 13,0 — _ Mesostyle—metastyle length 20 mm from base 22,5 -— 18,0 — — Max. ant.—post. diam. of postfossette . . 17,6 16,1 UO @ WAYS 15,0 6 ANNALS OF THE SOUTH AFRICAN MUSEUM which is from the ‘upper level’, is larger than L2109 in all other dimensions as well. Morphologically L2106 and L2109 differ most strikingly in the degree of development of the metastyle/hypostyle. In L2106 this region of the tooth is particularly well developed, forming the most posterior margin of the tooth and being separated from the hypocone by a deeply indented and narrow hypoglyph. In L2109 the metastyle/hypostyle is less prominent and it is the hypocone which forms the posterior margin of the tooth, while the hypoglyph is shallower and more wide open than in corresponding parts of L2106. There are some differences in the enamel patterns of the occlusal surfaces of the teeth, but they are not recorded because equid teeth are notoriously variable in this respect. In the past significance has been attached to minor differences in South African fossil Equus teeth and this contributed to the chaotic state of the group’s nomenclature. It would therefore seem inadvisable to make much of the dif- ferences observed in the two sets of Equus teeth from Baard’s Quarry, especially since the specimens involved are fragmentary and few. On the other hand, since these two sets of specimens are from different levels in the succession, the observable differences could be of evolutionary significance. It now appears likely that in addition to the living zebras and their ancestors there was one lineage of large Equus present in Africa for much, if not all, of the Pleistocene. If this was indeed the case then it is likely that there were some changes in, for example, size and tooth characters in successive members of this lineage. Indeed, the middle Pleistocene large Eguus from Elandsfontein near Langebaanweg does differ in certain respects from the late Pleistocene (‘classic’) E. capensis from other occurrences in the south-western Cape, the most obvious difference being the larger size of the latter. The South African Museum’s collections include many M?’s belonging to the large middle Pleistocene Equus, and in respect of both size and morphology the ‘lower levels’ L2109 is generally similar to them, whereas L2106 is larger and, in respect of the metastyle/hypostyle-hypoglyph—hypocone development, mor- phologically distinct. Unfortunately there were no M®’s available of large Equus of undoubted late Pleistocene age, but the ‘upper level’ M? (L2095) and lower molar (L866) are comparable in size to corresponding late Pleistocene teeth and larger than ones of middle Pleistocene age. Inadequate though the evidence may be, it is tentatively concluded that the ‘upper level’ Equus is close to or identical with late Pleistocene E. capensis from the south-western Cape, whereas the ‘lower levels’ Equus is probably closer to but not identical with the middle Pleistocene E. cf. capensis from this region. The material is identified accordingly in Table 4. Bovidae The Baard’s Quarry Bovidae will be dealt with in more detail by A. W. Gentry (in prep.), but he has kindly given permission for observations on them to be included in this paper. AGE OF THE FOSSILS FROM BAARD’S QUARRY, SOUTH AFRICA 7 A series of specimens numbered L21 include two left and two right upper molars and a lower molar of a reduncine. These teeth are closely matched by specimens belonging to the late Pleistocene Redunca cf. arundinum from Swart- klip near Cape Town. The teeth L1491A, D, F and J appear to belong to Hippotragus gigas, a species known from the early Pleistocene of east Africa, but which survived later in South Africa, being included in the middle Pleistocene fauna from Elands- fontein and possibly in the late Pleistocene fauna from Florisbad in the Orange Free State. The teeth of the living gemsbok, Oryx gazella, resemble those of H. gigas and there is a possibility that the Baard’s Quarry specimens belong to a gemsbok. The teeth L2110 and L2129B belong to the recently extinct blue antelope, Hippotragus leucophaeus, a species which is known only from the middle to late Pleistocene and Holocene of the south-western and southern Cape Province (Klein 1974a). Two incomplete horn-cores (L1523, L1643A) and two unnumbered man- dible fragments belong to Raphicerus. The teeth resemble those of the Raphicerus from the late Pleistocene Swartklip fauna, they are smaller than those of the middle Pleistocene Raphicerus from Elandsfontein and much smaller than the teeth of the “E’ Quarry Raphicerus. The horn-cores are correspondingly small. The ‘upper level’ assemblage includes several alcelaphine teeth (L1491B, C and H, and possibly L1292), which represent at least two species. Although not identified, these specimens are more advanced than the ‘E’ Quarry alcelaphines and are likely to be Pleistocene in age. Age of the ‘upper level’ assemblage The ‘upper level’ fauna is clearly of Pleistocene age and indications are that it dates from the latter part of this epoch. THE ‘LOWER LEVELS’ ASSEMBLAGE Although a large amount of material is included in the ‘lower levels’ assemblage, the number of identifiable specimens is limited. Tubulidentata A single metacarpal IV (L908) is assigned to this order. The specimen is larger in most respects than the corresponding bones in the available series of four modern Orycteropus afer. It is particularly distinctive in its overall length (c. 53,5 mm), which contrasts with the range of 39,9 to 46,4 mm in the compara- tive series. No aardvark metacarpals are represented in the ‘E’ Quarry assemblages, but there are at least six other postcranial bones and an isolated tooth, all of which fall within the size range observed in the four O. afer specimens. Indications are therefore that the Baard’s Quarry aardvark was larger than that from ‘E’ Quarry 8 ANNALS OF THE SOUTH AFRICAN MUSEUM and was either not conspecific or represented a different variety of the same species. In either case it is unlikely that the two animals were contemporaneous. Carnivora A specimen tentatively identified as a jackal (?Canis sp.) (L1478) is included in the ‘lower levels’ assemblage and has already been described (Hendey 1974: 195). This animal is not conspecific and probably not even congeneric with the only canid recorded from ‘*E’ Quarry, which is an as yet undescribed species of Vulpes (Hendey 1976). A fragmentary upper canine (L1479) may belong to a lion-sized felid. Although the identification is uncertain, the animal concerned is not conspecific with any of the ‘E’ Quarry carnivores. If the above specimens do indeed belong to a jackal and lion, this would indicate that they postdate the ‘E’ Quarry faunas, since these animals are known in Africa only from late Pliocene or post-Pliocene contexts. Three carnivores are included in the L179 series of specimens which in respect of their preservation differ from most other specimens in the ‘lower levels’ assemblage (see p. 5). Two have already been described (Hendey 1974: 242, 279). They were identified as Mellivora cf. capensis (L179/12) and Hyaena cf. bellax (L179/11). Although these identifications are tentative, the species con- cerned are not conspecific with any from ‘E’ Quarry and the Mellivora at least is clearly more advanced than its ‘E’ Quarry counterpart. Both species suggest a Pleistocene date. The third carnivore is represented by a mandible fragment (L179/13) which may belong to a canid. The only other potentially useful ‘lower levels’ carnivore specimens are two proximal radius fragments (L1400, L1706) belonging to a pinniped. They are similar to the radii of the ‘E’ Quarry Prionodelphis capensis, but positive identifi- cation of such fragmentary specimens was not possible. Proboscidea The best proboscidean specimens from Baard’s Quarry were not available during the present study. This is unfortunate in view of the importance attached to proboscideans in faunal dating and the confusion which now exists in respect of the species represented in the Baard’s Quarry assemblage. The first fossil from the Langebaanweg area to be described was a fragmen- tary elephant molar, which is one of the specimens no longer in the South African Museum. It was originally referred to Stegolophodon (Singer & Hooijer 1958), but subsequently identified as Mammuthus subplanifrons (Maglio & Hendey 1970; Maglio 1973). The revised identification was ignored by Wolff et al. (1973: 209) who claimed that three species of ‘elephant’ are present at “the site’, the ‘site’ including both Baard’s and ‘E’ Quarries. In their table 1 they listed Stegolophodon, presumably meaning the material identified elsewhere as M. sub- planifrons, Archidiskodon, which is in part a junior synonym of Mammuthus (Maglio 1973), Stegodon, which is now generally assigned to the Stegodontidae rather than the Elephantidae, and Anancus, which is a gomphothere. AGE OF THE FOSSILS FROM BAARD’S QUARRY, SOUTH AFRICA 9 The issues which have therefore to be settled are the identity of the pro- boscideans from the Langebaanweg sites and the source of the material. Since the observations which follow are confined to material in the South African Museum and since this material now requires further detailed study by an authority on proboscideans, a final statement is not possible. Parts of a Mammuthus subplanifrons individual from the Quartzose Sand Member are the only described proboscidean material from ‘E’ Quarry (Maglio & Hendey 1970; Maglio 1973), although many additional specimens, mainly isolated teeth, apparently belonging to this species are now available. The undescribed specimens are from bed 3aN of the Pelletal Phosphorite Member and are therefore somewhat younger than the described material. An unidentified anancine gomphothere is also known from ‘E’ Quarry. It is represented by many isolated teeth from the Pelletal Phosphorite Member. The Baard’s Quarry proboscidean specimens in the South African Museum are all from the ‘lower levels’ assemblage. The material, which is very fragmen- tary, apparently belongs to only two species. As in the case of *E’ Quarry, one is identified with the elephant, M. subplanifrons, while the other is an unidentified anancine gomphothere. Other taxa may be represented amongst the material now housed elsewhere. : Nothing constructive emerged from the comparison between the Baard’s and ‘E’ Quarry M. subplanifrons specimens. None of the Baard’s Quarry tooth frag- ments is exactly matched by corresponding parts of the more complete ‘E’ Quarry specimens, but, since the latter exhibit appreciable variation, no significance could be attached to the differences observed. Although M. subplanifrons is represented at both sites it does not necessarily follow that the animals concerned were contemporaries, since there are indica- tions that the species had a long history in South Africa (see p. 20). The only Baard’s Quarry gomphothere specimen is an incomplete cheek- tooth, apparently a left M, (L1179) of which parts of three cone-pairs and the talonid are preserved. Although gomphothere M,’s from ‘E’ Quarry vary both in size and morphology, L1179 is distinct in having the preserved lingual cones transversely elongated and plate-like compared with the conical structures of the ‘E’ Quarry specimens (Fig. 1). This characteristic is also not evident in east African anancine gomphotheres, which survived until about 4 m.y. ago (Maglio 1974; Beden 1976), and it is regarded as an evolutionary advance which indicates that the Baard’s Quarry gomphothere is younger than those from east Africa and ‘E’ Quarry. Although not directly relevant, it is worth noting that Anancus survived into the Pleistocene in Europe (Kurtén 1968). The Baard’s and ‘E’ Quarry proboscidean assemblages thus have an elephant species in common and each has a gomphothere, with the species from Baard’s Quarry being the more advanced. In view of the differences between the gompho- theres and the possibility that M. subplanifrons had a long history in South Africa, there is no sound basis for concluding contemporaneity between the two faunas on the basis of the proboscideans represented. 10 ANNALS OF THE SOUTH AFRICAN MUSEUM fm a rman MU iy Fig. 1. Occlusal views of gomphothere ?M, fragment from Baard’s Quarry (L1179) (below) and gomphothere M, from ‘E’ Quarry (L41018) (above). Specimens are in a similar state of wear. Scale in millimetres. AGE OF THE FOSSILS FROM BAARD’S QUARRY, SOUTH AFRICA 1) Rhinocerotidae Rhinoceros tooth fragments are probably the most commonly represented identifiable element in the ‘lower levels’ assemblage. Hooijer (1976a) has recorded the presence of Diceros bicornis in this assemblage and most of the available specimens apparently belong to this species. A few fragments are, however, too high-crowned for Diceros and probably represent a Ceratotherium. D. bicornis is not known in Africa from occurrences older than 4 m.y., whereas the “E’ Quarry rhinoceros, Ceratotherium praecox, is known only from 4 to 7 m.y.-old contexts (Hooier 1976a). The Diceros therefore suggests that the ‘lower levels’ assemblage postdates that from ‘E’ Quarry. The ‘lower levels’ Ceratotherium is of little significance because it could as well represent the post-4 m.y. C. simum as the pre-4 m.y. C. praecox. Equidae Both Equus and Hipparion are represented in the ‘lower levels’ fauna. The former was dealt with earlier (p. 5) and all that need be added here is that the presence of Equus is one of the firmest indications that the ‘lower levels’ fauna postdates that from ‘E’ Quarry. Equus is apparently not known anywhere in Africa from occurrences older than about 2 m.y. (Hooijer 19765; Eisenmann 1976), although it was present in Europe about 3 m.y. ago (Kurtén 1968). Indications are therefore that the first appearance of Equus in Africa cannot predate 2 to 3 m.y. There are several issues relating to Hipparion from the Langebaanweg sites which have yet to be resolved (Hendey 1976; Hooijer 1976a), but the only one to be dealt with here is whether or not the *‘E’ and Baard’s Quarry Hipparion assemblages are likely to be contemporaneous. The present study was confined to the lower cheekteeth from the two sites. The study was commenced with these specimens and since they alone indicate that at least two distinct (i.e. non-contemporaneous) populations are represented, other available material was not examined. Material representing two individuals from ‘E’ Quarry are excluded from the results of the study presented here. The premolar row L25197 was excluded because it is clearly atypical of the Lange- baanweg assemblage as a whole and is the only material tentatively assigned to H. namaquense by Hooijer (1976a: 26). The Quartzose Sand Member dentition L13102 was excluded because the immaturity of the individual concerned made the teeth appear to differ in size from their Baard’s Quarry counterparts to a far greater extent than other teeth from ‘E’ Quarry. The comparison was thus con- fined to specimens of roughly comparable ontogenetic age. The lower cheekteeth from Baard’s Quarry differ most obviously from the ‘E’ Quarry specimens in the following respects: 1. The Baard’s Quarry teeth have more pointed metaconids and metastylids, which results in their having more prominent vertical ribs on their lingual surfaces. 2. The Baard’s Quarry teeth are in general larger than those from “E’ Quarry. 12 ANNALS OF THE SOUTH AFRICAN MUSEUM They also seem to have more wrinkled enamel, at least on their buccal surfaces, and a heavier covering of cement, but no attempt was made to establish these and other possible differences conclusively, the two listed above being con- sidered sufficient for the purposes of the present study. Hooijer (1974: 30) has indicated that the development of ‘pointed rather than rounded loops’ in the metaconid and metastylid is ‘a progressive feature’ in Hipparion. For example, in Miocene H. primigenium the loops are rounded, whereas in Pleistocene H. ethiopicum they are pointed (Hooijer 1975). In this respect the Baard’s Quarry teeth are clearly more advanced than those from ‘E’ Quarry, in which the metaconid 1s almost invariably rounded and the meta- stylid is occasionally rounded (Fig. 2). Hooijer (1975) has commented on the difficulties which can be experienced in obtaining measurements of Hipparion teeth which can be meaningfully com- pared irrespective of the ontogenetic age of the individuals concerned. His methods, with one modification, are followed. The breadth measurements given here are of tooth enamel and not of the enamel plus cement. Measurements given elsewhere may or may not include the cement, this being dependent on the con- dition and/or ontogenetic age of individual specimens. There were a few instances in the present study where breadth measurements might be fractionally higher because of a thin cement covering, but in no instances were dimensions affected significantly. Only in the case of the P,’s were the Baard’s and ‘E’ Quarry assemblages not clearly distinct from one another in terms of size (Tables 2, 3). This may in TABLE 2 Dimensions of the Baard’s Quarry Hipparion lower cheekteeth P, P;/P4 M,/M, M; l b l b ] b 1 b L1592/L1446B . . c. 29,0 13,5 = — 26,8 13,0 = — TO4TS 21 a Te S80 14,2 = — — — — ~— 189441695) = 27,6 16,0 25,9 14,0 — — 1946 ee = 27,0 16,1 — -- — — IENA6S Bs ae = 28,2 15,5 — _ — _ ISU, gg cS = 28,2 16,3 = _ — — E9431.) ae ie ee ~ VIS 16,8 = - — -- JUIN Bg = 6.935 @, 1165 — _ ~ — L957 = = = - 29,4 15,9 — - L954 _ = -- = 28,6 16,1 = oh L959* = = = = 25,3 14,4 — — L952 - = = = 25,9 St _ — L1448 = = = = D55) 15,1 = — L1447B = = = = 26,6 15,9 - — L1635A = = = = 27,9 16,1 = — M465P 3. A = = = _ 24,6 13,5 — — E4556 %.0-0 by © BS = a = = = 32,0 12D 52k Sg oe ee = = = = = 31,6 1 LOSS roth), ea = = = = - WS e120 * See Figure 2. AGE OF THE FOSSILS FROM BAARD’S QUARRY, SOUTH AFRICA 13 ALIANT ATTY ETT 3 4 = Fig. 2. Occlusal and lateral views of Hipparion molars from Baard’s Quarry (L959) (right) and *E’ Quarry (L41747) (left). The specimens are close to the mean sizes for M,,. of the two samples and are in a similar state of wear. Scale in millimetres. 14 ANNALS OF THE SOUTH AFRICAN MUSEUM TABLE 3 Dimensions of the ‘E’ Quarry Hipparion lower cheekteeth 1B iBs 1B M, M2 M3; 1 b ] b ] b ] b ] b ] b L10954) \ 28:6, 12:5 26:5" 1320 DS) WS7/ 22,4 1230 2138) 109° — L20541 26,6 12:6 25,0 1358 22,8 13,6 — -- L2546 = = 925.5) sk// 23,4 13,5 Ziel 3) == — — — LS 104 =| Byles) AS) 26,1 3) = = = — -- — 22 Oe = = — re. 2239 “el i2 er 165. ae. ES — — — 120555 ae = = a = = 2059 12,3 23,0) 120n233eiies Py P,/P, M,/M2 M; b l b b b LA0G19= Se ee 264 13,0 = = Dl 150 = _ E46140/1 es 2955 1455) = = 23,0 11,0 = — L12865/L13423 . . — = ASS) (C5 NWS) 24,5 tiES = — 1220259 XG 14,4 = = == = = — WSOC Ge ve = == 25,0 1135.3) a = = — LAVAS Ae ae = = == = papi WT = a LASA oe) 5 eae = = =- = 22,6 tie7 = = LAGIS9 2 Se Rete = = oa = A2)se) 1353 a = E2AGIOR ee Vee = = = = 24,2 1355 = = Te22547 boa ee we = = = = ES) 74.5) = = T2545 ae es ee = = -— = = = 26,5 10,8 L20907— ee — — _ — -- _ 28,6 10,0 * See Figure 2. part, or wholly, be due to the fact that in equids the P, is one of the least useful teeth for distinguishing species (Cooke 1950). There are very clear size differences between the other lower cheekteeth (Figs 2-3), the Baard’s Quarry teeth being the larger. Some overlap in size might have been observed had the two assem- blages been larger, but the general size difference is indisputable. The size difference coupled with an advance in tooth morphology indicates that two non-contemporaneous populations of Hipparion are represented and that the one represented at Baard’s Quarry is the younger. The taxonomic implications of this conclusion were not pursued and the arrangement of Hooijer (1976a), whereby the Baard’s Quarry material is identi- fied as H. baardi and that from ‘E’ Quarry as H. cf. baardi, is followed. Hippopotamidae A few fragmentary Hippopotamus teeth are included in the ‘lower levels’ assemblage, but they are inadequate for specific identification and no conclusion about their age relative to the ‘E’ Quarry Hippopotamus is possible. Giraffidae Giraffids are poorly represented in the Baard’s Quarry assemblage, but one useful specimen is a Sivatherium metatarsal of an immature individual (L637). This specimen resembles metatarsals of the ‘E’ Quarry Sivatherium in terms of overall length and is appreciably longer than a middle Pleistocene specimen from Elandsfontein (see Harris 1976). The implication is that the Baard’s Quarry AGE OF THE FOSSILS FROM BAARD’S QUARRY, SOUTH AFRICA 0 0 . BAARD’s © ft) BREADTH BAARD’S 14 = = i—} < tad - 3 t--] 25 LENGTH 0 BAARD'S 0 BREADTH Fig. 3. Dimensions of lower cheekteeth of Hipparion from Baard’s and ‘E’ Quarries. 15 16 ANNALS OF THE SOUTH AFRICAN MUSEUM Sivatherium was at a stage in evolution closer to a Pliocene representative of this genus than one of middle Pleistocene age. It does not necessarily follow that the ‘E’ and Baard’s Quarry sivatheres must have been contemporaries. It has already been suggested that elephants survived longer in a primitive state in South Africa during the Pliocene/early Pleistocene and presumably the same might have applied to other large herbi- vores such as sivatheres. Bovidae The ‘lower levels’ Bovidae will be dealt with in more detail by A. W. Gentry (in prep.), but some observations on them are included here. At least seven species are represented, namely, a boselaphine, a reduncine, two alcelaphines, two antilopines and a neotragine. A single horn-core fragment (L1588A) is similar in all observable respects with corresponding parts of Mesembriportax acrae horn-cores from ‘E’ Quarry and it is identified with this species (Gentry 1974). Although regarded as con- specific with the ‘E’ Quarry species, it is not necessarily contemporaneous. M. acrae is represented in ‘E’ Quarry by specimens from the Quartzose Sand Member as well as beds 3aS and 3aN of the Pelletal Phosphorite Member, so evidently the species was present in the area for what might have been an appreciable period. Several incomplete horn-cores (e.g. L565, L1521) and a lower molar (L1487) belong to a reduncine which appears to have affinities with reduncines from east Africa and India that are older than 5 m.y. It is definitely not conspecific with either of the ‘E’ Quarry reduncines. This species is problematical since it is the only one from Baard’s Quarry which suggests a late Miocene or early Pliocene age. Since it is apparently inconsistent with the ‘lower levels’ fauna as a whole, there must be some doubt about its relationships to the east African and Indian species. Certainly if the lower molar does belong with the horn-cores, the species cannot predate those from ‘E’ Quarry since the latter have more primitive teeth. The tooth, at least, is not inconsistent with the age of the ‘lower levels’ fauna inferred on other grounds. A fragmentary horn-core (L9) and several fragmentary teeth (e.g. L2112, L1491E) belong to at least two alcelaphine species. The horn-core is a significant specimen because it resembles specimens from ‘E’ Quarry and might be related to the species to which they belong. Both the ‘E’ Quarry alcelaphines are primi- tive members of lineages apparently unrepresented elsewhere in Africa. One of them, ‘species A’, is commonly represented in both bed 3aS and bed 3aN and certain of the specimens from 3aN have features in common with L9. There is reason to believe that the 3aN ‘species A’ is a more advanced variety than that from 3aS and it is possible that L9 belongs to a still more advanced stage of the same lineage. If this is so, then the time interval between deposition of 3aN and the “lower levels’ at Baard’s Quarry must have been far greater than that between deposition of 3aS and 3aN, because L9 is much more advanced than the 3aN AGE OF THE FOSSILS FROM BAARD’S QUARRY, SOUTH AFRICA 17 species A, whereas the latter is not appreciably different from the 3aS species A. The ‘lower levels’ alcelaphine teeth represent at least two species which are distinguished on the basis of size. The smaller specimens may belong to the same species as L9. They correspond in size to alcelaphine teeth from ‘E’ Quarry, but are more hypsodont and in this respect at least are more advanced. The larger ‘lower levels’ alcelaphine is not conspecific with either of the “E’ Quarry species. Several horn-cores (e.g. L1670, L1645) belong to a small but unidentified species of Raphicerus, which is not conspecific with the much larger ‘E’ Quarry Raphicerus. In terms of size at least these specimens correspond to those in the ‘upper level’ assemblage. Five fragmentary horn-cores belong to a Gazella, which once again is not conspecific with its ‘E’ Quarry counterpart, but which appears most similar to the east African late Pliocene/early Pleistocene G. praethomsoni. Included in the L179 series, referred to earlier, are right and left horn-cores (L179/8, L179/10) and three lower molars (L179/4C, F, G) belonging to an Antidorcas, a genus which in east Africa is recorded as early as the late Pliocene, but which in South Africa is known only from post-Pliocene contexts. In general, indications are that the ‘lower levels’ and ‘E’ Quarry bovid faunas were not contemporaneous and that the former is younger. Age of the ‘lower levels’ fauna The ‘lower levels’ and ‘E’ Quarry faunas have little in common and there are more differences than similarities between them (Table 4). The situation may be summed up as follows: 1. Six of the twenty-one ‘lower levels’ species are more advanced than their ‘E’? Quarry counterparts (Mellivora, gomphothere, Hipparion, one alcela- phine, Raphicerus, Gazella). 2. Six of the ‘lower levels’ species are not represented in ‘E’ Quarry and either definitely or probably suggest a younger date (?Canis, ?Panthera, Hyaena, Equus, Diceros, Antidorcas). 3. Three of the ‘lower levels’ species are either not conspecific with or sub- specifically distinct from their ‘E’ Quarry counterparts and suggest that the two faunas are not contemporaneous (Orycteropus, reduncine, one alcelaphine). 4. Four of the ‘lower levels’ species may or may not be conspecific with their ‘E’ Quarry counterparts (?Prionodelphis, Ceratotherium, Hippopotamus, Sivatherium). 5. Two of the ‘lower levels’ species are conspecific with species from ‘E’ Quarry (Mesembriportax, Mammuthus). On balance the indications are that the ‘lower levels’ fauna postdates those from ‘E’ Quarry and that a late Pliocene or early Pleistocene age applies. One possible complication with the ‘lower levels’ assemblage is that it may not represent a single fauna. This is unlikely because there are no instances apparent where individual taxa are represented by more than one evolutionary 18 ANNALS OF THE SOUTH AFRICAN MUSEUM TABLE 4 The mammalian faunas from Baard’s Quarry, Langebaanweg, with a list of the nearest counterparts in the ‘E’ Quarry faunas ‘E’ QUARRY BAARD’S QUARRY ‘lower levels’ ‘upper level’ Orycteropus sp. Orycteropus sp. — Vulpes sp. ?Canis sp. -- Mellivora aff. punjabiensis Mellivora cf. capensis Homotherium sp. ?Panthera sp. -- ‘Percrocuta’ australis Ayaena cf. bellax — Prionodelphis capensis ?Prionodelphis capensis _- Mammuthus subplanifrons Mammuthus subplanifrons — Gomphotheriidae Gomphothertidae — Ceratotherium praecox Ceratotherium sp. -- Diceros bicornis — Hipparion cf. baardi Hipparion baardi — o Equus cf. capensis Equus capensis Hippopotamus sp. Hippopotamus sp. — Sivatherium hendeyi Sivatherium sp. — Mesembriportax acrae Mesembriportax acrae — Reduncini (2 spp.) Reduncini Reduncini = an Hippotragus gigas or Oryx gazella — — Hippotragus leucophaeus Alcelaphini (2 spp.) Alcelaphini (2 spp.) Alcelaphini (2 spp.) Gazella cf. vanhoepeni Gazella cf. praethomsoni — _ Antidorcas sp. = Raphicerus sp. Raphicerus sp. Raphicerus sp. Norte: unclassified or incompletely classified taxa listed in more than one column are not necessarily identical. stage, while the combination of species is for the most part what might be expected in an African fauna of late Pliocene/early Pleistocene age. The fauna is, in fact, unusual in some respects when compared with Plio/Pleistocene faunas in east Africa, but its peculiarities can be explained by the existence in the far south of the continent of endemic species and/or late surviving primitive forms. This aspect of faunas in the south-western Cape Province will be dealt with again later (p. 20). DISCUSSION When the present,study on the age of the Baard’s Quarry fossils was com- menced, there appeared to be only three possibilities which needed to be con- sidered. They are: 1. The Baard’s and ‘E’ Quarry fossils are contemporaneous (Wolff et al. 1973). 2. Part of the Baard’s Quarry assemblage is contemporaneous with that from ‘E” Quarry (i.e. Pliocene), but the rest is younger (i.e. Pleistocene) (Hendey 1970a, 1972, 1974). 3. All the Baard’s Quarry fossils postdate those from ‘E’ Quarry (Hendey & Deacon 1977). AGE OF THE FOSSILS FROM BAARD’S QUARRY, SOUTH AFRICA 19 The existence of at least two faunal units in the Baard’s Quarry assemblage, the younger of which is Pleistocene in age (i.e. the ‘upper level’ fauna), must now be beyond doubt. Since this fauna is limited in size and not problematical in respect of its age relative to the ‘E’ Quarry faunas, it can be excluded from further consideration and attention can be devoted to the main body of fossils from Baard’s Quarry, namely, those constituting the ‘lower levels’ assemblage (Table 4). The first of the three alternatives listed above, which was in fact not adequately substantiated when it was supported by Wolff et al. (1973), can now be dismissed. Wolff et al. (1973: 212) quoted faunal lists in Hendey (1969, 1970a, 19706) and their own table | in concluding that, “Virtually all the taxa listed for Baard’s are known from “E”’ Quarry’. Their table | did not record the provenance of the species listed, while the faunal list in Hendey (19700) referred only to species from ‘E’ Quarry and differed in only minor respects from that in Hendey (1970a). Consequently their conclusion must have been based on the lists in Hendey (1969, 1970a). Reference to these lists reveals only two species common to the ‘E’ and Baard’s Quarry faunas, namely, ‘Hipparion albertense baardi’ and ‘Libytherium olduvaiense’, which had been included on the basis of publications by Boné & Singer (1965) and Singer & Boné (1960). The alleged similarities observed by Wolff et al. referred to taxa which were not identified, incompletely identified or tentatively identified, an exceedingly weak basis for correlation. Their conclusion could not have been based on a personal knowledge of all the material concerned, since none of them has seen the specimens collected at Langebaanweg since 1967, that is, the vast majority of specimens from ‘E’ Quarry. The present study has shown that there are indeed two species common to the Baard’s ‘lower levels’ and “E’ Quarry assemblages. Even if these two are combined with the four other taxa which might be conspecific, they still con- stitute only 29 per cent of the ‘lower levels’ fauna. A comparison between the thirty-one named species from Members B and G of the Shungura Formation in Ethiopia reveals that about 29 per cent are common to both and yet there is an age difference of about | m.y. between them (Coppens & Howell 1976). If there are still adherents to the first alternative listed above, their case will have to be restated in the light of information now available. The second and third alternatives can be considered together because the one is a modification of the other, the earlier view that the Baard’s Quarry assemblage included Pliocene and Pleistocene elements being replaced by the opinion that the former is rather of late Pliocene/early Pleistocene age. The change in opinion came about largely because certain specimens and species previously included with the Pleistocene element had, because of their state of preservation, to belong with other specimens and species which were regarded as Pliocene in age. In other words, it was the combination of ‘Pliocene’ and ‘Pleistocene’ species to form what is now termed the ‘lower levels’ fauna which led to the conclusion that the early element in the Baard’s Quarry assemblage must postdate the faunas from ‘E’ Quarry. 20 ANNALS OF THE SOUTH AFRICAN MUSEUM The combination of species in the ‘lower levels’ fauna is admittedly not paralleled by any other late Pliocene/early Pleistocene fauna recorded in Africa, but its peculiarities can be explained by the location of the site near the southern continental extremity in a distinct geographic region far from other contem- porary or near contemporary occurrences. At least two, and possibly three, of the taxa, namely, Hipparion baardi, Mesembriportax acrae and possibly the L9 alcelaphine, belong to lineages not represented further north in Africa during the Pliocene and early Pleistocene. They are, or appear to be, southern endemics. The potential for endemic speciation amongst the larger herbivores of the south- western and southern Cape (i.e. Cape Biotic Region) has been suggested by the late Pleistocene bovids of this region (Hendey 1974: 14-19; Klein 19746: 43). While endemism in the Cape fauna may be due simply to the region’s distinct climate and vegetation, the region does, in addition, have the potential to become zoogeographically isolated from the rest of the subcontinent through aridification of areas immediately adjacent to it (Hendey 1974). It is not known if the Cape Biotic Region existed in its present form during the Pliocene, but by this time the climatic stability of the Tertiary was giving way to the instability of the Quaternary. Consequently it may have been during the Pliocene that aridification on the subcontinent first resulted in isolation of the more southerly parts of South Africa. The free interchange of mammals between southern Africa and regions further north is likely to have been impeded not only by the Namib and Kalahari deserts and adjacent semi-arid areas in the south-west of the sub- continent, but also by large rivers such as the Zambezi and Limpopo which traverse the eastern parts of southern Africa (Maglio 1973: 34). Isolation, even if it is incomplete, increases the potential for the evolution of endemic species and may also allow certain taxa to survive longer than their counterparts else- where. The latter is believed to apply in the case of Mammuthus subplanifrons. The situation in respect of South African M. subplanifrons is complicated and unsatisfactory. Maglio (1973: 53) found this taxon to be ‘represented [by] a heterogeneous assemblage with primitive molars whose relationships can only be clarified by discovery of cranial material’. Furthermore, although Maglio (1973: fig. 15 and elsewhere) records no occurrence of M. subplanifrons younger than 4 m.y., the dating of South African material is insecure. For example, Maglio (1973) judged the ‘primitive’ Quartzose Sand Member M. subplanifrons to be between 4 and 4,5 m.y. old, but there is now at least a possibility that the younger, although apparently equally primitive, bed 3aN representative of this Species may have been less than 4 m.y. old (vide supra). In addition, Maglio’s suggestion that the ‘progressive’ M. subplanifrons from Virginia, Orange Free State, dates back at least 4 m.y. was surprising since this material was said to occur in association with artefacts (Meiring 1955). The supposed antiquity of the Virginia mammoth has made it unique in the Orange Free State, a province which has otherwise produced only middle Pleistocene or younger fossils from similar occurrences. Maglio’s suggestions about its age were sufficiently striking to prompt a study of its geological associations by Butzer (1973). This study did AGE OF THE FOSSILS FROM BAARD’S QUARRY, SOUTH AFRICA 21 not contradict the Pliocene date, but there must be some doubt as to whether the geological and geomorphological observations actually confirm the date. Certainly Butzer’s study did not confirm the presence of artefacts in the deposits, but even so the age of this specimen cannot be regarded as settled beyond doubt. Maglio (1973) indicated a time range of only 0,5 m.y. for M. subplanifrons, but found that the range for Loxodonta adaurora in east Africa ‘must be taken as at least two million years, with very little change in molar structure throughout this period’ (p. 25). It now appears likely that his figure is an overestimate since L. adaurora does not occur later than Member B in the Shunguru Formation (Beden 1976), but a range of about 1,5 m.y. for the species still applies. Presumably a time range of this order might also have applied to South African M. subplanifrons, especially since there is no evidence of more advanced elephant species on the subcontinent during the late Pliocene and early Pleistocene. With the exception of problematical, but nevertheless primitive specimens from the Vaal River Gravels, the only other Plio/Pleistocene elephant recorded from South Africa is an Elephas ekorensis from Bolt’s Farm, Transvaal, which Maglio (1973: 34) believes may indicate that ‘an earlier stage of the (E. ekorensis) lineage persisted in southern Africa long after it was succeeded by a more progressive stage further north’. If this was indeed the case, then the late survival in South Africa of primitive Mammuthus is also likely. The possible late survival of E. ekorensis in the Transvaal and M. subplani- frons in the Orange Free State suggests that the zoogeographic isolation referred to earlier may have encompassed more of South Africa than just the Cape Biotic Region as it is now defined (Hendey 1974). In spite of the many uncertainties which exist there do appear to be grounds for believing that early elephant evolution in South Africa may not have paral- leled the well-documented east African pattern. The presumption that all M. sub- planifrons on the subcontinent must be at least 4 m.y. old seems less warranted than to presume that this heterogeneous taxon had a long local history. At the very least, the issue cannot be regarded as settled. If M. subplanifrons did indeed survive longer in a primitive state in South Africa, the same might also have applied to other large herbivores such as Sivatherium. This would account for the fact that the Baard’s Quarry Sivatherium resembles the Pliocene S. hendeyi rather than the Pleistocene S. maurusium, at least in terms of limb-bone proportions. The implication is that during periods when the more southerly parts of South Africa were isolated the large herbivores showed little or no evolutionary change (i.e. they remained ‘primitive’), whereas the smaller ones did evolve but, being isolated from their counterparts further north, represented locally endemic lineages. There is, however, an anomaly in this hypothesis since one of the large herbivores from Baard’s Quarry, the gomphothere, apparently exhibits an evolutionary advance over its ‘E’ Quarry counterpart. Mammals such as carnivores, which are less influenced by environmental factors and less likely to have their dispersal impeded by such factors, were not 22 ANNALS OF THE SOUTH AFRICAN MUSEUM necessarily affected by the isolation of parts of the subcontinent. Since in this case the isolating mechanism was aridification, isolation in respect of the mam- malian fauna could never have been complete, with some faunal interchange likely and accelerating as conditions in the arid and semi-arid regions ameliorated. Since faunal dating of the Langebaanweg sites is based on correlations with the east African record, this dating is likely to be complicated by the regional peculiarities of the local fauna. Consequently particular significance is attached to the first appearances of taxa in east Africa. The southerly part of South Africa is unlikely to have been a significant centre of mammalian evolution, and terres- trial species can otherwise only have moved into the area from points north. For example, if Maglio’s (1973) interpretation and dating of the origin of the Elephantidae is correct, then the earliest record of Mammuthus subplanifrons in the Langebaanweg area (i.e. the “E’ Quarry Quartzose Sand Member specimen) cannot be older than 5 m.y. and is most likely to be in the 44,5 m.y. range as Maglio suggested. Similarly, if the Shungura Formation Member G record of TABLE 5 Relative age of deposits in the vicinity of Langebaanweg Estimated age in millions ‘E? QUARRY BAARD’S of years QUARRY Less than SURFACE BED OCCURRENCES* ‘UPPER LEVEL’ 1 m.y. (5) Interval similar to (4)? (6) Interval greater than (5) About 2 m.y. “LOWER LEVELS’ (4) Interval greater than (3) = PELLETAL PHOSPHORITE MEMBER (GaN) e) & | (3) Interval less than (2) << 2 PELLETAL PHOSPHORITE MEMBER (3aS) S . | (2) Interval of unknown duration (a4 About | QUARTZOSE SAND MEMBER 4,5 m.y. < = (1) Interval of unknown duration jad Not more = GRAVEL MEMBER than 5 m.y. * Several isolated occurrences, none of which is necessarily contemporaneous with the ‘upper level’ at Baard’s Quarry. AGE OF THE FOSSILS FROM BAARD’S QUARRY, SOUTH AFRICA pegs Equus is the earliest for this taxon in Africa, then the Baard’s Quarry ‘lower levels’ Equus cannot be older than about 2 m.y. As more is learnt of the origin and evolution of African mammals, it should become possible to refine the faunal dating of the Langebaanweg assemblages. The present view on the relative ages of deposits in the vicinity of Langebaanweg is summarized in Table 5. ACKNOWLEDGEMENTS I am indebted to Drs A. W. Gentry (British Museum (Natural History)), J. M. Harris (Kenya National Museum), D. A. Hooijer (Rijksmuseum van Natuurlijke Historie), R. G. Klein (University of Chicago) and E. Vrba (Trans- vaal Museum) for direct and indirect assistance in preparing the manuscript of this paper. Particular thanks are due to Dr Gentry who generously made avail- able unpublished information on Langebaanweg and other bovids. I thank Mr N. J. Eden and Miss M. Sank for the photographs and Mrs J. Wallendorf for typing the manuscript of this paper. The South African Museum’s Langebaanweg Research Project is supported by Chemfos Limited, the South African Council for Scientific and Industrial Research and the Wenner-Gren Foundation for Anthropological Research (Grant no. 2752-1834) and the assistance of these organizations is gratefully acknowledged. The management and staff, especially Mr G. Benfield, of Chemfos Ltd at Langebaanweg have been extremely generous in the support which they have provided. REFERENCES BEDEN, M. 1976. Proboscideans from the Omo Group Formations. Jn: COPPENS, Y. ef al., eds. Earliest man and environments in the Lake Rudolf Basin: 193-208. Chicago: University Press. Bone, E. L. & SINGER, R. 1965. Hipparion from Langebaanweg, Cape Province and a revision of the genus in Africa. Ann. S. Afr. Mus. 48: 273-397. ButTzer, K. W. 1973. On the geology of a Late Pliocene Mammuthus site, Virginia, Orange Free State. Navors. nas. Mus. Bloemfontein 2: 386-393. CooKE, H. B. S. 1950. A critical revision of the Perissodactyla of southern Africa. Ann. S. Afr. Mus, 21: 393-479. CopPENS, Y. & HOWELL, F. C. 1976. Mammalian faunas of the Omo Group: Distributional and biostratigraphic aspects. In: COPPENS, Y. et al., eds. Earliest man and environments in the Lake Rudolf Basin: 177-192. Chicago: University Press. EISENMANN, V. 1976. Equidae from the Shungura Formation. In: CoppeENs, Y. ef al., eds. Earliest man and environments in the Lake Rudolf Basin: 225-233. Chicago: University Press. GENTRY, A. W. 1974. A new genus and species of Pliocene boselaphine (Bovidae, Mammalia) from South Africa. Ann. S. Afr. Mus. 65: 145-188. Harris, J. M. 1976. Pliocene Giraffoidea (Mammalia, Artiodactyla) from the Cape Province. Ann. S. Afr. Mus. 69: 325-353. HENDEY, Q. B. 1969. Quaternary vertebrate fossil sites in the south-western Cape Province. S. Afr. archaeol. Bull. 24: 96-105. HENDEY, Q. B. 1970a. A review of the geology and palaeontology of the Plio/Pleistocene deposits at Langebaanweg, Cape Province. Ann. S. Afr. Mus. 56: 75-117. 24 ANNALS OF THE SOUTH AFRICAN MUSEUM HENDEY, Q. B. 19706. The age of the fossiliferous deposits at Langebaanweg, Cape Province. Ann. S. Afr. Mus. 56: 119-131. HENDEY, Q. B. 1972. Further observations on the age of the mammalian fauna from Langebaan- weg, Cape Province. Palaeoecol. Afr. 6: 172-175. HENDEY, Q. B. 1973. Fossil occurrences at Langebaanweg, Cape Province. Nature, Lond. 244: 13-14. HENDEY, Q. B. 1974. The late Cenozoic Carnivora of the south-western Cape Province. Ann. S. Afr. Mus. 63: 1-369. HENDEY, Q. B. 1976. The Pliocene fossil occurrences in ‘E’ Quarry, Langebaanweg, South Africa. Ann. S. Afr. Mus. 69: 215-247. HENDEY, Q. B. & DEACON, H. J. 1977. Studies in palaeontology and archaeology in the Saldanha Region. Trans. R. Soc. S. Afr. 42: 371-381. Hoover, D. A. 1974. Hipparions from the Late Miocene and Pliocene of Northwestern Kenya. Zool. Verhand. 134: 1-34. Hoover, D. A. 1975. Miocene to Pleistocene hipparions of Kenya, Tanzania and Ethiopia. Zool. Verhand. 142: 1-80. Hoover, D. A. 1976a. The Late Pliocene Equidae of Langebaanweg, Cape Province, South Africa. Zool. Verhand. 148: 1-39. Hoover, D. A. 197656. Evolution of the Perissodactyla of the Omo Group deposits. In: CopPeENs, Y. et al., eds. Earliest man and environments in the Lake Rudolf Basin: 209-213. Chicago: University Press. KLEIN, R. G. 1974a. On the taxonomic status, distribution and ecology of the blue antelope, Hippotragus leucophaeus (Pallas, 1766). Ann. S. Afr. Mus. 65: 99-143. KLEIN, R. G. 19746. A provisional statement on terminal Pleistocene mammalian extinctions in the Cape Biotic Region (Southern Cape Province, South Africa). S. Afr. archaeol. Soc. Goodwin Ser. 2: 39-45. KurtTENn, B. 1968. Pleistocene mammals of Europe. London: Weidenfeld & Nicolson. Mac io, V. J. 1973. Origin and evolution of the Elephantidae. Trans. Am. phil. Soc. 63: 1-126. Mac tio, V. J. 1974. A new proboscidean from the late Miocene of Kenya. Palaeontology 17: 699-705. MacLio, V. J. & HENDEY, Q. B. 1970. New evidence relating to the supposed stegolophodont ancestry of the Elephantidae. S. Afr. archaeol. Bull. 25: 85-87. MEIRING, A. J. D. 1955. Fossil proboscidean teeth and ulna from Virginia, O.F.S. Navors. nas. Mus. Bloemfontein 1: 187-201. SINGER, R. & Hoorer, D. A. 1958. A Stegolophodon from South Africa. Nature, Lond. 182: 101-102. SINGER, R. & BONE, E. L. 1960. Modern giraffes and the fossil giraffids of Africa. Ann. S. Afr. Mus. 45: 375-548. TANKARD, A. J. 1975. Varswater Formation of the Langebaanweg-—Saldanha area, Cape Province. Trans. geol. Soc. S. Afr. 77: 265-283. Wo rr, R. G., SINGER, R. & BisHop, W. W. 1973. Fossil bear (Agriotherium Wagner, 1837) from Langebaanweg, Cape Province, South Africa. Quaternaria 17: 209-236. —— =~ 6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature (particularly Articles 22 and 51). Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov., etc. An author’s name when cited must follow the name of the taxon without intervening punctuation and not be abbreviated; if the year is added, a comma must separate author’s name and year. The author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is transferred from its original genus. The name of a subsequent user of a scientific name must be separated from the scientific name by a colon. Synonymy arrangement should be according to chronology of names, i.e. all published scientific names by which the species previously has been designated are listed in chronological order, with all references to that name following in chronological order, e.g.: Family Nuculanidae Nuculana (Lembulus) bicuspidata (Gould, 1845) Figs 14-15A Nucula (Leda) bicuspidata Gould, 1845: 37. Leda plicifera A. Adams, 1856: 50. Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b). Nucula largillierti Philippi, 1861: 87. Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. 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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. 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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 “a 3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced with 2,5 cm margins all round. First lines of paragraphs should be indented. 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REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: ‘Smith (1969) describes .. .’ ‘Smith (1969: 36, fig. 16) describes...’ ‘As described (Smith 1969a, 1969b; Jones 1971)’ ‘As described (Haughton & Broom 1927)...’ ‘As described (Haughton et al. 1927)...’ Note: no comma separating name and year 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. & RaAFFy, 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. THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. (continued inside back cover) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 75 Band February 1978 Februarie ant? ee Deel REDESCRIPTION OF PARABATH YNOMUS NATALENSIS BARNARD (CRUSTACEA, ISOPODA, CIROLANIDAE) 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, 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.), 701-4), 8, 901-2, 7), 10(1-3), 11(1-2, 5, 7, t.—p.i.), 15(4—5), 24(2), 27, 31(1-3), 32(5), 33 Price of this part/Prys van hierdie deel R1,70 Trustees of the South African Museum © Trustees van die Suid-Afrikaanse Museum 1978 ISBN 0 908407 37 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 ——_ a REDESCRIPTION OF PARABATHYNOMUS NATALENSIS BARNARD (CRUSTACEA, ISOPODA, CIROLANIDAE) By BRIAN KENSLEY South African Museum, Cape Town (With 3 figures) [MS. accepted 8 November 1977} ABSTRACT The cirolanid Parabathynomus natalensis, described by Barnard in 1924 and since over- looked, is redescribed. The validity of the genus is confirmed, and the genus compared with the closely related Bathynomus. The species, known from two specimens, is recorded from Natal and southern Mozambique. CONTENTS PAGE Introductiony Bes ms) 2. 9 25 Systematic discussion. . . . 26 Acknowledgements . . . . 30 INCiCheEnceSmen enn) Been eoil INTRODUCTION Interest in the giant isopod genus Bathynomus Milne Edwards is stirred each time a specimen is caught. Holthuis & Mikulka (1972) provided an excellent résumé of taxonomic and biological work which has been published on the (then) four species of Bathynomus. Shih (1972) and Griffin (1975) each describe a new species and provide notes on the genus. Nowhere, however, is mention made of the closely related genus Parabathynomus Barnard. This is not surprising, con- sidering the relatively obscure journal in which it was published (Barnard 1924) and that Parabathynomus has been mentioned only once in passing (Barnard 1940) since its description. The author, in developing a catalogue of southern African marine isopods, thought it necessary to re-examine the type of Parabathynomus and felt that redescription would be useful. The holotype of Parabathynomus natalensis was caught by the Fisheries Research vessel Pickle in 1920, off the coast of Natal. After description it was deposited in the British Museum (Natural History). A second specimen was found amongst the isopods collected in 1964 on the eighth cruise of the Anton Bruun during the International Indian Ocean Expedition. AS Ann. S. Afr. Mus. 75 (2), 1978: 25-31, 3 figs. 26 ANNALS OF THE SOUTH AFRICAN MUSEUM SYSTEMATIC DISCUSSION Parabathynomus natalensis Barnard Figs 1-3 Parabathynomus natalensis Barnard, 1924: 2, figs 1-4; 1940: 491. Description Male. Body parallel-sided, dorsally strongly convex, three and one-eighth times longer than wide; dorsal surface closely and minutely pitted, pits almost coalescing to form short striae on lateral portion of pleonites. Cephalon somewhat sunken into pereonite* |; frontal margin between eyes straight, faint impressed line present; faint line running along upper margin of eye, continuous across front. Antennular bases almost completely visible dorsally. Frontal lamina elongate—pentagonal, narrow, completely separating antennal bases. Clypeus wider than upper lip, lateral margins sinuous, anteriorly bifid at lamina base. Upper lip broadly bilobed. Eyes narrowly oblong, hardly visible in dorsal view, occupying most of anterolateral margin of cephalon. Pereonite 1 posterolateral angle rounded. Faint ridge on posterior half of lateral pereonite, with oblique continuous ridge ventral to former ridge. Coxae articulated on pereonites 2 to 7. Posterolateral angle of pereonite 2 rounded, 3 quadrate, 4 to 7 becoming progressively more acute; curved ridge present on all coxae. Pleon consisting of five free pleonites plus pleotelson. Pleonite 1 narrow, lateral margin hardly visible beneath coxa of pereonite 7. Pleonites 2 to 4 subequal in length. Pleonite 2 laterally narrowly acute, pleonites 3 and 4 laterally acute but with posterior subapical margin convex; pleonite 5 medially longer than preceding segments, lateral margin not extending as far as preceding seg- ments, overlapped by pleonite 4. Pleonite 2 with ventrolateral part of segment expanded into strong, flattened spinose process. Pleotelson with median length equal to basal width, distal margin evenly convex, with few irregular, low crenu- lations; very faint mid-dorsal impressed line just visible. Antennular peduncle three-segmented, terminal segment with distal oblique constriction, flagellum of about forty articles. Antennal flagellum reaching to pereonite 6; peduncle five-segmented, two distal segments subequal in length, flagellum of about seventy articles. Mandibles with incisor process strongly sclerotized, tridentate, innermost tooth of left mandible acute, with median edge folded on itself; lacinia oval, with short, curved teeth on raised rim on margin; molar process blade-like, apically acute, with strong denticles on median margin; palp three-segmented, second segment longest, with distal patch of fine, dense setae; terminal segment curved, with row of fine setae and few elongate distal setae. First maxilla with inner ramus short, bearing one slender and. three stout spines, with distal halves bearing fine hairs; outer ramus broad, with strongly sclerotized curved spines. * The author prefers this spelling. —Ed. REDESCRIPTION OF PARABATHYNOMUS NATALENSIS BARNARD i, Fig. 1. Parabathynomus natalensis. Holotype in dorsal and lateral view. Maxilliped endite triangular in cross-section, with two coupling hooks and about six plumose setae distally; palp five-segmented, third segment longest and broadest, all segments fringed with setae. Pereopod | ambulatory, dactylus curved, shorter than propodus; latter curved, with three short spines on ventral margin; carpus about half length of propodus, with two strong ventrodistal spines; merus with margin bilobed, distal lobe carrying two spines, proximal lobe with three spines, dorsal margin distally produced beyond carpus, with row of six spines; ischium with strong spine on distodorsal corner. Pereopods 2 and 3 similar to pereopod 1. Pereopods 4 to 7 similar, dactylus about half length of propodus, latter narrow, with strong spine at mid ventral point; carpus, merus, and ischium bearing several spines and setae on distally expanded margins. 28 ANNALS OF THE SOUTH AFRICAN MUSEUM Penial rami widely separated on sternite of pereonite 7, flattened, oval in outline. Pleopod | peduncle with eight coupling hooks on inner margin, and rounded membranous epipodite; endopod longer than exopod, median margin straight, outer margin curved; exopod with median margin convex, with single branchial tuft near base. Pleopods 2 to 5 similar, with large branchial tuft on peduncle next to inser- tion of the appendage, plus smaller tuft on outer margin, hidden by longer peduncular tuft; tiny epipodite present; endopod with slender stylet attached Fig. 2. Parabathynomus natalensis. A. Left mandible. B. Right mandible. C. First maxilla. D. Second maxilla. E. Maxilliped. 20 REDESCRIPTION OF PARABATHYNOMUS NATALENSIS BARNARD GEEZ 2 =- ~--. wee We eee Fig. 3. Parabathynomus natalensis. A. Antennule. B. Antenna. C. Pereopod 1. D. Pereopod 7. E. Pleopod 2. F. Pleopod 1. G. Frontal lamina, clypeus, and upper lip. H. Ventrolateral view of pleonites 1-3. 30 ANNALS OF THE SOUTH AFRICAN MUSEUM proximally, just extending beyond apex of ramus; exopod slightly shorter than endopod, distally broadly rounded, with well-developed branchial tuft at base. Uropodal peduncle produced into acute lobe extending along inner margin of endopod; exopod narrow, leaf-shaped, with nine spines on outer margin, six on inner margin, plus numerous fine plumose setae; endopod broad, longer than exopod, median margin distally curved, with outer distal angle blunt, six short spines on outer and inner margins. Female. Agrees in all details with male. Material Holotype 3 British Museum (Natural History) No. 1937.11.10.67. Locality: 29°49’S 31°46’E, 766 m Dimensions: total length 81 mm greatest width (at pereonite 5) 26 mm © United States National Museum No. 170251 Locality: 25°12’S 34°04’E, 230-295 m (taken in shrimp trawl) Dimensions: total length 69 mm greatest width 24 mm Remarks Shih (1972) defined Bathynomus as cirolanid isopods having seven free pereo- nites, five free pleonites, a telson with a strongly toothed posterior margin, free coxae on pereonites 2 to 7, ambulatory pereopods and pleopods adapted for swimming and respiration. The respiratory function of the pleopods is sup- plemented by highly ramified branchial tufts on the endopods. This feature most readily distinguishes Bathynomus from the rest of the cirolanids. The number and position of these branchial tufts is the most important feature separating Bathy- nomus and Parabathynomus. The other differences are given in the following table. Bathynomus Parabathynomus Frontal lamina Broad, often pear- Narrowly pentagonal shaped or triangular Pleotelson. . Distal margin dentate Distal margin faintly crenulate BYcS eee aebnianeular Elongate-oblong Branchial tufts Arise on bases of pleo- Arise on bases of pleopodal exopods, podal endopods and on pleopodal peduncles ACKNOWLEDGEMENTS My thanks are due to Miss J. Ellis of the British Museum (Natural History) for the loan of the type specimen of Parabathynomus, to Dr F. Ferrari of the Smithsonian Sorting Centre for the second specimen, and to Dr T. E. Bowman of the Smithsonian Institution for reading the manuscript and giving critical comments. REDESCRIPTION OF PARABATHYNOMUS NATALENSIS BARNARD 31 REFERENCES BARNARD, K. H. 1924. Description of a new genus and species of Isopod crustacean belonging to the family Bathynomidae, procured in South African Marine Survey. Rep. Fish. mar. biol. Surv. Un. S. Afr. 4 special rep. 2: 1-4. BARNARD, K. H. 1940. Contributions to the crustacean fauna of South Africa. XII. Further additions to the Tanaidacea, Isopoda, and Amphipoda, together with keys for the identifi- cation of the hitherto recorded marine and freshwater species. Ann. S. Afr. Mus. 32: 381-543. GRIFFIN, D. J. G. 1975. A new giant deep-water isopod of the genus Bathynomus (Flabellifera: Cirolanidae) from Eastern Australia. Proc. Linn. Soc. N.S.W. 100: 103-109. Hottuuis, L. B. & MIKULKA, W. R. 1972. Notes on the deep-sea isopods of the genus Bathy- nomus A. Milne Edwards, 1879. Bull. mar. Sci. 22: 575-591. SHin, C. 1972. Notes on the giant isopod genus Bathynomus Milne Edwards, 1879, with description of a new species. Publs Seto mar. biol. Lab. 21: 31-42. eae a nn t en yi 7 t < i 4 1 : i " : - MP i; I ; ey 7 - ce > — 2 = my % e = { i ss 7 a af > a 4 b nat i -~ { = } = j i aa . . 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. 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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, it not preceded by initials or full names e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene (c) Scientific names, but not their vernacular derivatives e.g. Therocephalia, but therocephalian Punctuation should be loose, omitting all not strictly necessary Reference to the author should be expressed in the third person Roman numerals should be converted to arabic, except when forming part of the title of a book or article, such as ‘Revision of the Crustacea. Part VIII. The Amphipoda.’ : Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial capital letter, provided the same generic name is used consecutively. Name of new genus or species is not to be included in the title: it should be included in the abstract, counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts. BRIAN KENSLEY REDESCRIPTION OF PARABATHYNOMUS NATALENSIS BARNARD (CRUSTACEA, ISOPODA, CIROLANIDAE) JOLUME 75 PART 3 MARCH 1978 : ISSN 0303-2515 507,68 OO ee -rti“(CwmTCTClCmUr CC! i = q - = - 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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For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list o, Scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number (only if independently paged) in parentheses, pagination (first and last pages of article). Examples (note capitalization and punctuation) BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140. FIscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. Zen. 74: 627-634. 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. THEELE, 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 75 Band March 1978 Maart Part 3 Deel REDESCRIPTION OF THE AMPHIPOD, CALLIOPIELLA MICHAELSENI SCHELLENBERG FROM SOUTH AFRICA, WITH COMPARISON TO A NEW GENUS FROM THE PACIFIC OCEAN 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(i-3, 5-8), 3(1-2, 4-5, 8, t.—p.i.), 511-3, 5, 7-9), 6(1, t.-p.i.), 71-4), 8, 911-2, 7), 10(1-3), 11(1-2, 5, 7, t.—p.i.), 15(4—-5), 24(2), 27, 31(1-3), 32(5), 33 Price of this part/Prys van hierdie deel R1,70 ISBN 0 908407 39 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 REDESCRIPTION OF THE AMPHIPOD, CALLIOPIELLA MICHAELSENI SCHELLENBERG FROM SOUTH AFRICA, WITH COMPARISON TO A NEW GENUS FROM THE PACIFIC OCEAN By J. LAURENS BARNARD Department of Invertebrate Zoology, Smithsonian Institution, Washington (With 3 figures) [MS. accepted 23 August 1977] ABSTRACT Calliopiella michaelseni Schellenberg is redescribed and compared with the Oregonian species, Calliopiella pratti J. L. Barnard. The two taxa are found to be distinct generically: a new genus Callaska is established for C. pratti. CONTENTS PAGE Introduction .. ED ccs tl eS Calliopiella Schellenberg . : 6 Ut 35 Calliopiella michaelseni Schellenberg ent 5 BG Callaska, gen. nov. . ce ik. ie SS Callaska pratti (J. L. Barnard) . a ee O Acknowledgements . . bbe toate, RAO INGAAS YTS ET ne INTRODUCTION This redescription of Calliopiella michaelseni Schellenberg is from material deposited in the South African Museum, and the comparison with the Oregonian species, Calliopiella pratti J. L. Barnard, is from material deposited in the Smithsonian Institution. Calliopiella Schellenberg Calliopiella Schellenberg, 1925: 147. Diagnosis Head with lateral lobe, rostrum weak. Antennae short, antenna | scarcely shorter than antenna 2, peduncles short; accessory flagellum articulate, scale-like, with three to four apical setules; antenna | bearing aesthetascs, no calceoli in male. Epistome not produced, upper lip pyriform, not excavate below. Mandibu- lar molars triturative, raker rows and incisors well developed, mandibular palp 33 Ann. S. Afr. Mus. 75 (3), 1978: 33-40, 3 figs. 34 ANNALS OF THE SOUTH AFRICAN MUSEUM ordinary, article 3 scarcely falcate, with B, D, and E setae. Lower lip lacking inner lobes. Inner plate of maxilla 1 fully setose medially, palp 2-articulate, right and left sides alike. Inner plate of maxilla 2 with full oblique facial row of setae. Maxillipeds ordinary. Coxae short but overlapping, coxa | smaller than but not hidden by coxa 2, posterior margin of coxa 4 weakly excavate. Morphology of gnathopod | similar to gnathopod 2, latter scarcely enlarged, wrists short, poorly lobate, hands elongate, narrow, subchelate, palms short, excavate and armed with four to five giant defining spines, dactyls short, curved, slightly overlapping palms. Pereopods 3-4 ordinary, dactyls short, article 2 of pereopods 5—7 narrowly ovate, increasingly larger from front to rear members, pereopods in general poorly armed. Pleon ordinary, untoothed. Uropods | and 2 poorly armed, each ramus with conspicuous armaments all terminal; uropod 3 short, peduncle weakly elongate, rami very short, scarcely longer than peduncle, weakly paddle- shaped, inner larger than outer, armaments sparse. Telson linguiform, with ventral keel, appearing almost fleshy but articulate and movable, cleft or entire. Sexes alike. Type species Calliopiella michaelseni Schellenberg, 1925 (monotypy). Composition Unique. Relationship Calliopiella differs from all other calliopiids, pontogeneiids and eusirids in the unusually reduced uropod 3 and poor spination on uropods 1-3. In its gnatho- pods, coxae, body form, accessory flagellum and antennae it bears close resem- blance to Atylopsis Stebbing but lacks inner lobes on the lower lip. It differs from Bouvierella Chevreux in the larger gnathopods almost identical to each other and in the presence of an accessory flagellum. It differs from Calliopius Liljeborg in the absence of male calceoli, presence of an accessory flagellum, the slightly elongate peduncle of uropod 3, the poorly lobate wrists of the gnathopods, and complete absence of inner lobes on the lower lip. The short gnathopodal wrists also distinguish Calliopiella from Cleippides Boeck, Apherusa Walker, Halirages Boeck and Liouvillea Chevreux. There is considerable resemblance to Atyloella Schellenberg and Liouvillea Chevreux, but Calliopiella lacks inner lobes on the lower lip, has a much more densely setose maxilla 1 (Atyloella), and lacks cusps on the epistome and head. Uropod 3 of Bovallia Pfeffer tends to approximate that in Calliopiella but otherwise the presence of an accessory flagellum and the elongate hands on the gnathopods distinguish Calliopiella. Eusiroides Haswell bears inner lobes on the lower lip, poorly setose maxilla 1 and elongate accessory flagellum. Harpinioidella Schellenberg has the first two characters mentioned for Eusiroides. 35 REDESCRIPTION OF THE AMPHIPOD CALLIOPIELLA MICHAELSENI SCHELLENBERG ™ ~~ S ss SEW s x N N S d 4 hi WU hy Von fh a, tl Fig. 1. Calliopiella michaelseni Schellenberg. a, female ‘a’ 10,32 mm; m, male ‘m’ 10,66 mm; g, male ‘g’? 7,22 mm. A. Lateral aspect. B. Right lacinia mobilis. C. Head. D. Left mandible. E. Head. F. Inner plate of maxilliped. G. Right mandibular apex. H. Prebuccal, anterior. I. Lower lip. J. Maxillipedal dactyl. K. Maxilliped. 36 ANNALS OF THE SOUTH AFRICAN MUSEUM Calliopiella might be an apomorph within the taxal group loosely charac- terized as the Paramoera—Pontogeneia complex, but differs from those many species in the peculiar uropods and the specially armed gnathopods. Calliopiella michaelseni Schellenberg Calliopiella michaelseni Schellenberg, 1925: 147-148; K. H. Barnard, 1940: 451-452, fig. 24; Vader, 1972: 14; Griffiths, 1974: 180-181; Griffiths, 1975: 118. Material South Africa, Cape Peninsula, 2 July 1971, female ‘a’ 10,32 mm, male ‘m’ 10,66 mm and 2 other specimens (SAM-—A15677); Schaapen Island, Saldanha Bay, 23 September 1957, LB 511, general collection, male ‘g’ 7,22 mm (SAM-A15678) and 2 other specimens from the same locality. Description With the characters of the generic diagnosis and accompanying illustrations. Figured female ‘a’ with 21 articles in primary flagellum of antenna 1, 22 articles in flagellum of antenna 2; aesthetascs of antenna 1 simple, example of formula, proximal to distal on female ‘a’ = 2-0-2-0 . . . (2-0 x 8)-0, on male ‘g’ = (1-2)-0-2-0-2-0-2-0-1-—0-2-0-2-0-2-0-0-0, parentheses indicating article 1 with one ventral and 2 apical aesthetascs. Right mandibular raker spines six plus one rudimentary spine, article 2 of palp with six setal spines. Cuticle covered with polygons, outlines clear and enclosing patches of densely villose surface, knobs irregularly scattered near bases of appendages, then lined up in rows near middle of appendages, then polygonal outlines lost near apices of appendages with knobs coalesced into striations, latter texture dominant on uropods and antennae. Each of coxae 2-7 with ovate, simple gill, scarcely pediculate. Brood plates very broad, fully setose, setae weakly bifid apically. Intraspecific variability Two sets of specimens from different sources indicate strong intraspecific phenotypy, or possible microspeciation: Set I: Cape Peninsula, specimens larger, eyes indiscernible, accessory flagellum lacking basolateral setule, right lacinia mobilis less complex (illus- trated), gnathopods with only two pairs of spines on palm, article 2 of pereopods 5—7 slightly thinner than in other group, epimeron 2 smaller, uropod 3 lacking apicomedial spine on peduncle, telson entire or scarcely excavate. Set 2: Saldanha Bay, specimens smaller, eyes well developed, accessory flagellum with basolateral setule, right lacinia mobilis more complex (illustrated), gnathopods with two triads of spines on palm, article 2 of pereopods 5-7 slightly stouter than in other group, epimeron 2 larger, uropod 3 bearing apicomedial spine on peduncle, telson distinctly notched. REDESCRIPTION OF THE AMPHIPOD CALLIOPIELLA MICHAELSENI SCHELLENBERG 37 Fig. 2. Calliopiella michaelseni Schellenberg. a, female ‘a’ 10,32 mm; g, male ‘g’ 7,22 mm. A. Right gnathopod 2. B-C. Accessory flagellum. D. Epimera 1-3. E. Apex of pereopod 3. F. Right gnathopod 2. G. Cuticle of coxa 5. H. Maxilla1. I. Right gnathopod 1. J. Mandibular palp. K. Maxilla 2. L. Right gnathopod 2, setae removed. 38 ANNALS OF THE SOUTH AFRICAN MUSEUM Some of the characters of set 2, the smaller specimens, do not correlate with size in terms of normal amphipodan development, for example, the more numerous gnathopodal spines and better development of accessory flagellum are out of phase with the larger specimens. The species is known to occur with various species of Patella so that these may be host-induced differences, though a very detailed study should be made on this problem. Illustrations Uropods 1-3 of male “b’ in dissected view magnified to same extent; uropod 3 of male ‘g’ drawn in situ; one row of apical setae omitted on outer plate of maxilla 2. Distribution Southern Africa, Cape Agulhas to South West Africa, intertidal, found under limpets, Patella spp., apparently obligatorily inquilinous. Callaska gen. nov. Etymology Contrived, feminine. Diagnosis Head with weak lateral lobe, rostrum well developed. Antennae of moderate length, antenna | shorter than antenna 2, peduncles short; accessory flagellum articulate, scale-like, very short, with three apical setules; antenna 1 bearing aesthetascs, no calceoli in male. Epistome unproduced, upper lip pyriform, not excavate below. Mandibular molars triturative, raker rows and incisors well developed, mandibular palp ordinary, article 3 not falcate, with B, D and E setae. Lower lip lacking inner lobes. Inner plate of maxilla 1 bearing only about four apical setae, palp 2-articulate, right and left sides alike. Inner plate of maxilla 2 with oblique partially facial set of about three setae. Maxillipeds ordinary. Coxae of medium extent, overlapping, coxa 1 smaller than but not hidden by coxa 2, posterior margin of coxa 4 weakly excavate. Morphology of gnathopod 1 similar to gnathopod 2 in female, distinctive in male, wrists short, poorly lobate, hands weakly elongate, narrow in female, broader in male, palms short except on male gnathopod 1, armed with two to three defining spines in female, four to five in male, dactyls short in female, longer in male, scarcely overlapping palms, gnathopod | in male larger than gnathopod 2. Pereopods 3-4 ordinary, dactyls of moderate length, article 2 of pereopods 5-7 expanded, ovate, increasingly larger from front to rear members, pereopods well armed (for the family group). Uropods 1 and 2 well armed; uropod 3 of medium length, peduncle elongate, rami subequal to or shorter than peduncle, lanceolate, bearing spines, no setae. Telson linguiform, laminar, movable, uncleft. REDESCRIPTION OF THE AMPHIPOD CALLIOPIELLA MICHAELSENI SCHELLENBERG 39 Type species Calliopiella pratti J. L. Barnard, 1954 (here selected). Composition Unique. Relationship Callaska pratti originally was placed in the genus Calliopiella but examina- tion of the type species of Calliopiella demonstrates adequate generic distinctions. Calliopiella is an unusual member of its family group in the loss of armaments on uropods 1-3, and the short rami of uropods I-3. In terms of uropods 1-3 Callaska is an ordinary member of the family but in which male gnathopod 1 is larger Fig. 3. Calliopiella michaelseni Schellenberg. a, female ‘a’ 10,32 mm; g, male ‘g’ 7,22 mm; m, male ‘m’ 10,66mm. A-B. Uropod 3, left. C-D. Telson. E-F. Uropod 1, left. G-H. Uropod 2 left. I. Pleon. 40 ANNALS OF THE SOUTH AFRICAN MUSEUM than gnathopod 2, a feature weakly developed in the allied Oligochinus J. L. Bar- nard from the same coastline. Callaska differs from Oligochinus in the elongate peduncle of uropod 3 and the uncleft telson, a variable character in the isolated South African Calliopiella but of firmer stability in calliopiids from other areas of the world. The short wrists of the gnathopods distinguish Callaska from a variety of otherwise similar genera such as Bouvierella Chevreux and Apherusa Walker. The latter genus also has weak inner lobes on the lower lip and Bouvierella lacks an accessory flagellum and has an elongate antenna 1. Calliopius Liljeborg lacks an accessory flagellum and bears inner lobes on the lower lip. Pontogeneoides Nicholls bears inner lobes on the lower lip, gnatho- pod 2 of the male is enlarged and the wrist lobate and the telson is elongate. Pontogeneia Boeck and Pseudopontogeneia Oldevig have deeply cleft telsons. Atylopsis Stebbing has inner lobes on the lower lip and unenlarged gnathopod 1. Dautzenbergia Chevreux has enlarged gnathopod 2 in the male and lacks an accessory flagellum. Callaska pratti (J. L. Barnard) Calliopiella pratti J. L. Barnard, 1954: 6-7, pls 6-8; J. L. Barnard, 1969: 95, figs 7-8. Distribution Coos Bay, Oregon to Goleta, California, intertidal. ACKNOWLEDGEMENTS I thank Dr C. L. Griffiths of the C.S.I.R. Oceanographic Research Unit, University of Cape Town, for identifying and lending me materials of C. michael- seni, 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. REFERENCES BARNARD, J. L. 1954. Marine Amphipoda of Oregon. Ore. St. Monogr. Stud. Zool. 8: 1-103. BARNARD, J. L. 1969. Gammaridean Amphipoda of the rocky intertidal of California: Monterey Bay to La Jolla. Bull. U. S. natn. Mus. 258: 1-230. BARNARD, K. H. 1940. Contributions to the crustacean fauna of South Africa. XII. Further additions to the Tanaidacea, Isopoda, and Amphipoda, together with keys for the identifi- cation of hitherto recorded marine and fresh-water species. Ann. S. Afr. Mus. 32: 381-543. GriFFITHS, C. L. 1974. The Amphipoda of southern Africa part 2. The Gammaridea and Caprellidea of South West Africa south of 20° S. Ann. S. Afr. Mus. 62: 169-208. 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. 67: 91-181. SCHELLENBERG, A. 1925. Crustacea VIII: Amphipoda. Jn MICHAELSEN, W. Beitrdge zur Kenntnis der Meeresfauna Westafrikas 3: 111-204. VADER, W. 1972. Associations between amphipods and molluscs. A review of published records. Sarsia 48: 13-18. $$ ——— 6. SYSTEMATIC papers must conform to the International code of zoological nomenclature (particularly Articles 22 and 51). Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov., etc. An author’s name when cited must follow the name of the taxon without intervening punctuation and not be abbreviated; if the year is added, a comma must separate author’s name and year. The author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is transferred from its original genus. The name of a subsequent user of a scientific name must be separated from the scientific name by a colon. Synonymy arrangement should be according to chronology of names, i.e. all published scientific names by which the species previously has been designated are listed in chronological order, with all references to that name following in chronological order, e.g.: Family Nuculanidae Nuculana (Lembulus) bicuspidata (Gould, 1845) Figs 14-15A Nucula (Leda) bicuspidata Gould, 1845: 37. Leda plicifera A. Adams, 1856: 50. Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b). Nucula largillierti Philippi, 1861: 87. Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. Note punctuation in the above example: comma separates author’s name and year semicolon separates more than one reference by the same author full stop separates references by different authors figures of plates are enclosed in parentheses to distinguish them from text-figures dash, not comma, separates consecutive numbers Synonymy arrangement according to chronology of bibliographic references, whereby the year is placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable. In describing new species, one specimen must be designated as the holotype; other speci- mens mentioned in the original description are to be designated paratypes; additional material not regarded as paratypes should be listed separately. The complete data (registration number, depository, description of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.: Poleupe 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. J. LAURENS BARNARD REDESCRIPTION OF THE AMPHIPOD, CALLIOPIELLA MICHAELSENI SCHELLENBERG FROM SOUTH AFRICA, WITH COMPARISON TO A NEW GENUS FROM THE PACIFIC OCEAN VOLUME 75 PART 4 FEBRUARY 1978 : ISSN 0303-2515 $07.63 OF THE SOUTH AFRICAN CAPE TOWN INSTRUCTIONS TO AUTHORS 1. MATERIAL should be original and not published elsewhere, in whole or in part. 2. LAYOUT should be as follows: (a) Centred masthead to consist of Title: informative but concise, without abbreviations and not including the names of new genera or species Author’s(s’) name(s) Address(es) of author(s) (institution where work was carried out) Number of illustrations (figures, enumerated maps and tables, in this order) (b) Abstract of not more than 200 words, intelligible to the reader without reference to the text (c) Table of contents giving hierarchy of headings and subheadings (d) Introduction (e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents (f) Summary, if paper is lengthy (g) Acknowledgements (h) References (i) Abbreviations, where these are numerous 3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced with 2,5 cm margins all round. First lines of paragraphs should be indented. Tables and a list of legends for illustrations should be typed separately, their positions indicated in the text. All pages should be numbered consecutively. Major headings of the paper are centred capitals; first subheadings are shouldered small capitals; second subheadings are shouldered italics; third subheadings are indented, shouldered italics. Further subdivisions should be avoided, as also enumeration (never roman numerals) of headings and abbreviations. Footnotes should be avoided unless they are short and essential. Only generic and specific names should be underlined to indicate italics; all other marking up should be left to editor and publisher. 4. ILLUSTRATIONS should be reducible to a size not exceeding 12 x 18 cm (19 cm including legend); the reduction or enlargement required should be indicated; originals larger than 35 x 47 cm should not be submitted; photographs should be rectangular in shape and final size. 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REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: ‘Smith (1969) describes...” ‘Smith (1969: 36, fig. 16) describes...’ “As described (Smith 1969a, 19695; Jones 1971)’ ‘As described (Haughton & Broom 1927)...’ ‘As described (Haughton et ai. 1927)...’ Note: no comma separating name and year Dagination indicated by colon, not p. names of joint authors connected by ampersand et al. in text for more than two joint authors, but names of all authors given in list of references. (b) Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, b, etc. to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a). For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list o, Scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number (only if independently paged) in parentheses, pagination (first and last pages of article). Examples (note capitalization and punctuation) BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140. FIscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. gén. 74: 627-634. Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320. Konn, A. J. 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) a ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 75 Band February 1978 Februarie Part 4 Deel A NEW MARINE ISOPOD FAMILY FROM THE SOUTH-WESTERN INDIAN OCEAN 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.), 5(1-3, 5, 7-9), 6(1, t.—p.i.), 711-4), 8, 911-2, 7), 10(1-3), 11(1-2, 5, 7, t—p.i.), 15(4-5), 24(2), 27, 311-3), 32(5), 33 Price of this part/Prys van hierdie deel R1,80 Trustees of the South African Museum © Trustees van die Suid-Afrikaanse Museum 1978 ISBN 0 908407 38 6 Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk., Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap A NEW MARINE ISOPOD FAMILY FROM THE SOUTH-WESTERN INDIAN OCEAN By BRIAN KENSLEY South African Museum, Cape Town (With 5 figures) [MS. accepted 30 November 1977] ABSTRACT A new family, Bathynataliidae, is created for Bathynatalia gilchristi Barnard, a marine isopod crustacean species from deep water off Natal. CONTENTS PAGE lintiroalieti@mn 5 « 6 .o 6 o 4 Systematic discussion . . . 41 Acknowledgements . . . . S50 INCiChENCES Bie Le) LO INTRODUCTION Barnard (1957) described Bathynatalia gilchristi from a single female taken by the R/V Pieter Faure in 880 metres off Durban, South Africa. With the brief description, Barnard (1957: 814) stated: ‘Taxonomic position doubtful. Super- ficially somewhat Serolid in appearance, but the similarities and the differences are equally apparent.’ Now that nine specimens including both sexes are avail- able, a fuller description can be given and the taxonomic position of this unusual species can be discussed. Several of Barnard’s statements can be amended and his diagnosis of the genus expanded. SYSTEMATIC DISCUSSION Family Bathynataliidae fam. nov. Diagnosis Body dorsoventrally flattened. Cephalon anterolaterally expanded, fused with pereonitr 1 medially, separated by a deep lateral slit. Pereonites 2 to 7 free, with articulating coxae. Pleon consisting of five pleonites (although these are quite distinct, there seems to be little movement possible between them) plus oval 41 Ann. S. Afr. Mus. 75 (4), 1978: 41-50, 5 figs. 42 ANNALS OF THE SOUTH AFRICAN MUSEUM pleotelson; only pleonites 2 and 3 with elongate lateral extensions. Antennule with 4, antenna with 5 peduncular segments, both flagella multiarticulate. Mandible lacking molar process; incisor broad, lacinia present on right side only; palp 3-segmented. Maxilla 1 curved, armed apically with cluster of spines. Maxilla 2 bilobed. Maxilliped with 3-segmented palp, broad endite, triangular exopod. Pereopod 1 robust and subchelate in both sexes, pereopods 2 to 7 slender, ambulatory in both sexes. Pleopod | indurate, exopod and endopod lying parallel, operculate over branchial chamber. Pleopods 2 to 5 biramous, mem- branous. Uropod subterminal, consisting of single segment, apically obscurely trilobed. Remarks Using several keys to the suborders and families of marine isopods (Barnard 1940; Menzies 1962; Schultz 1969), no family (or suborder) can be confidently arrived at for the present material. The species obviously does not belong to the Epicaridea (not being a parasite), the Anthuridea (not having an elongate body- form), the Gnathiidea (not having the characteristic sexual dimorphism of the gnathiids), the Asellota (lacking the characteristic pleopods of this group), or the Valvifera (not having valvate uropods). The pleopods, general disposition of the cutting and biting mouthparts, and the uropods, if one considers them to form a caudal fan with the pleotelson, indicate a flabelliferan affinity. The present material, however, cannot be fitted into any of the existing flabelliferan families, but would seem to be closest to the Serolidae. The similarities include the medial fusion of the cephalon and pereonite 1, both of which are laterally expanded, the structure of the multiarticulate antennules and antennae, the form of the broad incisor of the mandible, the presence of a large lacinia (albeit only on one side), the 3-segmented mandibular palp, the first maxilla (almost identical to such forms as Serolis completa Moreira 1971), the bilobed second maxilla (as seen in Serolis veaperta Moreira 1971), and the maxilliped structure. Several differences, however, immediately separate the Serolidae from the present material, and necessitate the erection of a new family for the latter. These differences include the second pereopods of the male (subchelate in Serolidae, ambulatory in Bathynatalia), the presence of five rather than three free pleonites, the operculate and non-setose first pleopod of Bathynatalia, the fourth and fifth pairs of pleopods (large and operculate in the serolids, delicate and membranous in Bathynatalia). Probably the most distinctive feature of Bathynatalia, and the most important difference between it and the Serolidae and all other flabelliferan families, is in the uropod. This clavate structure seems to represent an elongate peduncle, with the obscurely trilobed apex possibly representing the apex of the peduncle plus the bases of the rami, represented by two tiny, rounded processes. This interpretation is further supported by the presence of two narrow muscle- bands running the length of the appendage to two of the three apical lobes. The Bathynataliidae represents the only endemic marine isopod family from the southern African region. A NEW MARINE ISOPOD FAMILY FROM THE SOUTH-WESTERN INDIAN OCEAN 43 Genus Bathynatalia Barnard Bathynatalia Barnard, 1957: 814. Diagnosis As for the Family. Gender Feminine. Type-species Bathynatalia gilchristi Barnard, 1957. Bathynatalia gilchristi Barnard Figs 1-5 Bathynatalia gilchristi Barnard, 1957: 814, figs 2-4. Description Female Integument indurate, bearing numerous fine setules plus numerous elongate plumose setae amongst irregularly scattered tiny tubercles. Body broadest at pereonites 3 and 4. Cephalon lacking eyes; anterior margin hollowed to receive contiguous antennal bases, with low rostrum; anterolateral part lamellar, apically acute; posterior cephalon dorsally convex, irregularly tuberculate. Cephalon and pereonite 1 fused medially, fusion line marked by shallow groove; laterally separated by deep slit. Fused pereonite 1 expanded laterally into bilobed lamella; anterior part broadly rounded and contiguous with cephalon, posterior part triangularly acute; two middorsal tubercles present. Pereonites 2 to 4 similar, each with strong middorsal tubercle and elongate articulated coxae. Pereonites 5 to 6 narrower than preceding pereonites, lacking middorsal tubercles, with elongate coxae. Pereonite 7 very narrow middorsally, laterally acute but not elongate. Pleon of five pleonites plus pleotelson; pleonite 1 lacking free lateral mar- gins, pleonites 2 and 3 laterally produced into elongate posteriorly-directed processes; pleonites 4 and 5 short and narrow, lacking free lateral margins, pleonite 4 with middorsal tubercle. Pleotelson longer than broad, with globular proximocentral part bearing reticulate ridges and rounded tubercles, with lateral ridge running from proximal margin to strong distolateral spine, latter situated lateral to uropodal articulation, distal part of pleotelson triangular, lamellar, apically narrowly rounded; ventrolateral margins of pleotelson flexed towards midline and meeting pleopod 1. Antennular peduncle 4-segmented, segments 1 and 2 subequal in length, = =) aa 2) 2) = Z < 16) ea i, < x a =) fe) N aay an) ol ea (e) Nn 4] < Zz Z < Fig. 1. Bathynatalia gilchristi, 2 in dorsal view. A NEW MARINE ISOPOD FAMILY FROM THE SOUTH-WESTERN INDIAN OCEAN 45 Hir\\\ \I Fig. 2. Bathynatalia gilchristi. A. Antenna. B. Antennule. C. Pereopod 7. segment 3 slightly longer and more slender, segment 4 one-fifth length of third; flagellum of twelve articles. Antennal peduncle 5-segmented, segments | and 2 broad, 3 and 4 more slender, subequal in length, segment 5 one and one-half times longer than 4; flagellum of eleven articles. Mandibles indurate; palp 3-segmented, first segment slightly less than half length of second, latter with eight distal finely-fringed spines, distal segment slender, one-third length of second, with seven distal finely-fringed spines; left mandibular incisor of four broad teeth separated by small gap from short conical spine; right mandible with incisor of four teeth separated by small gap from 46 ANNALS OF THE SOUTH AFRICAN MUSEUM | Ul / ; i \w li Wi (| Wu, \W \i7 ne. i \ly Ww We \w \i We wh WY au AW Ww Ml S WP ly, wy ly ZZ; lw us (i Y ay We te we & \ \le Win Wy & Wee Wie oe Fig. 3. Bathynatalia gilchristi. A. Left mandible. B. Apex of right mandible. C. Apex of left mandible. D. Maxilla 2. E. Maxilliped. F. Maxilla 1. G. Apex of maxilla 1. a A NEW MARINE ISOPOD FAMILY FROM THE SOUTH-WESTERN INDIAN OCEAN 47 — Fig. 4. Bathynatalia gilchristi. A. Pereopod 1. B. Uropod. single conical spine at mediodistal corner, with lacinia on inner face bearing three (four?) obscure teeth. Maxilla 1 strongly curved and indurate, armed distally with ten large spines and one small spine, some of former distally finely serrate; no inner ramus present. Maxilla 2 distally bilobed, upper lobe slightly narrower than lower, bearing two elongate serrate spines; lower lobe with five serrate spines. Exopod of maxilliped broadly triangular, posterior (outer) face with setae and fine setiferous scales; endite broader than palp, distal edge straight, with single strong coupling 48 ANNALS OF THE SOUTH AFRICAN MUSEUM hook on median margin; posterior face of palp and endite densely setose; palp 3-segmented, two proximal segments relatively broad, distal segment small, rounded. : Pereopod | robust, subchelate ; dactylus meeting proximal spine on propodal palm; unguis about one-third dactylar length, with short spine at base; propodus proximally broad, palm with strong proximal spine plus three smaller spines; carpus, merus, and ischium together equal in length to basis. Pereopods 2 to 7 similar, slender, ambulatory; unguis half length of dactylus; propodus with five elongate plumose setae anterodistally; strong posteroventral sensory spine present; carpus, merus, and ischium subequal in length, basis equal to propodus, carpus, and merus combined. Marsupium formed by four pairs of oostegites on pereonites 1 to 4. (Although the marsupium was empty in the female dissected, the body cavity contained eight large incompletely-formed eggs.) Pleopod 1 indurate, operculate, meeting ventral pleon margins and entirely closing off the branchial chamber; basis half length of rami, with two diverging ridges on anterior surface; exopod shorter and narrower than endopod, both rami with outer (anterior) surface hollowed. Pleopod 2 with broad, almost square, basis bearing six retinaculae; exopod shorter than endopod, lying obliquely across endopod, distally truncate, with twelve distal plumose setae, latter longer than ramus and flexed ventrally into distal narrowed channel of pleon; endopod distally truncate, with eleven or twelve elongate plumose setae, endopods of left and right side linked along median edge by row of small plate-like scales. Pleopod 3 with basis enlarged on inner distal margin into broadly rounded lamella bearing two retinaculae; endopod basally broad, tapering slightly, shorter than exopod; latter with suture in distal half, distally rounded with nine plumose setae. Pleopods 4 and 5 similar, exopod of former with four plumose setae, of latter with two setae. Uropod consisting of single segment, extending beyond pleotelsonic apex, broadening distally into three barely separated lobes, two outer lobes each bearing single small, rounded process; entire appendage bearing fine setules plus irregular longitudinal row of elongate plumose setae. Male Very similar to female, body of adult male slightly narrower than that of ovigerous female. Sternites of pereonites 2 to 7 with single pair of low spines along midline. Pleopod 2 with exopod folded obliquely over endopod distally, with about ten distal plumose setae; endopod slightly longer than exopod, with eleven distal plumose setae; copulatory stylet extending from endopod base to slightly beyond apex, distally curved, apically rounded; basis with four retinaculae in rounded mediodistal lobe. 49 Z <= > Sap _ Sa SS Se Se SS > SS — SSS SS SS SS SSS <= A. Pleon in ventral view, showing operculate first pleopods. A NEW MARINE ISOPOD FAMILY FROM THE SOUTH-WESTERN INDIAN OCEAN Fig. 5. Bathynatalia gilchristi. B. Pleopod 2 2. C. Pleopod 3 2. D. Pleopod 4 . E. Pleopod 5 9. F. Pleopod 2 ¢. 50 ANNALS OF THE SOUTH AFRICAN MUSEUM Previous Records Holotype SAM-A10420, 2 TL 6,5 mm, off Durban, 880 m, April 1901. Material Meiring Naude station SM 129 30°53’S 30°31’E (off Natal) 850 m, May 1977. 2 2° (with oostegites) TL 6,8 mm 6,9 mm 2 63 TL 4,8 mm 5,6 mm 5 juvs TL 2,9 mm-4,2 mm ACKNOWLEDGEMENTS My thanks are due to the crew of the R/V Meiring Naude and the staff of the South African Museum who took part in the 1977 cruise. I am grateful to Mr R. Downes of the South African Museum for sorting the benthic samples, to Carolyn Bartlett Gast of the Smithsonian Institution, for assistance and guidance in producing Figure 1, and to Dr T. E. Bowman of the Smithsonian Institution, for comments and criticism of the manuscript. REFERENCES BARNARD, K. H. 1940. Contributions to the crustacean fauna of South Africa. XII. Further additions to the Tanaidacea, Isopoda, and Amphipoda, together with keys for the identifi- cation of the hitherto recorded marine and fresh-water species. Ann. S. Afr. Mus. 32: 381-543. BARNARD, K. H. 1957. Three additions to the fauna list of South Atrican Crustacea. Ann. Mag. nat. Hist. (12) 10: 814-816. MeEnzIES, R. J. 1962. The zoogeography, ecology, and systematics of the Chilean marine isopods. Acta Univ. lund. (2) 57: 1-162. Moreira, P. S. 1971. Species of Serolis Isopoda, Flabellifera) from southern Brazil. Bolm Inst. oceanogr. S. Paulo 20: 85-144. SCHULTZ, G. A. 1969. How to know the Marine Isopod Crustaceans. Dubuque, lowa: W. C. Brown. 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. Nnov., 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.: Releire 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 A NEW MARINE ISOPOD FAMILY FROM THE SOUTH-WESTERN INDIAN OCEAN VOLUME 75 PART 5 MARCH 1978 . ISSN 0303-2515 * // | ?. ‘ 1 ) | $ 7 ‘OF THE SOUTH AFRIC Ge MUSLIM | 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. 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REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: ‘Smith (1969) describes...’ ‘Smith (1969: 36, fig. 16) describes...” “As described (Smith 1969a, 1969b; Jones 1971)’ ‘As described (Haughton & Broom 1927)...’ ‘As described (Haughton et al. 1927)...’ Note: no comma separating name and year pagination indicated by colon, not p. names of joint authors connected by ampersand et al. in text for more than two joint authors, but names of all authors given in list of references. (b) Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, b, etc. to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a). For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list of 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. én. 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. Konun, A. J. 1960b. Spawning behaviour, eae masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): Sills THIELE, J. 1910. Mollusca: B. Polyelcophere 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 75 Band March 1978 Maart Part 5 Deel q Ay e SB B.B3:9 Oy “X) . N Se > \O Lp, 4 {oun no SS S UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA By MICHAEL R. COOPER Cape Town Kaapstad The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular intervals as material becomes available Obtainable from the South African Museum, P.O. Box 61, Cape Town Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM word uitgegee in dele op ongereelde tye na beskikbaarheid van stof Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad OUT OF PRINT/UIT DRUK 1, 201-3, 5-8), 3(1-2, 4-5, 8, t.—p.i.), 51-3, 5, 7-9), 6(1, t.—p.i.), 711-4), 8, 9(1-2, 7), 101-3), - 11(1-2, 5, 7, t.—p.i.), 15(4—-5), 24(2), 27, 31(1-3), 32(5), 33 Price of this part/Prys van hierdie deel R7,40 ISBN 0 908407 41 6 Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk.., Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA By MICHAEL R. COOPER Department of Geology, University of Oxford (With 39 figures) [MS. accepted 18 May 1977] ABSTRACT The classic ammonite locality at Salinas was visited and the section measured. Contrary to previous estimates, the earliest marine strata at this locality, resting conformably upon a fluvio-continental facies, are of latest Cenomanian age and clearly reflect marine transgression. The ammonite fauna from this locality is described, some of the more important elements being Metoicoceras gibbosum Hyatt, Euomphaloceras (Kanabiceras) septemseriatum (Cragin), Sciponoceras gracile (Shumard), Calycoceras naviculare (Mantell), Pseudocalycoceras angolaense (Spath), Gaudryceras isovokyense Collignon, Anagaudryceras cf. cassisianum (d’Orbigny), Puzosia (Austiniceras) intermedia orientalis Matsumoto, Puzosia (Anapuzosia) dibleyi (Spath), Tetragonites aff. blaisoni Collignon and Desmoceras (Pseudouhlighella) aff. ezoanum Matsumoto. This assemblage forms part of the world-wide Sciponoceras gracile Zone fauna of latest Cenomanian age. The presence of Watinoceras coloradoense (Henderson) and Vascoceras (Paravascoceras) cf. cauvini Chudeau in surface scree is taken as evidence for the basal Turonian zone of Watinoceras coloradoense. The Cenomanian—Turonian boundary problem is discussed at length and a modified zonation of the Lower Turonian proposed. Acanthoceratid phylogeny is commented upon, and the new subfamily Euomphaloceratinae is erected to house the following taxa: E. (Euomphaloceras), E. (Kanabiceras), Kamerunoceras, Schindewolfites, Yubari- ceras, Romaniceras (including Proromaniceras), Obiraceras, Shuparoceras and, tentatively, the Cenomanian Tunesites. CONTENTS PAGE Introduction ; ; : 5 , : ; I Geology . : : : : ; : 4 53 Systematics . ; 3 ; : ‘ ; 56 Age of the fauna ‘ ; ‘ : : so 1133 The Cenomanian—Turonian boundary problem ee SS Palaeogeographic significance f : ‘ 42 Summary . é : 5 s : 5 . 145 Acknowledgements . ; ; ; : . 146 References . ; : : : : . . 146 INTRODUCTION Salinas is situated a short distance to the north-west of the army camp of Posto de Sao Nicolau, 110 km to the north of Mocamedes (Fig. 1). Douvillé (1931) was the first to describe ammonites from Salinas when he recorded a ‘Barremian’ to ‘Turonian’ fauna, identifying the following species: Sl Ann. S. Afr. Mus. 75 (5), 1978: 51-152, 39 figs. a2 ANNALS OF THE SOUTH AFRICAN MUSEUM . Ponta das Salinas Fossil locality Luanda Benguella Mocamedes Grossa Fig. 1. Locality map. Barremian Pulchellia caicedoi Karsten Acanthoceras lyelli Leymerie Albian Desmoceras toucasi Jacob Knemiceras uhligi Choffat Acanthoceras borgesi Douvillé Gaudryceras salinarium Douvillé Prionotropis echinatus Douvillé Turonian Puzosia matheroni d’Orbigny i Vraconnian Cenomanian UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 35 Spath (1931), in a review of this fauna, considered all these ammonites to be of Cenomanian age, referring the Acanthoceras lyelli to Protacanthoceras, Acanthoceras borgesito Metacalycoceras and Prionotropis echinatus to Neocardio- ceras, while considering the Puzosia matheroni closer to the Cenomanian Austiniceras dibleyi than d’Orbigny’s Lower Cretaceous species. Haas (1942: 158, 163) tentatively referred Desmoceras toucasi Douvillé (non Jacob) to Puzosia spathi Venzo (=P. venzoi Breistroffer 1947), thereby suggesting the Cenomanian of Salinas to be underlain by Albian strata. He reasserted this opinion (Haas 1952: 16) when describing a specimen of Desmoceras latidorsatum var. inflata Breistroffer, supposedly from the Salinas exposure, stating that since ‘the specimen under discussion is undoubtedly conspecific with the specimens from Egito and Catumbela, whose Albian age need not be questioned, the Albian fauna may be assumed to be present beneath the Cenomanian one at Salinas’. Howarth (1965) followed Spath in considering the Salinas fauna to be Upper Cenomanian, but felt that Desmoceras latidorsatum vat. inflata admitted to the presence of Lower Cenomanian strata at Salinas. Kennedy (1971) considered the Salinas fauna to be of high Cenomanian or basal Turonian age, pointing out that Metacalycoceras borgesi (Douvillé) was conspecific with Calycoceras naviculare (Mantell), while confirming the puzosiid to be Austiniceras dibleyi. Neocardioceras echinatum (Douvillé) he referred to the genus Kanabiceras, while following Spath (1931) in considering the ‘Pulchellia’ a misidentified acanthocerate. The Stoliczkaia dispar var. attenuata Douvillé (1931: 29, pl. 2 (fig. 2a—b)), if a Stoliczkaia, he considered must ‘surely be Lower Cenomanian’. GEOLOGY The lowest beds in the stratigraphic succession at Salinas (Fig. 2) are exposed in the low sea-cliffs to the south of the Baia das Salinas. These beds comprise about 8 m of predominantly red, but also white, green and purple, laminated silts and clays, entirely lacking in fossils. That these beds attain a much greater thickness is evident in the sea-cliffs to the south of the Farol de Ponta Grossa, where at least 40 m of predominantly red siltstones and sandstones underlie the lowest limestone horizon. At Salinas these beds are overlain by 1,80 m of yellowish fine-grained sand- stone, also unfossiliferous, the upper 0,80 m with numerous well-rounded boulders and pebbles up to 0,20 m in diameter. This conglomeratic horizon is overlain by 1,0 m of highly fossiliferous white limestone, characterized by the abundance of the gasteropod Pseudomelania salenasensis Rennie. The succession can thus be interpreted as a typical transgressive sequence with initially highly coloured, unfossiliferous lagoonal sediments covered by receding dune sands or barrier deposits, then beach conglomerates and finally shallow-water open marine limestones. The ‘Pseudomelania Beds’ are crammed with Rhynchostreon suborbiculatum (Lamarck) (Fig. 3), together with Exogyra (Costagyra) olisiponensis Sharpe, ANNALS OF THE SOUTH AFRICAN MUSEUM Brownish silts with lines of gray calcareous nodules yielding Kanabiceras, Pseudocalycoceras, Calycoceras, Metoicoceras and Sciponoceras Poorly exposed brownish silts with more resistant portions, yielding rare C.naviculare (Mantel1) Unfossiliferous brownish limestones weathering with a highly jagged surface 5m "Pseudomelania Beds" - fossiliferous white limestone with Pseudomelania, Veniella etc. Unfossiliferous, yellowish fine-grained sandstone, the upper portion conglomeratic Predominantly red, but also white, green and purple, laminated silts and clays, lacking in fossils Fig. 2. Geological section through the Salinas exposure. For explanation see text UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 39) Fig. 3. A-C. Rhynchostreon suborbiculatum (Lamarck). D-E. Trigonarca sp. F. Pseudomelamia salenasensis Rennie. All specimens in the South African Museum collections. x 1. 56 ANNALS OF THE SOUTH AFRICAN MUSEUM Protocardia hillana (J. Sowerby) and Veniella forbesiana (Stoliczka), but are entirely lacking in ammonites. Three metres of unfossiliferous brownish lime- stones, weathering with a highly jagged surface, and interbedded silts separate this horizon from the first unit containing Calycoceras naviculare (Mantell). However, it is only 2,50 m higher up, in the lowest horizon of calcareous nodules that ammonites become relatively abundant. At this point the section becomes difficult to follow but there would appear to be, in all, some eight such nodular horizons in a thickness of about 6 m. These nodules are extremely hard and highly fossiliferous, being characterized by the abundance of the bivalve Rhyncho- streon suborbiculatum (Lamarck). SYSTEMATICS All catalogue numbers refer to the collections housed in the South African Museum. Measurement abbreviations are as follows: D, diameter; H, whorl height; W, whorl width (intercostal unless otherwise specified); Ui, diameter of umbilicus between umbilical seams; Uo, diameter of umbilicus between umbilical bullae; W/H, whorl width/height ratio. All measurements are given in milli- metres, and percentages follow in parentheses. Class CEPHALOPODA Cuvier, 1797 Subclass AMMONOIDEA Zittel, 1884 Order LYTOCERATIDA Hyatt, 1899 Suborder LYTOCERATINA Hyatt, 1899 Superfamily TETRAGONITACEAE Hyatt, 1900 Family Tetragonitidae Hyatt, 1900 Genus Tetragonites Kossmat, 1895 Type species Ammonites timotheanus Pictet, 1847 1895 Lytoceras (Tetragonites) Kossmat 1925 Epigoniceras Spath 1935 Saghalinites Shimizu (nom. nud.) 1935 Neoepigoniceras Shimizu (nom. nud.) 1935 Eoepigoniceras Shimizu (nom. nud.) 1954 Epigoniceras (Saghalinites) Wright & Matsumoto Discussion Much attention has recently been paid to the Tetragonitidae (Wiedmann 1962, 1973; Murphy 1967a), and a relatively stable taxonomic classification appears to have been produced. Wiedmann (1973: 588) rejected Murphy’s (1967a) emphasis on constrictions as a basis for classification since ‘number and course of constructions vary with age’, consequently basing his taxa on whorl section and the persistence of con- constrictions. Sutural differences were regarded as of limited taxonomic use. A number of workers (Howarth 1958: 9; Wiedmann 1962: 131, 171) have pointed out that Epigoniceras is indistinguishable from Tetragonites, while Wiedmann (1973: 589) does not regard the development of distinct umbilical UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 5 shoulders as an adequate basis for the generic separation of Saghalinites. The new genus Carinites Wiedmann was proposed for the keeled Cenomanian T. spathi (Fabre 1940: 214, pl. 6 (fig. 1), text-fig. 26), but appears to be of dubious status. Tetragonites sp. nov.? aff. blaisoni Collignon, 1964 Figs 4P-Q, 5 Compare Tetragonites blaisoni Collignon, 1964: 31, pl. 324 (fig. 1448). Wiedmann, 1973: 601, pl. 1 (fig. 4), pl. 6 (figs 5—7) text-fig. 7. Tetragonites collignoni Murphy (?non Breistroffer), 1967a: 66, pl. 5 (figs 2-3), text-fig. 36a only. Tetragonites subtimotheanus Murphy (non Wiedmann), 1967a: 62, pl. 6 (figs 5-8), ? text-fig. 35a—d only. Description The recrystallized test of the specimen has been preserved and is moderately involute, with about 75 per cent of the preceding whorl covered. The umbilicus is rather narrow (+ 28% of diameter) and fairly deep, with broad, inclined umbilical walls and angular umbilical shoulders. The flanks are flat and con- verge noticeably towards the flat venter. The whorl section is depressed, trape- zoidal, with the greatest width at the umbilical shoulder. The ornament comprises weak, periodic, strongly prorsiradiate collars on the flanks, which cross the venter with a very slight concave-adoral arch. Measurements No. D H W W/H Ui SAM-PCA2818 31 13 (42) 16 (52) 1,23 9 (29) a 2D 9 (41) 10 (45) lel 6 (27) Discussion The unique Angolan example is fragmentary, hence comparison with other members of the genus is difficult. The relative proportions of the Salinas specimen are virtually identical to those of Kossmat’s holotype of 7. subtimotheanus, and easily fall within the range of morphometric variation shown by this species (Wiedmann 1973). The whorl section is also closely comparable, except that in 7. subtimotheanus Wied- mann the umbilical wall is almost vertical, whereas in the Angolan example it is very distinctly inclined (Fig. 5). In the fragmentary Angolan specimen it is not possible to decide how many constrictions there were per whorl, but in general form they closely resemble those of T. subtimotheanus. In North America, T. subtimotheanus ranges from the upper Lower Albian (hulenense Zone) to the Lower Cenomanian, although elsewhere it appears to be confined to the Upper Albian. In view of the different stratigraphic ages and the slight but distinct 58 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 4. A-C, H-K. Pseudocalycoceras angolaense (Spath). A-—C. SAM-PCA2777. H-I. SAM-PCA2719. J-K. SAM-PCA2767. D-E. Rhynchostreon suborbiculatum (Lamarck), SAM-PCA2801. L-M. Calycoceras (Calycoceras) naviculare (Mantell), SAM—PCA2762. F—G, N-O. Euomphaloceras (Kanabiceras) septemseriatum (Cragin), SAM-—K 2878, K2779. P-Q. Tetragonites sp. nov. ? aff. blaisoni Collignon, SAM-—PCA2818. x 1. UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 59 Fig. 5. Tetragonites sp. nov. ? aff. blaisoni Collignon. Whorl section of SAM—PCA2818. x1. difference in whorl-section, the Angolan example can only be regarded as a closely allied, but different, species. Tetragonites rectangularis ampakabensis Collignon, from the Lower Ceno- manian of Madagascar, differs in being more involute (U=20-23°% of diameter), somewhat more depressed, and with steeper umbilical walls. Tetragonites blaisoni Collignon has similar relative proportions to the Salinas example, while the con- figuration of the constrictions is also closely comparable. It differs, however, in having much steeper umbilical walls and in being a Lower Cenomanian form. According to the amended diagnosis of Tetragonites glabrum Jimbo given by Matsumoto (19425: 671), this species has a small umbilicus (20-26% of diameter), with slightly depressed whorls. The flanks converge to a rounded venter, while constrictions are few and irregular in development. Tetragonites epigonum (Kossmat) has an almost square whorl section (Usher 1952: 55), with not more than three prorsiradiate constrictions per whorl which appear to be effaced across the venter. Tetragonites jurinianus (Pictet) lacks constrictions at all growth stages, while in T. nautiloides (Pictet) constrictions are present only at diameters of less than 10 mm. The Angolan specimen is the only example of a Tetragonites yet recorded from uppermost Cenomanian strata, and in view of the slight, but distinct, differences from all previously recorded species of this genus, it probably merits a new specific name. The author hesitates to do so, however, due to the poor preservation of the unique Angolan example. Family Gaudryceratidae Spath, 1927 Discussion The family Gaudryceratidae has been subdivided into a number of sub- families—Gaudryceratinae (Spath 1927), Gabbioceratinae (Breistroffer 1953), Kossmatellinae (Breistroffer 1953) and Vertebritinae (Wiedmann 1962). The use- fulness and desirability of these groupings are questionable. Vertebrites is herein regarded as only a subgenus of Gaudryceras and consequently the Vertebritinae must fall into synonymy (cf. Henderson 1970: 22). Moreover, the close affinities of Kossmatella and Anagaudryceras are shown by the fact that the newly proposed species Kossmatella (Murphyella) enigma Matsumoto, Muramoto & Takahashi, type of the subgenus, is herein considered a junior subjective synonym of Anagaudryceras buddha (Forbes) (= A. sacya Forbes), thereby suggesting the superfluity of the subfamily Kossmatellinae. The subfamily Gabbioceratinae Breistroffer was considered by Murphy 60 ANNALS OF THE SOUTH AFRICAN MUSEUM (19675) to include only two genera, Gabbioceras and Jauberticeras, and charac- terized by a wide crater-like umbilicus, with flat, sloping umbilical walls, and a prominent ventro-lateral angulation of the whorls. Maximum width of the whorls was at this angulation. Ornament comprises fine, sinuous lirae and con- strictions as in Gaudryceras. According to Murphy (19676: 604), “The principal difference between Jauberticeras and Gabbioceras is in the position of the angula- tion or, alternatively expressed, the relative curvature of the venter. In Jauberti- ceras, the angulation is very high and the curvature of the venter low; in Gabbio- ceras, the angulation is at about mid flank and the degree of curvature is high.’ The differences are not herein considered of generic importance, and the Gabbio- ceratinae thus becomes a monogeneric subfamily of doubtful systematic value. Genus Gaudryceras de Grossouvre, 1894 Type species Ammonites mitis Hauer, 1866 1894 Gaudryceras de Grossouvre 1926 Vertebrites Marshall 1934 Epigaudryceras Shimizu 1934 Pseudogaudryceras Shimizu 1934 Hemigaudryceras Shimizu 1935 Neogaudryceras Shimizu Discussion Wright (in Arkell et al. 1957: L200) considered the following features to be diagnostic of the genus Gaudryceras: “Lirae coarser than in Anagaudryceras, close or distant, simple or branching, moderately to very sinuous; last whorl may bear variable close or distant strong foldlike ribs. Suture with retracted auxilia- ries. May be divisible into subgenera but nominal groups cited in synonymy are not well differentiated.’ Wright & Matsumoto (1954) provided a detailed discussion of this genus, concluding that the genera Epigaudryceras (type species: Ammonites crassicosta- tum Jimbo), Hemigaudryceras (type species: Ammonites denmanensis Whiteaves), Neogaudryceras (type species: Gaudryceras tenuiliratum Yabe) and Pseudogaudry- ceras (type species: Gaudryceras tenuiliratum var. infrequens Yabe) were junior subjective synonyms of Gaudryceras. Anagaudr yceras Shimizu (type species: Ammonites sacya Forbes, 1846: 113, pl. 14 (fig. 10)) was regarded as a synonym of Gaudryceras by Wiedmann (1962: 156-158). As noted by Howarth (1965: 357), however, ‘The ornament of all but the adult stage of Anagaudryceras is so fine that the shell appears to be smooth, while Gaudryceras is characteristically covered with fine ribs. This difference is sufficient for generic distinction in keeping with the scale of differences usually adopted for Cretaceous genera.’ Mesogaudryceras Spath (type species: Ammonites leptonema Sharpe, 1855: 32, pl. 14 (fig. 3)) differs from Gaudryceras in possessing compressed inner whorls, broad convex flanks, and a narrowly arched venter. It is also more involute than Gaudryceras. UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 61 Vertebrites Marshall (type species : Vertebrites murdochi Marshall, 1926: 139, pl. 20 (fig. 9), pl. 30 (figs 1-2), pl. 40 (fig. 2)) is an Upper Senonian—Maastrichtian genus, more evolute than Gaudryceras and retaining a flattened venter to larger growth stages, so that the ventro-lateral shoulders tend to become subangular. Fine flank lirae increase by bifurcation and intercalation across the venter. These ribbing characteristics are developed in a number of typical Gaudryceras, e.g. Gaudryceras vertebratum Kossmat, Gaudryceras stefaninii Venzo (W. J. Kennedy pers. comm.), and thus the only distinct difference between Vertebrites and Gaudryceras is that the former is more evolute and tends to become serpenticone, while the tabulate venter is retained to large diameters. The writer does not regard these differences as generic and prefers to treat Vertebrites as a subgenus of Gaudryceras (cf. Matsumoto 1959b: 141). Eogaudryceras Spath (type species: Ammonites numidus Coquand in Sayn 1890: 144, pl. 1 (figs 3-4)) is a Lower Cretaceous genus, moderately evolute and with an initially trapezoidal whorl section becoming rounded with age. Ornament comprises fine, dense, slightly sinuous lirae and constrictions on the inner whorls only. It thus superficially resembles Anagaudryceras. Gaudryceras (Gaudryceras) isovokyense Collignon, 1964 Figs 6A—B, 7 Gaudryceras isovokyense Collignon, 1964: 31, pl. 324 (fig. 1447). Description One well-preserved specimen, partially retaining recrystallized test, is assigned to this species. The shell is entirely septate to a diameter of 74 mm. It is very evolute, with about 30 per cent of the preceding whorl covered. The whorl section at the adoral end of the outer whorl is oval, slightly depressed, with convex flanks converging slightly towards the evenly rounded venter. The preceding whorl, however, shows a more depressed whorl section with a broad, almost flat, venter (Fig. 7), as in the holotype. The umbilicus is wide and shallow, with a moderately steep umbili- cal wall and a well-rounded umbilical shoulder. The ornament comprises fine, sharp lirae which begin at the umbilical seam, are slightly prorsiradiate on the umbilical wall, curve forward more strongly on the umbilical shoulder, sweep across the flanks and recurve so as to pass straight across the venter. At two places on the outer whorl, about 340° apart, 2-4 adjacent ribs are crowded together, thicker and slightly flared. This feature is associated only with the external shell as no constrictions are visible on the internal mould. Across the venter, the flank ribs become finer and increase by intercalation and bifurcation. Thus 19 flank ribs correspond to 28 ribs across the venter. On the anterior portion of the outer whorl there are 12 ribs within a 10 mm distance along the venter, whereas on the posterior position there are 16 in the same distance. 62 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 6. A-B. Gaudryceras (Gaudryceras) isovokyense Collignon, SAM-—PCA2713. C-H. Vasco- ceras (Paravascoceras) cf. cauvini Chudeau. C-D. SAM-—PCA2816. E-F. SAM-—PCA2727. G-H. SAM-PCA2796. I-J. Pseudocalycoceras sp., SAM-—K2760. x1. UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 63 Fig. 7. Gaudryceras (Gaudryceras) isovokyense Collignon. Whorl section of SAM-—PCA2713. x1. Measurements No. D H W W/H Ui SAM-PCA2713 74 25 (34) 28 (38) LW 29 (26) < 54 17 (31) Die(G9)) L743) WS) (2D) is 37 11 (30) 17 (46) eS 21 (18) Discussion So far as can be judged, the penultimate whorl of the Salinas example is identical with the holotype of G. isovokyense from the Cenomanian ‘Zone a Mantelliceras mantelli et Calycoceras newboldi’ of Madagascar (Collignon 1964). Gaudryceras varagurense Kossmat (1895: 122, pl. 17 (fig. 4), pl. 18 (figs 2a—c)), known from the Cenomanian to uppermost Campanian, differs from G. isovokyense in retaining an evenly rounded venter throughout ontogeny, having more frequent flared ribs and in that the rib density remains the same across the venter. Gaudryceras stefaninii (Venzo) (1936: 21, pl. 2 (figs 3-4)) from the Cenoma- nian of Zululand bears a close resemblance to the Angolan material. This species differs, however, in having much finer, denser, Vertebrites-like ribbing across the venter. Gaudryceras vertebratum Kossmat (1895: 126, pl. 15 (figs 4-5)) shows a flattened venter which is apparently ‘smooth’, with prorsiradiate lirae on the flanks. Gaudryceras (Vertebrites) murdochi (Marshall) (Henderson 1970: 22, pl. 3 (fig. 1)) is an Upper Campanian species, very evolute, serpenticone with strongly depressed whorls and a broad flat venter. Gaudryceras varicostatum (van Hoepen) (1921: 7, pl. 2 (figs 10-12)) shows dense fine lirae across the venter, but the latter is not nearly so flattened as in 64 ANNALS OF THE SOUTH AFRICAN MUSEUM G. isovokyense. Constrictions also appear to be more frequent in Van Hoepen’s species. Gaudryceras (?Vertebrites) kayei (Forbes) (1846: 101, pl. 8 (fig. 3)) is a very evolute, serpenticone form, with a broad, convex venter and about four constric- tions on the outer whorl. The specimen figured by Stoliczka (1865: 156, pl. 77 (figs 1-2)) shows the same features as Forbes’s example, but with a somewhat more flattened venter. The ornament comprises fine prorsiradiate flank ribs which, according to Stoliczka (1865: 156), “bi-, or tri-furcate at the edge of the umbilicus (= ventro-lateral shoulder)’. This species differs from both G. stefa- ninit (Venzo) and G. vertebratum Kossmat in having a convex and not flattened venter. Occurrence This species is currently known only from Angola and Madagascar. Genus Anagaudryceras Shimizu, 1934 Type species Ammonites sacya Forbes, 1846 1934 Anagaudryceras Shimizu 1934 Paragaudryceras Shimizu 1972 Kossmatella (Murphyella) Matsumoto, Muramoto & Takahashi Discussion Anagaudryceras is a long-ranging genus known from the Albian to the Maastrichtian. A host of species have been described but, as noted by Howarth (1965), the majority are nominal species, of little taxonomic value. Differences from G. (Gaudryceras) and G. (Vertebrites) are noted above. The newly proposed subgenus Kossmatella (Murphyella) was diagnosed (Matsumoto et al. 1972: 208) as follows: ‘The Kossmatella type ribs occur on relatively early inner whorls (with diameters from several millimetres to 10 or 15 mm), but on the late inner whorls the ribs and furrows are sparse and rather weak. In other words smoothish part is predominant over the ribbed or con- stricted part in the middle growth stage. In the late septate whorl and the adult body-whorl the fold ribs and intervening furrows are distinct.’ Thus, Murphyella supposedly differs from the contemporaneous Anagaudry- ceras only in possessing fold-like ribs and constrictions at a very early stage in growth, although, as noted by Matsumoto ef al. (1972), these are variable in persistence and intensity. However, McLearn (1972) has recently shown identical features to occur in Anagaudryceras sacya (= A. buddha), and the author has little hesitation in regarding Kossmatella (Murphyella) as a junior subjective synonym of Anagaudryceras. UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 65 Anagaudryceras cf. cassisianum (d’Orbigny, 1850) Figs 8-9, 10J, 11B—D Compare Ammonites cassisianus @Orbigny, 1850: 146. Ammonites madraspatanum Stoliczka, 1865: 151, pl. 75 (fig. 2). Anagaudryceras madraspatanum (Stoliczka) Matsumoto, 1959a: 74, pl. 22 (fig. 3). Gaudryceras salinarium Douvillé, 1931: 42, pl. 1 (fig. 3), text-fig. 5. Description Two examples, one a juvenile and the other a fragment of an adult, both with recrystallized test preserved, are close to d’Orbigny’s species. Fig. 8. Anagaudryceras cassisianum (d’Orbigny). D’Orbigny’s syntypes from the Cenomanian of Cassis, France. Photo: W. J. Kennedy. x1. The juvenile example, SAM—PCA2769 (Fig. 11 B—D), is very evolute (umbili- cus 46-50% of diameter), with about one-sixth of the preceding whorl covered. The umbilicus is wide and shallow, with a steep umbilical wall and a well-rounded umbilical shoulder. The flanks are strongly convex and converge towards the evenly rounded venter. The ornament comprises extremely fine, indistinct, flexuous lirae which arise at the umbilical seam, pass forward across the umbilical wall, swing forward more strongly on the flanks, then recurve gently so as to cross the venter transversely. There appear to be occasional, very weak constrictions on the earliest whorls. SAM-—PCA2726 (Fig. 10J) is a fragment of a mature individual. The whorl section is unknown. Ornament comprises fine lirae which arise at the umbilical seam and pass forward (prorsiradiate) to the umbilical shoulder where they increase by bifurcation and intercalation. The lirae are almost rectiradiate across the flank. A prominent collar, following much the same course as the lirae, splits into three fine ribs across the flanks. Although a much larger diameter, this specimen very closely resembles d’Orbigny’s larger syntype of Ammonites cassisianus. 66 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 9. Anagaudryceras cf. cassisianum (d’Orbigny). The holotype of Gaudryceras salinarium Douvillé (after Douvillé 1931). x1. Measurements No. D H W W/H Ui SAM-PCA2769 39 15 (38) 18 (46) 1,20 u s 28 8 (29) 12 (43) 1,50 13 (46,4) < oD) 6 (27) 10 (45) 1,70 11 (50) Discussion D’Orbigny’s syntypes (Fig. 8) show the following features: the shell is evolute (umbilicus 41-45 % of diameter), with a wide, shallow umbilicus, evenly rounded umbilical shoulder, slightly convex flanks, and a rounded venter. The whorl section is almost circular, about as wide as high. On the inner whorls, extremely fine lirae arise at the umbilical seam and pass forward (prorsiradiate) to the umbilical shoulder where they increase by bifurcation and intercalation. On the flanks the lirae are initially prorsiradiate, but quickly recurve so as to become rectiradiate or slightly rursiradiate on the ventral half of the flanks. There are periodic collars which follow much the same course as the constric- tions. On the outer whorls, the lirae become coarser and Gaudryceras-like, nar- rower than the interspaces, while collars split into coarse lirae on the flanks. Gaudryceras salinarium Douvillé is based upon a fragmentary specimen with a circular whorl section, and showing broad Anagaudryceras-like flank ribs on which are superimposed Gaudryceras-like lirae (Fig. 9). Thus, the holotype of G. salinarium appears to differ from A. cassisianum only in what appear to be the broad flank ribs typical of mature Anagaudryceras. Of the new material from the Salinas locality, a large fragment (Fig. 10J) appears indistinguishable from d’Orbigny’s species, but lacks the broad flank ribs of G. salinarium, while the juvenile (Fig. 11B—D) differs in possessing a depressed whorl section and very indistinct collars. Neither d’Orbigny’s nor Douvillé’s species are well known and the differences are well within the range of intraspecific variation observed in UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 67 Fig. 10. A-E. Euomphaloceras (Kanabiceras) septemseriatum (Cragin). A. SAM—PCA2738. B-C. SAM-PCA2797. D-E. SAM-PCA2832. F-—G. Pseudocalycoceras angolaense (Spath). F, SAM-PCA2743. G. SAM-—PCA2772. H-I. Pseudocalycoceras aff. haugi (Pervinquiére), SAM-PCA2775. J. Anagaudryceras cf. cassisianum (d’Orbigny), SAM—PCA2726. x 1. 68 ANNALS OF THE SOUTH AFRICAN MUSEUM many Cretaceous ammonite species. However, without additional material the problem cannot be resolved in this paper, and the Angolan material is referred to Anagaudryceras cf. cassisianum (d’Orbigny). Other broadly contemporaneous species of Anagaudryceras include A. buddha (Forbes), A. sacya (Forbes), A. involvulum (Stoliczka), A. revelatum (Stoliczka), A. madraspatanum (Stoliczka), A. utaturense Shimizu, A. multiplexus (Stoliczka), A. limatum (Yabe), A. aurarium (Anderson) and A. enigmum (Matsu- moto, Muramoto & Takahashi). Since 1865, many workers (Stoliczka 1865, Kossmat 1895, Matsumoto 1959a) have realized that Ammonites sacya Forbes (1846: 113, pl. 14 (fig. 10)), the type of the genus Anagaudryceras, was merely based upon the inner whorls of Ammonites buddha Forbes (1846: 112, pl. 14 (fig. 9)), a body chamber frag- ment. Since A. buddha clearly has priority, it includes among its synonyms (cf. Howarth 1965) A. sacya (Forbes), A. revelatum (Stoliczka), A. limatum (Yabe) and A. enigmum (Matsumoto, Muramoto & Takahashi). Anagaudryceras buddha (Forbes) differs from A. cassisianum in maintaining extremely fine liration to maturity. Anagaudryceras multiplexum (Stoliczka 1865: 151, pl. 75 (fig. 2)) differs from A. cassisianum in being somewhat more evolute (umbilicus 50°, of diameter), with a steeper umbilical wall and more abrupt umbilical shoulder, as well as in retaining extremely fine liration to maturity. Anagaudryceras madraspatanum (Blanford) (Stoliczka 1865: 151, pl. 75 (fig. 2)) was considered to show the following features: ‘Shell consisting of numerous round whorls, which are only slightly involute, increasing regularly in height and thickness, and crossed by dense fine flexuous striae. On the casts of some of our specimens, slight transverse furrows are seen. . . . Umbilicus large and deep, three-fifths of the inner whorls being exposed; aperture circular.’ Matsumoto (1959a: 74, pl. 22 (fig. 3)) recorded A. madraspatanum from the Chitina Valley of Alaska and considered the diagnostic features to be ‘the circular cross section of the whorl, little involution, fairly wide umbilicus, fine and dense striae and lirae on the surface of the shell that curve gently forward near the umbilical margin, but are nearly rectiradiate on the main part of the whorl, and the faint constrictions’. Of these characters, the prominence of constricted collars and the shape of the whorl section are known to vary with ontogeny, while the width of the umbilicus and the degree of involution of the shell are comparable with A. cassisianum. Adult specimens of A. madraspatanum are unknown and consequently the writer regards Stoliczka’s (1865) species as a junior subjective synonym of A. cassisianum. Anagaudryceras involyulum (Stoliczka) (1865: 150, pl. 75 (fig. 1)) is a com- pressed species with rapidly expanding whorls and about five constrictions on the outer whorl at 44 m diameter. This species has recently (Howarth 1968) been recorded from the Turonian of Ponta Grossa, Angola. It would seem to differ from A. cassisianum in being more compressed, and in not developing Gaudryceras-type lirae in maturity. UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 69 Fig. 11. A. Puzosia (Anapuzosia) dibleyi (Spath), SAM-—PCA2792. B-D. Anagaudryceras cf. cassisianum (d’Orbigny), SAM—PCA2769. x1. 70 ANNALS OF THE SOUTH AFRICAN. MUSEUM - Occurrence Anagaudryceras cassisianum is at present recorded from France, Alaska, southern India and possibly Angola. Suborder ANCYLOCERATINA Wiedmann, 1966 Superfamily ANCYLOCERATACEAE Meek, 1876 Family Baculitidae Meek, 1876 Subfamily Baculitinae Meek, 1876 Genus Sciponoceras Hyatt, 1894 Type species Hamites baculoide Mantell, 1822 Sciponoceras gracile (Shumard, 1861) Fig. 12B—D Baculites gracilis Shumard, 1861: 596. Stanton, 1893: 1166, pl. 34 (figs 1-3). Grabau & Shimer, 1910: 179, fig. 1432. Adkins, 1928: 206, pl. 24 (fig. 3). Moreman, 1942: 210. Baculites ovatus White (non Say), 1876: 199, pl. 19 (figs 4-5). Sciponoceras gracile (Shumard) Cobban, 1951: 2185. Matsumoto, 1959b: 106. Matsumoto & Obata, 1963: 23. Cobban, 1955: 202, pl. 2 (fig. 3). Cobban & Scott, 1972: 47, pl. 17 (figs 9-29). Wright & Kennedy, 1973 (in Juignet et a/. 1973): 21, pl. 1 (figs 2-6). Baculites n. sp. aff. bohemicum Fritsch, Kossmat, 1895: 154, pl. 19 (fig. 18). Baculites kossmati Nowak, 1908: 348. Baculites cfr. gaudini Pictet & Campiche, Matsumoto, 1942a: 229. Sciponoceras kossmati (Nowak) Matsumoto, 19595: 106, pl. 31 (figs 2-3). Matsumoto & Obata, 1963: 13, pl. 3 (fig. 2), pl. 4 (fig. 1), pl. 5 (figs 1-3), pl. 6 (figs 3-5). Cyrtochilus stylus Anderson, 1958: 188, pl. 11 (fig. 5). Description A single fragment, with the recrystallized test preserved, is the only specimen assignable to this genus from Salinas. The shell tapers slowly adapically and has a very compressed, elliptical whorl section, with flattened sides. The ornament, which is very faint, comprises weak, prorsiradiate constrictions, most prominent across the venter but becoming effaced across the lower flanks and dorsum, and very faint ribs which follow the course of the constrictions. There would appear to be one, possibly two, such constrictions within a distance equal to the major diameter. At a major diameter of 9 mm the width is only 6 mm. Discussion Shumard (1861: 596) described Sciponoceras gracile as follows: ‘Shell very slender, gradually tapering to apex; transverse section varying from broad ovate to subcircular; surface in some specimens nearly smooth, but usually marked with moderately prominent, rounded costae, which on the dorsum (venter) are distinct and arched towards the aperture, and on the sides curve obliquely back- wards and downwards to the ventral margin (dorsum), before reaching which they become nearly obsolete. Some of the specimens exhibit very fine flexuous Striae of growth.’ ee UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 71 Shumard’s type material has never been figured or adequately redescribed, nor has a lectotype been designated. According to Wright & Kennedy (in Juignet et al. 1973: 21) the type material is probably in the University of Texas collections. Wright & Kennedy (in Juignet et al. 1973: 21) have recently discussed this species on the basis of topotype material from the Sciponoceras gracile/Metoico- ceras whitei Zone (Cobban 1951) of the Gulf Coast and Western Interior of North America, and material from the late Cenomanian of southern England and northern France. These authors show this species to be characterized by a compressed, elliptical whorl section with flattened sides and broad prorsiradiate constrictions spaced at intervals slightly greater than the major diameter. Between constrictions there are 5—6 ribs. Both ribbing and constrictions are most prominent across the venter, becoming obsolete before the dorsum is reached. Shumard’s (1861) descriptions leave little doubt that there is a fair degree of variability in the prominence of the ornament of this species, and consequently the weak ornament of the Angolan example is not considered significant. Sciponoceras roto Cieslinski differs in having an almost circular whorl section at all growth stages, while S. baculoide (Mantell) has more distant con- strictions (one every 1,5—2 major diameters), a less compressed whorl section, and coarser, more robust ventral ribbing (on the body chamber at least). Sciponoceras glaessneri Wright (1963: 599, pl. 81 (figs 2—3)) from the Middle Cenomanian of Bathurst Island, Australia, is more inflated at large growth stages, with more convex flanks, and lacks distinct ventral ribbing. Sciponoceras kossmati (Nowak) (= Baculites sp. nov. aff. bohemicum Fritsch, Kossmat, 1895: 154, pl. 19 (fig. 18)) is the most important synonym of S. gracile. Sciponoceras kossmati was said (Matsumoto 19595: 107) to have ‘narrower and more deeply incised saddles . . . while the external saddle is much broader than the lateral saddle in S. gracile, the former is nearly as broad as and somewhat lower than the latter in S. kossmati. The general outline of the sutural elements is reversed trapezoidal in S. kossmati, but is rectangular in S. gracile.’ However, not only are they morphologically indistinguishable (Matsumoto & Obata 1963), but they also occur at the same stratigraphical horizon. Moreover, Matsumoto & Obata (1963) have noted considerable variation in the suture of S. kossmati. In Japan, S. kossmati is associated with Calycoceras naviculare (Mantell), Euomphaloceras (Kanabiceras) septemseriatum (Cragin), Tarranto- ceras faustum (Matsumoto & Muramoto) and Pseudocalycoceras angolaense (Spath). These are typical uppermost Cenomanian forms and the author regards S. kossmati as a junior subjective synonym of S. gracile. Occurrence This species is recorded from the top Cenomanian of California, the Western Interior, Texas, England, France, and Angola, and the uppermost Cenomanian-—?basal Turonian of India and Japan. UP ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 12. A. Calycoceras (Calycoceras) naviculare (Mantell), SAM—PCA2764. B—D. Sciponoceras gracile (Shumard), SAM-PCA2724. E-H. Euomphaloceras (Kanabiceras) septemseriatum (Cragin). E-F. SAM-—PCA2761. G-H. SAM-PCA2835. x 1. UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA W3 Superfamily HOPLITACEAE Douvillé, 1890 Family Desmoceratidae Zittel, 1895 Subfamily Puzosiinae Spath, 1922 Genus Puzosia Bayle, 1878 Type species Ammonites planulatus J. de C. Sowerby, 1827 1878 Puzosia Bayle 1900 Pleuropachydiscus Hyatt 1922 Austiniceras Spath 1954a Anapuzosia Matsumoto 1954a Mesopuzosia Matsumoto Discussion Since the inception of Bayle’s genus Puzosia in 1878, a host of names have been proposed for closely allied forms. The genus Mesopuzosia Matsumoto (type species Mesopuzosia pacifica Matsumoto, 1954a: 82, pl. 14 (fig. 1), pl. 15 (figs 1-2), pl. 16 (figs 1-3)) was said (Matsumoto 1954a: 79) to be ‘similar to Puzosia in shell form and suture-line but with much better developed costae. In the middle and adult stages of growth many of the costae (including both longer and some shorter ones) extend below the middle of the flanks and are of equal strength on the outer half of the whorls. Although there are longer and shorter ribs, the differentiation to broader (or stronger) and narrower ones is not found as in Parapuzosia (s.s.) or in Austiniceras and the costation is relatively regular. The shell often attains a huge size and in the last whorl the ornamentation disappears.’ Wiedmann & Dieni (1968) considered Mesopuzosia to be a ‘stratigraphische Gattung’ and placed it in synonymy with Puzosia s.s. However, the differences between the types of Mesopuzosia and Puzosia are at least as great as between the latter genus and Austiniceras. Consequently, the writer prefers to retain Meso- puzosia as a subgenus of Puzosia. Anapuzosia Matsumoto (type species Puzosia buenaventura Anderson, 1938: 185, pl. 40 (fig. 3), pl. 41 (figs 1-2)) was erected as a subgenus of Puzosia, and separated from Puzosia s.s. on body chamber characteristics. In Puzosia s.s., ribbing remains consistently fine to the aperture, whereas in Anapuzosia the ‘adult body chamber is ornamented with periodic, flared or tuberculated, strong ribs of little or no flexuosity’ (Matsumoto 1954a: 71). According tothe emended diagnosis of Renz (1972: 704), in Anapuzosia ribs arise at the umbilical shoulder, are flexuous and either bifurcate or are joined by intercalatories across the venter. In some cases, there may be up to three intercalatories between main ribs. These differences are in keeping with the other generic and subgeneric separations within the Puzosiinae, and consequently the writer follows Renz (1972) in regarding Anapuzosia as a valid subgenus of Puzosia. The writer is not inclined to attach much weight to the change in ornament on the body chamber since, where sexual dimorphism in ammonites has been proved, the macroconch (female) dimorph commonly shows a marked change of ornament on the body chamber. 74 ANNALS OF THE SOUTH AFRICAN MUSEUM The genus Austiniceras Spath (type species Ammonites austeni Sharpe, 1855: 28, pl. 12 (fig. la—b only)) was diagnosed by Kennedy (1971: 37) as follows: ‘Medium-sized to large puzosiids with a compressed, convergent whorl section and a narrowly arched venter. Moderately evolute, with 2 distinct orders or ribbing; distant, long flexuous main ribs with many shorter, flexuous intercalated ribs.’ The lectotype of Austiniceras austeni is a gigantic specimen 410 mm in diameter and juveniles of this species are not yet known. In southern England (Kennedy 1971), A. austeni ranges from low in the mantelli Zone to at least the Middle Turonian, and is one of the commonest Upper Cenomanian—Lower Turonian ammonites in this area. It is clearly obvious from Kennedy’s (1971) excellent figure of the holotype that the inner whorls of this species cannot satis- factorily be distinguished from Puzosia. Moreover, there are a number of described Puzosia species from the Upper Cenomanian and Turonian which probably represent nothing more than the inner whorls of Sharpe’s species. In Japan and California they are known as Puzosia orientale Matsumoto, in Madagascar as P. orientale and P. praecompressa Collignon, in Portugal as P. cf. gaudama (Forbes), and in India as P. intermedia Kossmat. It is clear, therefore, that Austiniceras is distinguished from some Puzosia species only by its large size and, where present, the change in ornament on the body chamber. Within the genus Puzosia s.s.. Wiedmann & Dieni (1968) recognized two species groups. The species group of Puzosia quenstedti (Parona & Bonarelli) was characterized by a moderately wide umbilicus (umbilical ratio 25—33 %), flattened sides which converge towards the narrowly rounded venter and 6-7 constrictions per whorl. The constrictions are strong, radial, and with a strong convex ventral sinus. Species are separated on whorl section differences. The species group of Puzosia mayoriana (d’Orbigny) was characterized by a relatively wide umbilicus (31-40% of the diameter), flattish, subparallel flanks and a broadly rounded venter. There are 4-6 rather deep, sigmoidal constrictions per whorl which form a chevron across the venter. These are the characters of P. planulata Sowerby (= P. subplanulata Schliiter), the type of the genus. The genus Puzosia may now be subdivided as follows: P. (Puzosia). Shell compressed, moderately evolute with 4—7 sinuous collars (constrictions on the internal mould) per whorl. Flanks flattish, subparallel, with a broadly rounded venter. Ribbing simple, distinct only on outer half of whorls. Age: Lower Albian—Upper Turonian. P. (Anapuzosia). Similar to P. (Puzosia), but with ribs arising from near the umbilical shoulder and increasing by intercalation and bifurcation above mid-flank. All ribs and constrictions flexuous. Age: Lower Albian—Upper Cenomanian. P. (Austiniceras). Similar to P. (Puzosia) but large, with flat convergent flanks and a narrowly arched venter. Generally straight, rigid constrictions, strongly projected on the venter. May attain large size, when body chamber UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 75 ornamented with close, dense, robust long ribs extending to the umbilical margin. Age: ? Albian—? Campanian, mainly Cenomanian—Turonian. P. (Mesopuzosia). Similar to P. (Puzosia), but with straight, rigid constric- tions, while ribbing may arise close to the umbilical shoulder. Age: Turonian—Coniacian. Subgenus Austiniceras Spath, 1922 Type species Ammonites austeni Sharpe, 1855 Puzosia (Austiniceras) intermedia orientalis Matsumoto, 1959 Ammonites planulatus Stoliczka (non J. de C. Sowerby), 1865: 134, pl. 67 (figs la—b only). Puzosia gaudama Kossmat (non Forbes), 1898: 118, pl. 16 (figs 2-3), pl. 22 (fig. 2). Puzosia cf. gaudama (Forbes) Choffat, 1898: 82, pl. 17 (fig. 2), pl. 18 (fig. 1). Puzosia orientale Matsumoto, 1954a: 74, pl. 13 (figs 1-2). Collignon, 1961: 32, pl. 5 (fig. 1). Puzosia orientale kossmati Matsumoto, 1954a: 75. Puzosia praecompressa Collignon, 1964: 55, pl. 332 (fig. 1493). Puzosia intermedia orientalis Matsumoto, 1959b: 16, pl. 4 (fig. 1). Diagnosis A moderately evolute (umbilicus 27-34 % of diameter), compressed (W/H = 0,70—0,78) species of Puzosia with about 4-6 straight, prorsiradiate constrictions which project strongly forward on the venter. Numerous fine, strongly prorsi- radiate ribs intercalated on outer one-third of flanks between constrictions and projected strongly across the venter. Description The single specimen from these beds has the recrystallized test preserved. It is strongly compressed and moderately involute, with slightly more than half the preceding whorl covered. The umbilicus is fairly narrow and rather shallow, with a steep, slightly undercut umbilical wall. The broad flanks are flat, converging slightly to the narrowly arched venter. The ornament comprises probably 4-5 straight prorsiradiate constrictions per whorl which project strongly forward across the venter. Between the constrictions are about 22 fine ribs which are present only on the outer one-third of the flanks. Fig. 13. Puzosia (Austiniceras) intermedia orientalis Matsumoto, SAM—PCA2824. x1. 76 ANNALS OF THE SOUTH AFRICAN MUSEUM Measurements No. D H W W/H Ui P. orientale (holotype) . 126,3 50,0 (40) 38,7 GI) 0578 43,3 (34) Kossmat, 1898: pl. 22 (fie5 2) : . 80,0 32,0(40) 23,0(29) 0,72 26,0 (32) Choffat 1898: pl. 17 (fig. 2), Plea Sasa 1) . 125,0 57,0(46) 40,0(32) 0,70 34,0 (27) P. praecompressa (holotype) . -~ *67,0> > 27:20:40) =) 2 01Ci OMS 17,0 (25) SAM-PCA2824 _. . 40,0 17,0 (42) 12;0'G0) 0570 12,0 (30) a - = 30.0 13,0 (43) 10:0:63)) Oni 8,0 (27) Discussion The Angolan example clearly belongs to Wiedmann & Dieni’s (1968) group of Puzosia quenstedti. It differs from the Upper Albian—Lower Cenomanian Puzosia communis Spath (1923: 47, pl. 2 (fig. 3a—e)) in possessing fewer constric- tions per whorl. Puzosia orientale orientale Matsumoto (1954a: 74, pl. 13 (figs 1-2)) was proposed for ‘a species of Puzosia characterized by relatively compressed whorls, narrowly arched venter, numerous, regular, fine and narrow costae on the external half and the prorsiradiate course on sides and conspicuous forward bend on the periphery of the costae and the constrictions’. Matsumoto’s holotype, from the Lower Turonian of Japan, is poorly preserved, but judging from his description, his rather poor figures and the measurements, the Angolan example appears to be conspecific. Puzosia orientale kossmati Matsumoto (= P. gaudama Kossmat (non Forbes), 1898: 118, pl. 16 (figs 2-3)) differs only in being slightly more evolute. Puzosia muelleri Grossouvre (= Ammonites hernensis Schliter, 1871: 40, pl. 9 (figs 12a—b, 14) (non Schliiter 1867)) is very similar to the Angolan specimen, differing only in being more compressed (W/H = 0,65) (? crushed). It is thus possibly conspecific with P. compressa Kossmat. Puzosia subcorbarica Matsumoto (1954a: 73, pl. 9 (fig. 1), pl. 12 (fig. 1)) is a compressed form from the Upper Albian of Japan which is more compressed (W/H = 0,51—0,69) than P. orientale, with (?) slightly sigmoidal constrictions. The Lower Cenomanian Puzosia takei Popovici-Hatzeg (1899: 12, pl. 2 (fig. 2)) is very similar to P. orientale, but is somewhat more evolute, with more constrictions. Puzosia praecompressa Collignon (1964) is a junior subjective synonym of P. intermedia orientalis, as is clearly shown by Collignon’s figure, description and measurements. It is very doubtful whether P. intermedia orientalis can be distinguished from the inner whorls of the contemporaneous Puzosia (Austiniceras) austeni (Sharpe), UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA TF Fig. 14. A. Pseudocalycoceras angolaense (Spath), SAM-—PCA2732. B-C. Puzosia (Anapuzosia) dibleyi (Spath), SAM—PCA2749. D-E. ?Protacanthoceras sp., SAM—PCA2804. F. Exogyra (Costagyra) olisiponensis Sharpe, SAM—PCA2752. x1. 78 ANNALS OF THE SOUTH AFRICAN MUSEUM a species based solely on gigantic mature individuals. However, since the inner whorls of Sharpe’s species are still unknown, the sinking of Matsumoto’s species must await knowledge of the ontogenetic variation in P. (A.) austeni. Occurrence Puzosia intermedia orientale ranges from the Upper Cenomanian into the Lower Coniacian, and is known from Japan, India, Portugal, California, Madagascar and Angola. Subgenus Anapuzosia Matsumoto, 1954 Type species Puzosia buenaventura Anderson, 1938 Puzosia (Anapuzosia) dibleyi (Spath, 1922) Figs 11A, 14B-C, 15B—C Ammonites austeni Sharpe, 1855: 28, pl. 12 (fig. 2 only). Austiniceras dibleyi Spath, 1922: 127. Wright & Wright, 1951: 19. Kennedy, 1971: 39, pl. 13 (figs 1-2), pl. 14 (fig. 4). Puzosia (Austiniceras?) dibleyi (Spath) Spath, 1931: 316. Puzosia matheroni Douvillé (non d’Orbigny), 1931: pl. 2 (figs Sa—b). Diagnosis A medium-sized rather coarsely ribbed Anapuzosia, with 8—9 flexuous main ribs per whorl, between which are 5—9 coarse flexuous intercalatories, some of which arise close to the umbilical shoulder. Description This species is a common member of the Salinas fauna and, like most of the fossils from these beds, almost invariably has the recrystallized test preserved. The shell is compressed to slightly depressed (W/H = 0,83-1,06), and moderately involute (umbilicus 27-32% of diameter), with about 65 per cent of the preceding whorl covered. The umbilicus is fairly narrow and moderately deep, with steep umbilical walls and an evenly rounded shoulder. The flanks are convex and converge towards the evenly arched venter. The whorl section is about as wide as high. There are 8-9 prominent sinuous main ribs per whorl, between which are 5—9 flexuous intercalatories, a number of which arise near the umbilical shoulder whilst the remainder are only intercalated on the outer one-third of the flanks. Measurements No. D H W W/H Ui SAM-—PCA2792 165 71 (43) 74 (45) 1,04 49 (30) a 103 44 (43) 45 (44) 1,02 33162) SAM-PCA 2749 90 37 (41) 39 (43) 1,06 2 CIE) 99 70 30 (43) 29 (41) 0,97 20 (28,5) SAM-—PCA2748 97 40 (41) 41 (42) 1,03 30 (30,9) » iS 32 (43) 31 (41) 0,97 22 (30) : 27 12 (44) 10 (37) —0,83 ? UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 79 Discussion Puzosia (Anapuzosia) dibleyi has long been referred to Austiniceras. At present, this species is known only from north-western Europe and Angola. The British material is based upon crushed composite internal moulds, whereas the well-preserved Angolan material has the recrystallized test preserved and is thus more valuable in determining the generic status of this species. As already pointed out, the differences between Austiniceras and Puzosia s.s. are slight, and the author prefers to treat the former as a subgenus of the latter. Austiniceras austeni differs from Puzosia subplanulata (Schliiter) in its flat con- verging flanks, narrowly arched venter, generally rigid constrictions and its large size. Puzosia dibleyi (Spath) differs from P. (Austiniceras) austeni in its smaller size, flexuous main ribs, convex flanks, evenly arched venter, and in that many of the intercalated ribs arise close to the umbilical shoulder. These are the exact features which are herein considered to be diagnostic of Anapuzosia and the author therefore has no hesitation in referring Spath’s species to this subgenus. The Albian species P. (Anapuzosia) saintoursi Collignon (19635: 61, pl. 263 (fig. 1150), 68, pl. 266 (fig. 1157)), P. (A.) colusaensis (Anderson) (1902: 96, pl. 5 (figs 128-129), pl. 10 (fig. 200)), P. (A.) multicostata Renz (1972: 707, pl. 2 (figs 1-2), pl. 3 (fig. 1), pl. 9 (fig. 4)) all have finer, more numerous intercalatories. Occurrence This species is known only from southern England and Angola. Subfamily Desmoceratinae Zittel, 1895 Genus Desmoceras Zittel, 1884 Type species Ammonites latidorsatus Michelin, 1836 1884 Desmoceras Zittel 1908 Desmoceras (Latidorsella) Jacob 1925 Phyllodesmoceras Spath 1938 Desmoceras (Pseudouhligella) Matsumoto (nom. nud.) 1942a Desmoceras (Pseudouhligella) Matsumoto 1947 Lunatodorsella Breistroffer 1954a Onitshoceras Reyment 1971 Desmoceras (Moremanoceras) Cobban Discussion Wiedmann & Dieni (1968) followed Wright (in Arkell et al. 1957) in including Latidorsella and Phyllodesmoceras within the synonymy of Desmoceras s.s., while retaining Lunatodorsella and Pseudouhligella as valid subgenera. Desmoceras (Moremanoceras) Cobban (type species Tragodesmoceras scotti Moreman, 1942: 208, pl. 33 (fig. 8)) was said to differ (Cobban 1971: 5) from Desmoceras ‘chiefly by having a simpler suture with broad lateral lobe and shorter auxiliary lobes. Constrictions, which are present by the juvenile stages, disappear and then are rejuvenated in the late adult stage.’ Moremanoceras was also said to resemble the Coniacian Onitshoceras in form and sculpture, but with a very different suture-line. ANNALS OF THE SOUTH AFRICAN MUSEUM 80 TX “S6LTWOd-INVS “(©4ZOI[OIS) DuDISaquof DjJaIUeA *Q “S~LCWOd-WVS ‘D “8pLCVOd WYVS “4 “(yyeds) 1dajqip (vIsozndvuy) visoznd ‘Q-A “WIT “Bid OSTe 998 “yOLCVWOd-WVS ie OO <(jaiURI\]) advjnaiavu (sp4az094]DD) Sd1ad0IADD “YW “ST “SI UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 81 Onitshoceras Reyment (type species O. matsumotoi Reyment, 19546: 249, pl. 3 (figs 1-2), pl. 5 (fig. 7)) was diagnosed by Reyment (1955: 19) as follows: ‘Flanks slightly inflated, venter broad, gently rounded, umbilicus narrow. Ornament consists of numerous irregular prorsiradiate ribs, swinging forwards and thickening on the venter. In the young they begin at mid-flank, but in the adult at the umbilical margin.’ The differences between Onitshoceras and Desmoceras are no greater than between the latter and Moremanoceras, and consequently Reyment’s genus is probably also best treated as a subgenus of Desmoceras. The differences between the various subgenera of Desmoceras are generally subtle and of questionable taxonomic value. At present they are no more than chronostratigraphic taxa. However, until objective population studies are undertaken, and the intraspecific and ontogenetic variation within the various subgenera better understood, the following subgenera are recognized within Desmoceras: D. (Desmoceras). Shell inflated, moderately to very involute, with depressed subquadrate to suboval whorl section. Ornament comprises strong sig- moidal ribs (constrictions on the internal mould), between which are dense growth striae or weak ribs on the outer part of the flanks and venter. Age: Upper Aptian—Upper Cenomanian. D. (Lunatodorsella). Similar to D. (Desmoceras), but shell cadicone. Doubt- fully separable from Desmoceras s.s. Age: Upper Albian. D. (Pseudouhligella). Similar to D. (Desmoceras), but with biconcave con- strictions, while whorl section generally compressed in maturity, and fre- quently with narrowly arched venter. Possibly dimorphic. Suture con- siderably incised, with symmetrical trifid lateral lobe and numerous auxiliary elements decreasing in size away from the lateral lobe. Age: Upper Albian— Upper Cenomanian. D. (Onitshoceras). Similar to D. (Desmoceras), but with fine irregular ribs arising near the umbilicus and strengthening across the venter. Suture-line with very narrow lobes and one or more saddles projecting beyond line of suture. Age: Coniacian. D. (Moremanoceras). Similar to D. (Onitshoceras), but possessing simpler suture, with broad lateral lobe and short auxiliary lobes. Age: Uppermost Cenomanian. Subgenus Pseudouhligella Matsumoto, 1942 Type species Desmoceras whiteavesi var. japonica Yabe, 1902 Desmoceras (Pseudouhligella) aff. ezoanum Matsumoto, 1942 Fig. 16 Compare Desmoceras toucasi Douvillé (non Jacob), 1931: 40, pl. 2 (fig. 6a—b). Desmoceras (Pseudouhligella) ezoanum Matsumoto, 1942a: 26, text-fig. 1b; 19546: 260, pl. 3 (figs 1-4). 82 ANNALS OF THE SOUTH AFRICAN MUSEUM Description Douvillé’s (1931) example, the whereabouts of which is unknown, would seem to belong here. Judging from Douvillé’s figure (Fig. 16), the specimen clearly has the recrystallized test preserved and is rather involute (umbilicus 25% of the diameter), with a rounded? umbilical shoulder, convex flanks, and a narrowly arched venter. The whorl section is compressed, with maximum width below mid-flank. The shell is ornamented only with flexuous growth striae and frequent, slightly prorsiradiate, flexuous constrictions which are weakly projected across the venter. Discussion Unfortunately, the exact characters of the shell cannot properly be judged from Douvillé’s figure (Fig. 6), but the Angolan specimen appears to be related to D. (P.) ezoanum Matsumoto (1942a: 26, text-fig. 1b; 19545: 260, pl. 3 (figs 1-4)), from which it differs largely in possessing a (?) rounded and not angular umbilical shoulder. In this respect it approaches D. (P.) poronaicum Yabe (Matsumoto 19545: 259, pl. 3 (figs 5—7)) which differs, however, in having an elliptical whorl section. Noteworthy is the fact that Yabe (1902) considered D.(P.) poronaicum possibly a juvenile D. (P.) japonicum Yabe, whilst Matsumoto (19545) at first regarded it as an extreme variant of D. (P.) ezoanum. In Japan, D. (P.) japonicum Yabe, D. (P.) poronaicum Yabe, and D. (P.) ezoanum Matsu- moto are contemporaneous species in the Paleogyliakian (Cenomanian); D. (P.) poronaicum is based upon juvenile material (less than 25 mm diameter) which may represent the inner whorls of either D. (P.) japonicum or D. (P.) ezoanum; D. (P.) japonicum is based upon large (118 mm diameter) adult material charac- terized by frequent constrictions and flat parallel flanks, while D. (P.) ezoanum Fig. 16. Desmoceras (Pseudouhligella) aff. ezoanum Matsumoto. The specimen figured by Douvillé (1931) as Desmoceras toucasi Jacob (after Douvillé 1931). x1. UPPERMOST CENOMANIAN-BASAL. TURONIAN AMMONITES FROM SALINAS, ANGOLA 83 is based upon medium-sized (52 mm diameter) adult material which differs from D. (P.) japonicum in that the adult whorls are more compressed, with convergent flanks and a narrowly arched venter, and fewer less prominent constrictions. Matsumoto’s (19545) variant, D. (P.) japonicum compressior (= D.(P.) japonicum s.s.) was said to differ from D. (P.) ezoanum only in having flat parallel and not convergent flanks. Clearly these differences are due to a comparison of the juvenile whorls of D. (P.) japonicum with the adult whorls of D. (P.) ezoanum. It is doubtful, therefore, whether the inner whorls of D. (P.) japonicum and D.(P.) ezoanum can be satisfactorily distinguished. The fact that D. (P.) ezoanum attains maturity at smaller diameters than D. (P.) japonicum, and is characterized by a change in shape of the adult whorls would seem to suggest that the differences are those between sexual dimorphs. Clearly, however, objective population studies on the Japanese material are required to confirm this suggestion. The Upper Albian D. (P.) dawsoni Whiteaves (1884: 205, pl. 26 (fig. 1); 1900: 286, pl. 37 (fig. 3)) is more strongly compressed, with a narrower umbilicus (14-16% of diameter) and higher whorls, but the differences are slight. Material figured by Matsumoto (1959a) from Alaska clearly shows constrictions to increase in number and prominence on the body chamber. Desmoceras (Pseudouhligella) calabarense Reyment (1955: 18, pl. 2 (fig. 8), pl. 3 (fig. 1)) was said to differ from D. (P.) japonicum only in being more evolute and in having less strongly flexed constrictions. The latter clearly become more flexuous on the body chamber (cf. Matsumoto 1959a) and hence the Nigerian species differs only in having a slightly wider umbilicus (19 % at 53 mm diameter as opposed to 14-15% in D. (P.) japonicum). For the present, the Nigerian species is best treated as a subspecies of D. (P.) japonicum although population studies may show even such a separation to be superfluous. Superfamily ACANTHOCERATACEAE Hyatt, 1900 Family Acanthoceratidae Hyatt, 1900 Subfamily Mantelliceratinae Hyatt, 1903 Genus Calycoceras Hyatt, 1900 Type species Ammonites navicularis Mantell, 1822 1900 Calycoceras Hyatt 1926 Metacalycoceras Spath 1959 Lotzeites Wiedmann 1972 Conlinoceras Cobban and Scott 1972 Gentoniceras Thomel 1972 Newboldiceras Thomel 1972 Subeucalycoceras Thomel 1972 Mourreiceras Thomel Discussion Within recent years the rather broadly perceived genus Calycoceras has been subdivided into a number of subgenera (and genera). 84 ANNALS OF THE SOUTH AFRICAN MUSEUM The genus Metacalycoceras Spath (1926) was proposed for the specimen of Ammonites navicularis figured by d’Orbigny (1842, pl. 103). However, according to the Code of Zoological Nomenclature (p. 59), the type of ‘a nominal genus is a nominal species’ and hence Metacalycoceras becomes a junior objective synonym of Calycoceras. Lotzeites Wiedmann (type species Acanthoceras aberrans Kossmat, 1895: 202, pl. 14 (fig. 4)) was erected for small Cheloniceras-like homeomorphs, with a coronate whorl section and moderately wide umbilicus. Distant flank ribs are ornamented with sharp, pointed umbilical bullae and prominent lower ventro- lateral tubercles. The latter are joined across the broad, convex venter by paired ribs, between which is an intercalary. All ribs are ornamented with weak upper ventro-lateral and siphonal tubercles. As pointed out by Kennedy (1971: 70), however, some species referred to Lotzeites, e.g. C. (L.) bathyomphalum (Koss- mat), have outer whorls indistinguishable from C. subgentoni (Spath) while ‘there is every transition from bathyomphalum type nuclei to subgentoni type nuclei in contemporary populations from the Middle Cenomanian of southern England’. Moreover, according to Kennedy & Juignet (1973), the holotype of Calycoceras (Lotzeites) crassum Thomel appears to be a poorly preserved Acanthoceras hippo- castanum (J. de C. Sowerby). Gentoniceras Thomel (type species Ammonites gentoni Brongniart, 1822: 150, pl. 11 (fig. 6a—b)) was proposed as a subgenus of Calycoceras for forms characterized by having a much more compressed whorl section than Calycoceras S.S., With dense ribbing which is always dominant over the tuberculation. Ventral tuberculation is weak, with lower ventro-lateral and siphonal tubercles frequently lost at an early stage. C. (Gentoniceras) may be a synonym of the earlier C. (Hourcquiceras) Collignon (1939) (see discussion under Euomphaloceratinae). Calycoceras s.s. was diagnosed by Thomel (1972) as comprising strongly inflated forms, always with a very depressed whorl section and with prominent, robust ribbing. Ventral tuberculation weakens considerably on the outer whorls or may even be lost completely. Newboldiceras Thomel (type species Acanthoceras newboldi Kossmat, 1897: 111, pl. 12 (figs 2-3), pl. 14 (fig. 2)) was erected as a new genus within the Acantho- ceratinae for Calycoceras-like homeomorphs in which the whorl section is com- pressed, subquadratic. As in Calycoceras s.1., ribbing is generally fine and masked by the tubercles at early growth stages. Unlike Calycoceras, siphonal tubercles are retained to large diameters. Its characters are ‘quelque sorte hybrides entre ceux des genres Calycoceras et Acanthoceras’. It is as yet uncertain whether this group comprises mantelliceratinid homoeomorphs of Acanthoceras or acantho- ceratinid homoeomorphs of Calycoceras. At present, the writer is inclined to treat Newboldiceras as a subgenus of Calycoceras. Within Newboldiceras, Thomel (1972) erected the subgenera Subeucalyco- ceras (type species Acanthoceras baylei Pervinquiére, 1907= Acanthoceras sartha- cense Bayle, 1878: pl. 62 (figs 1-2)) and Mourreiceras (type species N. (M.) mourrei Thomel, 1972: 118, pl. 48). Subeucalycoceras was proposed for forms UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 85 with flat, subparallel flanks and dense, rather straight, rursiradiate ribbing, the flanks forming a right angle with the venter. Judging from Bayle’s (1878) figure, however, N. (S.) baylei (Pervinquiere) appears to be a Gentoniceras. Mourreiceras was erected for denser and more finely ribbed forms homoeomorphous with Euomphaloceras, but probably does not bear separation from Newboldiceras sensu stricto. Conlinoceras (type species Calycoceras (Conlinoceras) gilberti Cobban & Scott, 1972: 61, pl. 1, pl. 2 (figs 5—9, 13-18), pl. 3 (figs 5-7, 11)) was proposéd as a subgenus of Calycoceras for distantly ribbed forms, with a sloping umbilical wall and poorly developed umbilical tubercles. Within the genus Calycoceras sensu lato it now appears possible to recog- nize the following subdivisions: C. (Calycoceras). Strongly inflated forms with a very depressed whorl section and coarse, robust, rursiradiate ribbing, generally with prominent umbilical bullae. In some species, weakly developed ventral tuberculation persists to maturity. Age: Upper Middle Cenomanian—Uppermost Cenomanian. C. (Lotzeites). Small, Cheloniceras homoeomorphs with ribbing looped across the venter between prominent lower ventro-lateral tubercles. Between looped ribs are intercalatories. There are three rows of weakly developed tubercles along the venter. Age: Upper Cenomanian. C. (Conlinoceras). Rather compressed forms with coarse distant ribbing, a sloping umbilical wall and poorly developed umbilical bullae. Ventral tuberculation becomes obsolete at large growth stages. Age: Middle Cenomanian. C. (Newboldiceras). Compressed forms with flattish flanks and a subquad- rate whorl section. Ribbing rather fine, dense, masked by tuberculation on inner whorls. Ventral tubercles persist to large diameters and thus homoeo- morphous with Acanthoceras. Age: Middle-low Upper Cenomanian. C. (Gentoniceras). Compressed forms commonly with strongly tuberculate inner whorls and strongly ribbed, non-tuberculate outer whorls. Umbilicus wide, with steep umbilical walls and prominent umbilical bullae. Age: Middle Cenomanian. Calycoceras (Calycoceras) naviculare (Mantell, 1822) Figs 4L-M, 12A, 15A, 17, 18A-B Ammonites navicularis Mantell, 1822: 198, pl. 22 (fig. 5). Stoliczka, 1864: 73, pl. 39 (figs 2-4). Crick 1919: 154, pl. 4. Acanthoceras naviculare (Mantell) Kossmat, 1897: 11. Choffat, 1898: 72, pl. 4 (fig. 6a—b), pl. 6 (figs 1-2). Fagesia navicularis (Mantell) de Grossouvre, 1912: 26, pl. 3 (fig. 1). Metacalycoceras grossouvrei Spath, 1926: 431. ?Metacalycoceras boehmi Spath, 1926: 431. Acanthoceras borgesi Douvillé, 1931: 32, pl. 2 (figs 3-4). Calycoceras (Metacalycoceras) stoliczkai Collignon, 1937: 48. 86 ANNALS OF THE SOUTH AFRICAN MUSEUM Calycoceras naviculare (Mantell) Hyatt, 1900: 589; 1903: 113. Spath, 1923: 144; 1937: 278. Wright & Wright, 1951: 26. Arkell et al., 1957: L431, fig. 531, 6, fig. 532. Cobban, 1971: 13, pl. 1 (figs 1-3), pl. 10 (figs 1-8), pl. 11 (figs 1-5), pl. 12 (figs 1-2), pl. 13 (figs 1-5), pl. 14 (figs 1-3), pl. 15 (figs 1-2), pl. 16 (figs 1-2), pl. 17. Kennedy 1971: 71, pl. 33 (fig. 1a—b), pl. 34 (fig. la—b), pl. 35 (figs 1-2), pl. 36 (figs 1-4), pl. 37 (figs 1-3), pl. 47 (figs 1a—c, 3a-b, 5a_b). Thomel 1972253: Calycoceras borgesi (Douvillé) Wright & Wright, 1951: 26. Thomel, 1972: 55. ?Calycoceras obrieni Young, 1957: 1171, pl. 150 (figs 1-4). Calycoceras (Metacalycoceras) auspicum Anderson, 1958: 243, pl. 20 (fig. 8). Mantelliceras oregonense Anderson, 1958: 244, pl. 8 (fig. 4), pl. 14 (fig. 1). Calycoceras stoliczkai Collignon, Matsumoto, 1959b: 78, pl. 21 (fig. 1). Thomel, 1972: 56, pl. 17 (figs 8-10), pl. 18 (figs 5-7). ?Calycoceras boehmi (Spath) Wright & Wright, 1951: 26. Calycoceras grossouvrei (Spath) Collignon 1965: 10, pl. 3 (fig. 2). Diagnosis A coarsely ribbed, evolute (umbilicus 23-32% of diameter) Calycoceras, generally strongly inflated and commonly with a very depressed whorl section. There are between 30 and 50 rursiradiate ribs per whorl (rectiradiate in juveniles), generally alternating long and short. Except for prominent umbilical bullae on the long ribs and ventro-lateral angulations on some specimens, tuberculation is absent except at small growth stages (generally less than 50 mm diameter). Description This is a common form in the Angolan fauna, and all specimens have the recrystallized test preserved. The shell is strongly inflated (W/H = 1,31-1,68), with a deep, rather narrow umbilicus (22-30% of diameter), and broad, convex, slightly overhanging umbilical walls. The whorl section (Fig. 17) is strongly depressed, reniform, with a broadly rounded venter sometimes showing ventro-lateral angulations. Recti- radiate to slightly rursiradiate ribs begin at the umbilical seam and terminate in prominent bullae on the umbilical shoulder. From the bullae arise coarse, robust, slightly rursiradiate flank ribs which alternate with shorter intercalatories arising Fig. 17. Calycoceras (Calycoceras) naviculare (Mantell). Whorl sections. A. SAM—PCA2715. B. SAM-PCA2764. x1. UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 87 somewhat below mid-flank. There are generally about 30 ribs per whorl on the outer whorls, although juvenile whorls are somewhat more densicostate with up to 36 ribs per whorl. Ribs are broader than the interspaces. Ribs arising from a bulla on one flank are commonly intercalated on the opposing flank. In all the Angolan material, ranging from 34 mm to 115 mm in diameter, ventral tubercula- tion appears to be lacking, although distinct ventro-lateral angulation of the ribbing is present to large diameters. Measurements No. D H We Wi Wi/H Ui Uo SAM-PCA2717 87 32(37) 46(53) 42(42) 3,31 ? 42 SAM-PCA2715 84 35(42) 58(69) 53(63) 1,51 25(30) 44 re 53 Z(G) BCI) AY) Iss I (WSK) 4) 34 «=? 26 v ? 9 (26,4) 16 SAM-PCA2723. 95 40(42) 58(61) 54(57) 1,35 26 (27,3) 45 xd VW 3G) 452) 4l@o) lee 2l@s) 85 SAM-PCA2733 72 35 u z r 16 (22,2) 39 SAM-PCA2762 34 16(47) 22(64) 21(62) 1,31 10(30) 17 SAM-PCA2764 115 48(42) 73(63) 63(55) 1,31 30(26) 46 Discussion The degree of inflation and the rib density of the Angolan material seem to be rather variable. The confusion which has long surrounded Mantell’s species has recently been clarified by Cobban (1971: 13) and Kennedy (1971: 72). The writer cannot agree with Thomel’s (1972) finely drawn species limits and thus follows the previous authors in their treatment of this species. Calycoceras boulei Collignon (1937: 43, pl. 5 (figs 2-4), pl. 8 (figs 9-11)) is a somewhat earlier form, first appearing late in the Middle Cenomanian, and differing largely in the retention of upper and lower ventro-lateral tubercles to large diameters. Calycoceras guerangeri (Spath) was originally figured only in side view (Gueranger 1867, pl. 4 (fig. 4)) and without seeing the holotype, which is still extant, it is difficult to assess. As interpreted by Thomel (1972: 61) this species differs in being of somewhat earlier age, with a more compressed whorl section and 2-3 intercalatories between main ribs. Calycoceras bruni (Fabre) (1940: 230, pl. 8 (figs 1—-2)), the type of which is apparently lost (Thomel 1972: 60), shows the following features: shell inflated, with a depressed whorl section, and about 30 strong, slightly rursiradiate ribs on the outer whorl. On the adapical portion of the outer whorl, the ribs are slightly flexuous whereas adorally they are rigid. Main ribs arise from prominent umbili- cal bullae and are separated by 1-2 intercalatories. All ribs are ornamented with ventro-lateral and siphonal tubercles to large growth stages; but lateral tubercles appear to be lacking. It differs from C. naviculare, therefore, in being of older age and in the persistence of ventral tuberculation to large diameters. 88 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 18. A-B. Calycoceras (Calycoceras) naviculare (Mantell), SAM-—PCA2715. C—D. Watino- ceras coloradoense (Henderson) (2), SAM—PCA2730. E-F. Pseudocalycoceras angolaense (Spath), SAM-PCA2808. G-—H. Euomphaloceras (Kanabiceras) septemseriatum (Cragin), SAM-PCA2826. Note the (?)superficial resemblance to Metasigaloceras. x1 UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 89 Calycoceras robustum Thomel (1972: 63, pl. 19 (figs 1-5)) is a compressed form with robust, rursiradiate ribbing. Main ribs arise from prominent umbilical bullae and alternate with shorter intercalatories. Occasionally there are 2 inter- calatories between main ribs, with about 30 ribs on the outer whorl. Main ribs have lateral tubercles, while all ribs are ornamented with ventro-lateral and siphonal tubercles. Calycoceras letullieri Collignon (1964: 131, figs 1603-1604) is a strongly inflated form belonging to the boulei group. On the outer whorl there are 32 ribs, generally alternating long and short. Long ribs are ornamented with prominent umbilical, lateral, ventro-lateral and weak siphonal tubercles at small growth stages, but only the umbilical and lateral tubercles are retained to maturity. While the differences between C. boulei, C. bruni, C. letullieri and C. robus- tum are probably no more than the intraspecific variation observed in C. navicu- lare, the author hesitates to place the first 4 species into synonymy until objective population studies have been made. They form a closely related group of slightly older species, generally less inflated, in which ventral tuberculation is retained to large diameters. Calycoceras thieuloyi Collignon (1964: 133, figs 1605-1607) is a strongly inflated globose species. In the holotype, 22 prominent umbilical bullae give rise to coarse, robust, rursiradiate primary ribs with sporadic intercalatories, so that there were probably about 36 ribs on the outer whorl. At early growth stages there may be up to 3 intercalatories between main ribs. Lateral and ventral tuberculation is lacking at all growth stages figured. According to Kennedy (1971: 73), this species may be a Mantelliceras. In time, there may be some justification for recognizing a number of sub- species within C. naviculare. At present, however, population studies are not far enough advanced to allow such a grouping. Occurrence At present, C. naviculare is recorded from Japan, the Western Interior, California, Texas, England, France, Portugal, Angola, Madagascar and southern India. Subfamily Acanthoceratinae Hyatt, 1900 Genus Protacanthoceras Spath, 1923 Type species Ammonites bunburianus Sharpe, 1853 Protacanthoceras? sp. A Fig. 19C—D Compare Ammonites bunburianus Sharpe, 1853: 25, pl. 9 (fig. 3a-c). Pictet & Campiche, Ké60 Silo. Diener, 1925; 24: Protacanthoceras bunburianum (Sharpe) Spath, 1923: 144. Kennedy, 1971: 97 pl. 31 (fig. la-c). 90 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 19. A-B. Pseudocalycoceras angolaense (Spath). An internal mould, SAM—PCA2734. C_-D. ?Protacanthoceras sp., SAM-PCA2778. E-F. Watinoceras coloradoense (Henderson) (2), SAM-PCA2753. G-L. Euomphaloceras (Kanabiceras) septemseriatum (Cragin). G—H. SAM-—PCA2736. Note prominent umbilical bullae, break-up of keel into isolated tubercles, and virtual disappearance of ventral sulci. I-J. SAM-—-PCA2756. K-L. SAM-PCA2742, a (?)pathological specimen showing abrupt loss of ventral ornamert. XI. UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 91 Description A single, rather poorly preserved example, with the recrystallized test preserved, shows affinities to Sharpe’s species. The shell is strongly compressed (W/H = 0,66), involute, with flat, parallel flanks and a narrow umbilicus (22 % of diameter). The venter is narrow, with an evenly rounded intercostal section, and is ornamented by three rows of closely spaced, weakly clavate tubercles situated on weak ribs. The lower ventro-lateral tubercles are very indistinct. Ribbing seems to be lacking on the lower flanks, first appearing at the lower ventro-lateral tubercle and becoming distinct only across the venter. On the outer whorl, there are about 18 ribs across the venter per half whorl. Measurements No. D H W W/H Ui SAM-—PCA2778 23 9 (39) 6 (26) 0,66 5 (22) Discussion The above specimen is only tentatively referred to this genus because of its poor preservation. From what can be seen of the Angolan example, it cannot be distinguished from P. bunburianum. The author’s reservations stem from the fact that laterally crushed inner whorls of Pseudocalycoceras angolaense would probably be equally difficult to distinguish from P. bunburianum. Protacanthoceras? sp. B Fig. 14D-E Description This specimen is also tentatively assigned to Protacanthoceras, although the preservation leaves much to be desired. It is small, involute and compressed, but appears to be laterally crushed. The little that is preserved of the ornament shows there to have been rather prominent rectiradiate ribs, at least on the outer flanks, with small umbilical and lower ventro-lateral tubercles and three rows of clavi closely spaced across the venter. Discussion The relatively coarse ribbing of this example would appear to distinguish it from juveniles of Pseudocalycoceras angolaense (Spath). Consequently, the rather coarse ornament is taken to indicate closer affinities with Protacanthoceras than Pseudocalycoceras. Protacanthoceras compressum (Jukes-Browne) is similar, but has fewer more prominent ribs, with distinct umbilical and lower ventro-lateral tubercles. 92 ANNALS OF THE SOUTH AFRICAN MUSEUM Genus Tarrantoceras Stephenson, 1955 Type species Tarrantoceras rotatile Stephenson, 1955 1955 Tarrantoceras Stephenson 1969 Sumitomoceras Matsumoto Discussion Tarrantoceras was originally (Stephenson 1955: 59) separated from Eucalyco- ceras on differences in the suture-line. In the emended diagnosis given by Cobban & Scott (1972: 64), this genus comprises moderately large, compressed forms (Fig. 20) in which prominent prorsiradiate flank ribs are ornamented with umbilical, upper and lower ventro-lateral and siphonal tubercles. Of these, the upper ventro-lateral tubercles are the most prominent, rising above the level of the siphonal tubercles. The latter are very weakly developed in the adult growth stages and are finally lost about half-way up the body chamber, while the lower ventro-lateral tubercles are lost at the start of the body chamber. Fig. 20. Tarrantoceras rotatile Stenhenson. The holotype, after Stephenson (1955). x1. Sumitomoceras Matsumoto (in Matsumoto et al. 1969: 283) was erected for moderately evolute forms with flat flanks and a compressed whorl section. Long ribs arise from umbilical tubercles and are separated by 1-2 intercalatories. All ribs are prorsiradiate and ornamented with upper and lower ventro-lateral tubercles, while at very immature growth stages weak siphonal tubercles are present. In the middle and late growth stages the ventro-lateral tubercles weaken and disappear, constrictions develop and this genus homoeomorphs Kossmaticeras. However, the holotype of Tarrantoceras multicostatum Stephenson (1955: 61, pl. 6 (figs 21-23)), considered a synonym of T. rotatile Stephenson by Cobban UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 93 & Scott (1972), shows the same loss of tuberculation on the body chamber as does Sumitomoceras, whilst the juvenile whorls are also comparable. The author regards Sumitomoceras as a junior subjective synonym of Tarrantoceras. Another genus closely allied to Tarrantoceras is Eucalycoceras Spath. As diagnosed by Kennedy (1971: 80), this genus comprises involute forms with com- pressed whorls and broad flattish flanks. Ribbing is rather dense, generally with umbilical, upper and lower ventro-lateral and siphonal tubercles. At large growth stages, some or all of the ventral tubercles may be lost. Tarrantoceras is thus very close to Eucalycoceras. Besides the sutural differences noted by Stephenson (1955), Tarrantoceras seems invariably to lose its siphonal tubercle and to be adult at smaller diameters than Eucalycoceras, whilst the latter genus is typically higher-whorled and more involute. Other Upper Cenomanian genera with five rows of tubercles across the venter are the micromorph Protacanthoceras and Pseudocalycoceras, the latter distinguished by its flexuous, rursiradiate ribbing. Tarrantoceras ? sp. juv. indet. Fig. 21D-E Description A single small, fragmentary specimen with recrystallized test preserved is assigned to this genus. The shell is moderately compressed, with a polygonal costal and rounded intercostal section. It would appear to have been moderately evolute. There are small but distinct umbilical tubercles from which arise prorsiradiate main ribs. Between long ribs there is almost invariably a short rib intercalated at about mid- flank. All ribs bear prominent lower ventro-lateral tubercles and slightly clavate upper ventro-lateral and siphonal tubercles. Discussion The solitary Angolan fragment closely resembles juvenile forms of Eucalyco- ceras, such as were figured by Kennedy (1971). So far as the author is aware, however, this genus has not been recorded from strata of this age (top Ceno- manian), and thus he prefers to refer the Angolan specimen to the closely allied Tarrantoceras. At a similar growth stage, Pseudocalycoceras angolaense (Spath) is very weakly ribbed or smooth, while species of Protacanthoceras lack the strong flank ribbing seen in the Angolan specimen. Genus Pseudocalycoceras Thomel, 1969 Type species Ammonites harpax Stoliczka, 1864 1969 Pseudocalycoceras Thomel 1969 Neocalycoceras Thomel 1972 Pseudocalycoceras (Haugiceras) Thomel ANNALS OF THE SOUTH AFRICAN MUSEUM 94 "TX “p8LZ7VOd-WYS “Ant ‘oR][NG [eoI[Iquin yeoM AISA 910N “YIL7WOd-WVS = CO yy ‘ds spsav0juvAsv I, “A-A ‘WeAP] Wwnsoggis spsav02N0JaW “D-V “I~ “SI UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 95 Discussion This genus was erected (Thomel 1969: 651) for moderately evolute forms with coarse rursiradiate, flexuous ribbing and five rows of tubercles across the venter. Differences from other genera are noted under the discussion of Watinoceras. Neocalycoceras was erected without description or diagnosis and is thus a nomen nudum. Haugiceras (type species Acanthoceras haugi Pervinquiere, 1907: 270, pl. 14 (fig. la—b)) (Fig. 22) was proposed as a subgenus of Pseudocalycoceras for rather inflated forms in which the whorl section was wider than high, the ribs were more rigid and less robust, rursiradiate, but less recurved than in Pseudo- calycoceras s.s. The tuberculation was said to be entirely different, in Haugiceras the umbilical tubercles are not twisted into an S, while the tubercles across the venter are rounded and not clavate, and closer together. Into this subgenus, Thomel (1972) placed Pseudocalycoceras angolaense (Spath), P. haugi (Pervin- quiere) and P. robustum Thomel. However, Thomel’s separation of Haugiceras from Pseudocalycoceras appears to be artificial, and the author regards the two names as synonyms. Fig. 22. Pseudocalycoceras haugi (Pervinquiére). The holotype, after Pervinquiére (1907). x1. 96 ANNALS OF THE SOUTH AFRICAN MUSEUM Thomel (1972) also included a number of typical Pseudocalycoceras species under his heterogenous group of Eucalycoceras (Proeucalycoceras) (type species Calycoceras (Eucalycoceras) besairiei Collignon, 1937: 13, pl. 3 (figs 1-4), pl. 8 (fig. 5)). This subgenus was erected for moderately evolute forms with a com- pressed, quadrate whorl section, flat flanks, and dense, fine, flexuous ribbing at immature growth stages. Ribs are ornamented with small umbilical bullae, upper and lower ventro-lateral tubercles, of which the former are more prominent, and siphonal tubercles which are lost at an early stage. Of the species assigned to this subgenus, Jarrantoceras bentonianum (Cragin), Pseudocalycoceras dentonense (Moreman) and its synonym P. /ewisvillense (Moreman) (cf. Cobban & Scott 1972), together with Calycoceras leonense (Adkins) and Mantelliceras lymense (Spath) do not belong here. As presently defined, this taxon (Proeucalycoceras) is of little value. Pseudocalycoceras angolaense (Spath, 1931) Figs 4A—C, H-K, 6I-J, 10F—G, 14A, 18E—-F 19A-B, 23-25, 26F—K Acanthoceras rhotomagense Taubenhaus (non Brongniart), 1920: 13. Acanthoceras sp. A, Moreman, 1927: 95, pl. 15 (fig. 2). Acanthoceras lyelli Douvillé (non Leymerie), 1931: 31, pl. 1 (fig. 1). Protacanthoceras angolaense Spath, 1931: 316. Basse, 1940: 448, pl. 6 (fig. 3). Avnimelech & Shoresh, 1962: 531. Eucalycoceras dentonense Moreman, 1942: 205, pl. 33 (figs 4-5), text-fig. 2k. Matsumoto, 19595: 97, text-fig. 51. Eucalycoceras indianense Moreman, 1942: 206, pl. 33 (figs 9-10), text-fig. 21. Matsumoto, 1959b: 98. Eucalycoceras lewisvillense Moreman, 1942: 206, pl. 33 (figs 6-7), text-fig. 2n, u. Eucalycoceras underwoodi Powell, 1963a: 315, pl. 31 (fig. 17). Pseudocalycoceras (Neocalycoceras) angolaense (Spath) Thomel, 1969: 651. Eucalycoceras (Proeucalycoceras) dentonense Moreman, Thomel, 1969: 650. Eucalycoceras (Proeucalycoceras) lewisvillense Moreman, Thomel, 1969: 650. ‘Protacanthoceras’ angolaense Spath, Kennedy, 1971: 115. Pseudocalycoceras angolaense (Spath) Cooper, 1972: 248; 1974: 86. Pseudocalycoceras (Haugiceras) angolaense (Spath) Thomel, 1972: 97. Pseudocalycoceras dentonense (Moreman) Cobban & Scott, 1972: 63, pl. 13 (figs 11-29), pl. 15 (figs 1-7, 10-13). Pseudocalycoceras sp. aff. P. dentonense (Moreman) Matsumoto & Kawano, 1975: 13, pl. 1 (fig. 1), text-fig. 3. Description The shell is moderately evolute (umbilicus 18-41 % of diameter), somewhat compressed (W/H = 0,67-1,12) to slightly depressed, with a fairly wide, deep umbilicus. The best preserved specimen, SAM-—PCA2731 (Fig. 26I-K), retains recrystal- lized test, and is entirely septate to 57 mm diameter. About one-third of the preceding whorl is covered to just below the lower ventro-lateral clavi by the outer whorl. The umbilicus is fairly broad and deep, with a steep umbilical wall, slightly undercut at the seam, and an abruptly rounded umbilical shoulder. The UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 97 Fig. 23. Pseudocalycoceras angolaense (Spath). The holotype (after Douvillé 1931). x1. whorl section is slightly compressed, almost quadrate, with a polygonal costal section. On the outer whorl of SAM—PCA2731, about 18 ribs are weakly developed on the umbilical wall and pass radially to the umbilical shoulder where they give rise to 1—2 flank ribs. The umbilical tubercles are irregularly developed, and there are only 12 on the outer whorl. The flank ribs are flexuous, rursiradiate, and number about 30 on the outer whorl. At the prominent lower ventro-lateral clavi the ribs flex forwards to the upper ventro-lateral clavi and then cross the venter transversely. All ribs bear prominent siphonal clavi. The penultimate whorl of this specimen is very weakly ornamented and shows the earliest whorls to have lacked ribbing, at least on the lower portion of the flanks, and umbilical tubercles to a diameter of about 20 mm. Together with the above specimen, three other examples, SAM—PCA2767, SAM-PCA2777 and SAM-PCA2772, provide an almost complete ontogenetic sequence for this species. In SAM—PCA2767 (Fig. 4J—K) the flanks are entirely smooth to a diameter of 11 mm, at which stage, in SAM—PCA2777 (Fig. 4A—C), ventral tuberculation appears, as well as very faint ventral ribbing. By 15 mm diameter the ribbing has become more prominent, but is still confined to the outer flanks, while the umbilical shoulder bears occasional weak umbilical tubercles. At this stage the whorl section is oval, with slightly convex flanks con- verging to the narrowly arched venter. In costal section the venter is almost tabulate and trituberculate. In this same specimen the lower ventro-lateral 98 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 24. Pseudocalycoceras angolaense (Spath). Whorl sections. A. SAM-PCA2731. B. SAM—PCA2767. C. SAM-PCA2732. x1. tubercles, although faint, are evident at 15 mm diameter, whereas in SAM-—PCA2767 they are still absent at 28 mm diameter, whilst the latter also shows no signs of umbilical tubercles at the same stage. In SAM—PCA2772 (Fig. 26G), at about the same diameter, the umbilical tubercles are weak but noticeable, with upper and lower ventro-lateral and siphonal tubercles. The ribbing is fairly strong but is not connected to the umbilical tubercles. In SAM-—PCA2731 the umbilical tubercles become prominent at about 28 mm diameter. At about this same stage the flanks become flat, or slightly convex, and almost parallel, with the greatest width at mid-flank. In this same specimen there are 11 siphonal clavi within 40 mm along the venter at a diameter of 35 mm, but only 6 at 57 mm diameter. In SAM—PCA2734, with about one-third of a whorl of body chamber preserved, there are only 5 siphonal clavi in an equal distance at 68 mm diameter. This specimen also shows, as does the holotype (Fig. 23), the simple, coarse, straight ribbing on the body chamber. Fig. 25. Pseudocalycoceras angolaense (Spath). Suture-line of SAM-—PCA2719 at about 60 mm diameter. UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 99 A further example, SAM-—PCA2721, is coarsely ornamented at an early growth stage, providing further evidence of the great variability in ontogenetic development of this species. It is already showing well-developed umbilical tubercles at a diameter of about 13 mm, while at 25 mm diameter ribbing is already prominent and connected to the umbilical tubercles. This specimen is also slightly more inflated than typical examples. Measurements No. D H W W/H Uo Ui SAM-PCA2731 a1 24 (42) 23 (40) 0,96 U3) 17 (30) x 44 21 (48) 20 (45) 0,95 7 13 (30) SAM-—PCA2721 41 16 (39) 18 (44) ee, 17 12 (29) i 29 BE WE Le 9 9 (41) SAM-PCA2772 37 iV@e 15Gb G88 15 10 (27) ‘ 25 DES) 8665) O75 8 6 (24) SAM-PCA2777 DD 11 (50) 816) 0a) oe 4 (18,2) i 15 9(60) 6(49) 067 ? BB) SAM-PCA2734 1 20(G) MGS) O85 S80 BAe) : AON (A). Me? = 19 14 (30) SAM-PCA2767 3 ISS) WO On 2 9 (24,3) r 10 6(60) 4(40) 067 2 2 (20) Discussion Cobban & Scott (1972) have dealt comprehensively with Pseudocalycoceras dentonense (Moreman), including P. /ewisvillense (Moreman) and P. indianense (Moreman) within the synonymy of this species. The North American material differs from the Angolan material only in that many individuals have more coarsely ribbed inner whorls, while the ventro-lateral and siphonal tubercles generally weaken and become obsolete between 25 and 80 mm diameter. The differences are clearly not specific, and P. dentonense is considered a junior sub- jective synonym of P. angolaense. Eucalycoceras underwoodi Powell (1963a: 315, pl. 31 (fig. 17)) is a contemporaneous species which also does not bear separation from P. angolaense. Pseudocalycoceras haugi (Pervinquiére) (Fig. 22) is more coarsely ribbed, with more prominent umbilical tubercles, less flexuous ribbing, and more swollen lower ventro-lateral tubercles. It represents a larger growth stage than observed in P. angolaense, and until its ontogenetic and intraspecific variation is known, I prefer to regard them as distinct. Thomel (1972) described a number of Pseudocalycoceras species from south- east France. Of these, P. equituriense Thomel, P. flandrini Thomel and P. planum Thomel appear better referred to the genus Thomelites. Pseudocalycoceras dromense Thomel is possibly better assigned to Lotzeites. The holotype of Pseudocalycoceras pseudorbignyi Thomel was not figured, while the plesiotype is a body chamber fragment which is difficult to assess. According to Thomel 100 ANNALS OF THE SOUTH AFRICAN MUSEUM (1972: 95), the holotype is about one-half whorl of an internal mould. The shell is rather evolute (umbilical ratio 32%), with flat subparallel flanks. On the outer whorl, there are 17 flexuous ribs per half whorl, of which 7 arise from bullae at the umbilical shoulder. The venter is narrow and ornamented with three rows of small tubercles. Pseudocalycoceras robustum Thomel (1972: 98, pl. 47 (figs 6-7)) is a moderately evolute species. The inner whorls (up to 36 mm diameter) are com- pressed, with the maximum width near the umbilical shoulder. Ornament com- prises alternating long and short ribs, rigid, robust, and totalling 25 on the outer whorl. Main ribs arise from prominent, rounded umbilical tubercles, while all ribs are ornamented with upper and lower ventro-lateral and siphonal clavi. At large diameters the siphonal tubercles are lost and the ribs rise to form a sharp crest (? crushed) on the venter. It differs from P. angolaense in having more robust, less flexuous ribs which do not arise in pairs from the umbilical shoulder, and is thus probably a junior subjective synonym of P. haugi. Pseudocalycoceras judaicum (Taubenhaus) (Avnimelech & Shoresh 1962: 531, pl. 15 (fig. 2)) has similar alternating long and short flank ribs, about 16 per half whorl, but is strongly crushed laterally and difficult to compare. It is probably not separable from P. haugi. Pseudocalycoceras harpax (Stoliczka 1865: 72, pl. 39 (figs 1, la—b)) is a moderately evolute (umbilical ratio 27-30%), compressed species, with flattish, slightly convex flanks. Main ribs arise from umbilical tubercles (initially rounded but later bullate) and are flexuous, rursiradiate, with 1-2 intercalatories sepa- rating main ribs. The evenly rounded venter is ornamented with 5 rows of closely spaced tubercles, upper and lower ventro-lateral and siphonal clavi. There are 38 ribs on the outer whorl, of which 15 are main ribs. This appears to be a Middle or low Upper Cenomanian species which differs from P. angolaense in that main ribs do not commonly arise in pairs from the umbilical tubercles, the whorl section is more rounded, not distinctly polygonal as in P. angolaense, and the ventral tuberculation is not as prominent nor so markedly clavate. The forms assigned by Thomel (1972) to P. harpax and its varieties appear to belong to other species. Occurrence Pseudocalycoceras angolaense has been recorded from Texas, the Western Interior, Angola, northern France, Japan and Israel. Pseudocalycoceras aff. haugi (Pervinquiere, 1907) Figs 1OH-I, 22, 27 Compare Acanthoceras haugi Pervinquiére, 1907: 270, pl. 14, figs la—b. Protacanthoceras judaicum (Taubenhaus) Avnimelech & Shoresh, 1962: 531, pl. 15 (fig. 1) (holotype refigured). ————— UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 101 Fig. 26. A-B. Euomphaloceras (Kanabiceras) septemseriatum (Cragin), SAM-—PCA2827, x1. C-E. Undetermined gasteropods. F—K. Pseudocalycoceras angolaense (Spath), x 1. F-H. SAM-PCA2721. I-K. SAM-PCA2731. 102 . ANNALS OF THE SOUTH AFRICAN MUSEUM Protacanthoceras aft. compressum Avnimelech & Shoresh (non Jukes-Browne), ISLS S8V2, ols NS CUS, 2), Pseudocalycoceras (Haugiceras) robustum Thomel, 1972: 98, pl. 47 (figs 6-7). Description A rather poorly preserved Angolan example shows affinities to Pervinquiére’s species. The preservation of the Salinas material, together with the fact that the Tunisian form represents a different ontogenetic stage, makes comparison difficult. The specimen has the recrystallized test preserved. The Angolan example is strongly compressed and moderately evolute, with about half the preceding whorl covered. The umbilicus is fairly wide and rather shallow, with a steep umbilical wall and subrounded umbilical shoulder. The flanks are broad and flat, almost parallel, with a narrow, evenly rounded venter intercostally, but polygonal costally. The venter is ornamented with three rows of closely spaced clavi. The flank ornament is seen to comprise almost radial, alternating long and short ribs, although there may occasionally be two inter- calatories between main ribs. All the ribs have weakly developed lower ventro- lateral tubercles and prominent upper ventro-lateral and siphonal clavi, while the long ribs also have sporadic, faint umbilical tubercles. The last rib on the adoral portion of the outer whorl shows a fairly distinct umbilical tubercle, which would seem to suggest that they became more prominent with age. There are about 15 ribs per half whorl on the outer whorl. Measurements No. D H W W/H Ui SAM-—PCA2775 32 16 (50) ONG) 0,63 8 (25) as 2S) 11 (44) 7 (28) 0,64 6 (24) Discussion As noted under the discussion of P. angolaense, the species P. haugi, P. robustum and P. judaicum (Fig. 27) are probably not specifically separable, while the above material differs from P. angolaense in being more coarsely ornamented at small diameters, with less flexuous, rigid flank ribs. The close resemblance to P. judaicum probably relates to the fact that the Angolan material has also been laterally crushed. Pseudocalycoceras harpax (Stoliczka) has denser, flexuous ribbing. Occurrence Pseudocalycoceras haugi appears to be present in Tunisia, southern France, Israel and possibly Angola. Subfamily Euomphaloceratinae. New subfamily Diagnosis Multituberculate derivatives of Acanthoceras, typically evolute, with a depressed, subquadrate to compressed, elliptical whorl section, and more or less UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 103 Fig. 27. Pseudocalycoceras cf. haugi (Pervinquiére). < 1. The holotype of Protacanthoceras judaicum (Taubenhaus), after Avnimelech & Shoresh (1962). x 1. prominent constrictions on the early whorls. In almost all genera, there are umbilical, upper and lower ventro-lateral and siphonal tubercles, which in some genera are supplemented by one to two rows of lateral tubercles. Range: Middle Cenomanian (possible, but doubtful, Lower Cenomanian members are discussed below) to uppermost Turonian. Discussion As herein interpreted, the Euomphaloceratinae includes the following genera: Euomphaloceras Spath, Kanabiceras Reeside & Weymouth, Kameruno- ceras Reyment, Schindewolfites Wiedemann, Romaniceras Spath, Yubariceras Matsumoto, Saito & Fukada, Shuparoceras Matsumoto and Obiraceras Matsu- moto. Also tentatively included here is the Lower? Cenomanian Tunesites Pervinquiere. The origins of the subfamily lie in the group of Euomphaloceras cunningtoni (Sharpe) which was derived from depressed and spinose Acanthoceras rhoto- magense (Brongniart) during Turrilites costatus Zone times, by a weakening of the flank ribs and the development of intercalated ventral ribs (Kennedy 1971). Acanthoceras rhotomagense pseudeuomphalum Thomel may be a passage form. Euomphaloceras s.s., i.e. euomphalum (Sharpe), arose from this stock during pentagonum Zone times by a further weakening of ribbing relative to tubercula- 104 ANNALS OF THE SOUTH AFRICAN MUSEUM tion, and the appearance of constrictions in the early and middle growth stages (it should be noted that the earliest growth stages of FE. cunningtoni are unknown and it too may possess constrictions on the juvenile whorls). Kanabiceras first appears in the uppermost Cenomanian (gracile Zone), and is easily derived from E. euomphalum merely by a joining of the siphonal -tubercles so as to form a serrated keel. Passage forms are known (Kennedy 1971). During the Lower Turonian, Kanabiceras gave rise to Schindewolfites and Kamerunoceras. Kamerunoceras Reyment is known with certainty only from the Lower Turonian of Nigeria and the Cameroons. Reyment (1955) regarded it as a probable offshoot of the Protacanthoceras stock that produced Mammites, Watinoceras and Benueites. The inner whorls of Kamerunoceras, however, bear a low crenulated keel with shallow sulci on either side, which therefore link it rather to the uppermost Cenomanian Kanabiceras. Schindewolfites Wiedmann (1959) includes evolute forms with constricted early whorls, irregular ornament, and prominent tuberculation. In maturity, this genus may develop a row of lateral tubercles in addition to the umbilical, upper and lower ventro-lateral, and siphonal tubercles. The inner whorls of this genus bear a close morphological resemblance to Kanabiceras and a close phyletic relationship seems probable. Thomel (1972: 158) considered Acanthoceras ver- gonsense Thomel as the probable ancestor of Schindewolfites; this appears wholly untenable since there is a considerable temporal gap of several ammonite zones between the last appearance of Acanthoceras and the first appearance of Schinde- wolfites. Matsumoto (1975) suggested that Schindewolfites might best be treated as a Subgenus of Yubariceras. Matsumoto ef al. (1957) interpreted Yubariceras as a multituberculate derivative of Acanthoceras s.s. although, more recently, Matsumoto (1975) con- sidered the origins of the group should be sought in what he termed ‘. . . a certain species group of Euomphaloceras as well as in a transitional form between Acanthoceras and Euomphaloceras’. It is difficult to derive Yubariceras from either Acanthoceras or Euomphaloceras s.s. since, as with Schindewolfites, there is a considerable time gap between the first appearance of Yubariceras and the last appearance of Acanthoceras or Euomphaloceras s.s. Yubariceras first appears in the Lower Turonian and, in common with Kanabiceras, is very evolute, with a depressed whorl section, multituberculate ornament, and it bears constrictions on the early whorls. It is thus in this latter taxon (or one of its early Turonian derivatives) that the ancestry of Yubariceras should be sought. Matsumoto ef al. (1957) considered Romaniceras as a multituberculate derivative from the group of Calycoceras newboldi—spinosum. These two groups are indeed superficially similar, although the resemblances are simply due to homoeomorphy since the newboldi group die out early in the Upper Cenomanian (pentagonum Zone) and Romaniceras does not appear until about the Middle Turonian. As noted by Matsumoto (1975), Romaniceras and Yubariceras are closely allied, and it seems more likely that the typically Middle to Upper Turonian Romaniceras was derived from an early form of Yubariceras. UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 105 Wiedmann (1959) erected the subgenus Romaniceras (Proromaniceras) for upper Lower Turonian forms allied to Calycoceras orientale (Kossmat). Yet again, the resemblance is merely one of homoeomorphy, and the author would follow Matsumoto (1975) in considering Proromaniceras as a synonym of Romaniceras S.s. Shuparoceras is a Lower to Middle Turonian genus “. . . allied to the sub- group of Calycoceras choffati in various respects’ (Matsumoto 1975: 110). The group of Calycoceras choffati (Kossmat) died out early in the Upper Cenomanian, and clearly did not provide the ancestor of Shuparoceras. Again, the constricted inner whorls, evolute form, and multituberculate ornament of this latter genus are typical of the Euvomphaloceras—Kanabiceras lineage, and it is here that the ancestry of Shuparoceras should be sought. Obiraceras is a late Turonian genus allied to both Yubariceras and Schinde- wolfites, and Matsumoto (1975) considers its derivation from the latter genus the most likely. From a morphological and stratigraphical point of view, this deriva- tion is quite acceptable. The author does not agree with Matsumoto (1975) that the Turonian multituberculate acanthoceratids are polyphyletic. Indeed, they form a closely allied (af excessively split) and intimately related group which clearly have their origins in the late Cenomanian Euomphaloceras—Kanabiceras lineage. Two other constricted acanthoceratid genera remain to be discussed. Tunesites Pervinquiere (1907) was erected for evolute (umbilicus 30-39% of diameter) pyritic nuclei, with compressed whorl sections. The flanks bear slightly flexuous, prorsiradiate ribs ornamented by upper and lower ventro-lateral and siphonal tubercles, while there are three more or less prominent constrictions per whorl. Pervinquiere (1907) gave the range of his genus as uppermost Albian—Cenomanian, whereas Wright (in Arkell et al. 1957) considers it as an exclusively Lower Cenomanian taxon. According to Pervinquiére’s (1907) original account, the holotype of 7. choffati Pervinquiere comes from beds yielding Forbesiceras obtectum (Sharpe), a typically Middle Cenomanian species (Kennedy 1971). In terms of acanthoceratid phylogeny, Tunesites shows advanced morphological features which are not compatible with an Upper Albian—Lower Cenomanian age and, as with some of Pervinquiere’s other stratigraphic data, the horizon is perhaps doubtful. Consequently, any attempt to fit Tunesites into a phylogenetic scheme must, at present, be speculative. Hourcquiceras Collignon (1939) was erected as a subgenus of Calycoceras for forms characterized by the presence of prominent constrictions on the juvenile whorls, and was considered a possible synonym of Tunesites by Wright (in Arkell et al. 1957). The subgenus is of Middle or Upper Cenomanian age, and closely resembles Calycoceras (Gentoniceras) in maturity. Collignon’s (1939) figures do not show obvious constrictions, only periodic more strongly developed flank ribs (an identical feature to that seen in juvenile Gentoniceras). As the writer has no material before him, further speculation is not possible, but he prefers to leave Hourcquiceras in the Mantelliceratinae. 106 ANNALS OF THE SOUTH AFRICAN MUSEUM Genus Euomphaloceras Spath, 1923 Type species Ammonites euomphalus Sharpe, 1855 1923 Euomphaloceras Spath 1931 Kanabiceras Reeside & Weymouth 1937 Cunningtoniceras Collignon Discussion According to Wright (in Arkell et al. 1957: L414), the genus Kanabiceras differed from Euvomphaloceras ‘in that paired tubercles on venter are oblique, not transverse, and siphonal tubercles now form a nodose keel’. Kanabiceras first appears in the uppermost Cenomanian and is easily derived from the earlier (pentagonum Zone) Euomphaloceras euomphalum (Sharpe) merely by a joining of the siphonal tubercles so as to form a serrated keel. Kennedy (1971: 90), how- ever, records intermediates between the two forms while noting that a nodose keel may sometimes be present in E. euomphalum (Sharpe). Moreover, with the wide range of variation seen in the Angolan material of Kanabiceras septem- seriatum (Cragin), and also admitted by other authors (Matsumoto 19595), together with the variability shown by E. euomphalum (Wright 1963), the distinc- tions between the two genera blur. The differences are not herein regarded as of generic status, and consequently Kanabiceras is treated as a subgenus of Euom- phaloceras. The genus Evomphaloceras now may be subdivided as follows: E. (Euomphaloceras). Shell very evolute, inflated, with depressed whorl sec- tion; prominent umbilical and lower ventro-lateral tubercles on some or all of main ribs. All ribs cross venter with small upper ventro-lateral and siphonal tubercles, more numerous than lower ventro-lateral tubercles. Inner and middle whorls with distinct constrictions in some species. Age: Middle-Upper Cenomanian. E. (Kanabiceras). As for E. (Euomphaloceras) but with siphonal tubercles joining to form a prominent keel. Upper ventro-lateral tubercles more or less oblique to keel. Age: Upper Upper Cenomanian—Lower Turonian. Subgenus Kanabiceras Reeside & Weymouth, 1931 Type species Scaphites ? septem-seriatus Cragin, 1893 Euomphaloceras (Kanabiceras) septem-seriatum (Cragin, 1893) Figs 4N—O, 10A-E, 12E-H, 18G—H 19G-L, 26A-B, 28 Scaphites ? septem-seriatus Cragin, 1893: 240. Adkins 1928: 259. Acanthoceras ? kanabense Stanton, 1893: 181, pl. 36 (figs 6-8). Acanthoceras kanabense Stanton, Moreman, 1927: 95, pl. 13 (fig. 5). Kanabiceras kanabense (Stanton) Reeside & Weymouth, 1931: 12. Neocardioceras septem-seriatum (Cragin) Moreman 1942: 213, pl. 33 (figs 11-12). Prionotropis echinatus Douvillé, 1931: 34, pl. 3 (figs 1-5), pl. 4 (figs 1-3). Neocardioceras (echinatum) (Douvillé) Spath, 1931: 316. UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 107 Kanabiceras septemseriatum (Cragin) Matsumoto, 19595: 99, pl. 24 (fig. la—c). Powell 1963a: 316, pl. 31 (figs 9-10). Matsumoto et al., 1969: 279, pl. 37 (figs 1-3). Cobban & Scott, 1972: 25 pila I2i(iss S27): Lyelliceras stanislausense Anderson, 1958: 247, pl. 8 (fig. 5). Diagnosis Shell very evolute (umbilicus 32-44% of diameter), inflated, with wide shallow umbilicus. Whorl section varies from almost quadrate to strongly depressed (W/H = 1,12-1,57). Ornament highly variable. Ribbing very weak to prominent, prorsiradiate, with prominent umbilical and lower ventro-lateral septate spines. Upper ventro-lateral spines or clavi more numerous than lower ventro-lateral spines. Ribbing projected strongly forwards on the venter so as to meet the serrated siphonal keel at an acute angle. The keel may be strongly serrated or only a row of weakly joined tubercles. On either side of the keel are sulci which may be very deep or almost non-existent. Intercalated ribs and striae of various strength are common on the flanks. Description This species is one of the most abundant members of the Salinas fauna and consequently the intra-specific variation can be more readily judged. The orna- mentation of this species is highly variable. All the specimens have their recrystal- lized shell preserved. The shell is very evolute, the preceding whorl being covered to the lower ventro-lateral tubercles, with a generally depressed whorl section; polygonal costally but with a rounded intercostal section. The umbilicus is wide and shal- low, the umbilical shoulder well rounded and occasionally impressed to accom- modate the prominent, but irregularly developed, lower ventro-lateral spines of the preceding whorl. The ribbing of this species is highly variable; in some examples the ribbing dominates the ornament (Fig. 4F—G) whereas in others ribbing is subordinate and spines predominate (Fig. 1OB—C). There is a morphological gradation of specimens between these two end members. In SAM-PCA2761 (Fig. 12E-F) ribs arise at the umbilical seam and are faintly rursiradiate to the umbilical bullae, which are most strongly developed on the earlier whorls. From these bullae the ribs pass forward (prorsiradiate) to the lower ventro-lateral tubercles, which are again very irregularly developed. On the earlier whorls they seem to have been, almost invariably, obliquely clavate, but as diameter increases they become swollen and rounded and many form the bases of long septate spines, although not every tubercle developed into a spine. From these lower ventro-lateral tubercles the ribs flex strongly forwards to join prominent upper ventro-lateral clavi close to the siphonal line. There are far more upper ventro-lateral clavi than lower ventro-lateral tubercles. The upper ventro-lateral clavi are separated from the serrated keel by prominent sulci across which the ribbing is very weakly developed. There are more serrations to 108 ANNALS OF THE SOUTH AFRICAN MUSEUM Z f\ eae \ f MNS “7 \ NX “f/f 7 - PE eae ~ —-=5 4) Ww // NEN of XN eas G » / ve 7 Fig. 28. Euomphaloceras (Kanabiceras) septemseriatum (Cragin). < 1. Whorl sections. A. SAM-PCA2736. B. SAM-PCA2810. C. SAM-PCA2742. x 1. the keel than upper ventro-lateral clavi. Fine ribs and growth striae may be inter- calated between main ribs. In this example tuberculation forms a subordinate part of the ornament. The greatest width is at mid-flank. In SAM-PCA2835 (Fig. 12G—H) tuberculation is again very poorly developed, whilst there are only about seven major ribs per half whorl. Between these, however, are numerous thread-like, flexuous lirae, all of which bear upper ventro-lateral clavi, but rarely lower ventro-lateral tubercles. The inner whorls of SAM-—PCA2876 are very similar to the preceding example, but on the outer whorl the fine ribbing is virtually lost and it becomes even more coarsely ribbed than SAM-—PCA2761. SAM—PCA2279 shows a further stage of development towards the effacement of ribbing. The major ribs are very distant, with only a few intercalated ribs, whilst the former have very prominent umbilical bullae, lower ventro-lateral tubercles and upper ventro-lateral clavi. The sulci separating the keel are very deep. SAM-PCA2797 (Figs 10B-—C) is an extreme individual in which virtually all ribbing is lost and tubercles dominate the ornament. Weak, prorsiradiate ribbing is evident to about 10 mm diameter, with short spines already impressed in the succeeding whorl. The ribbing, however, is far coarser at this stage than at later growth stages of many other examples, i.e. SAM-—PCA2827 (Fig. 26A-B). Beyond this growth stage ribbing 1s effaced entirely and the ornament comprises occasional, very irregularly developed, umbilical bullae, of which there are 5 on the outer whorl, equally irregular lower ventro-lateral tubercles of which there are 13 on the outer whorl, 7 of which represent the bases of long, septate spines which pointed diagonally outwards. These are preserved impressed in the penulti- UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 109 mate whorl where they attain a length of 5 mm at only 10 mm diameter. The upper ventro-lateral clavi are more distant than in the other specimens, but still far outnumber their lower counterparts. The ventral sulci are weakly developed, forming two shallow grooves on either side of the keel. The serrated keel is formed by the joining together of the asymmetrical siphonal tubercles which are steepest adapically. SAM-—PCA2826 (Fig. 18G—H) is very similar to the previous example but differs in the extreme depression of the whorl section. A probable pathological example is SAM—PCA2742 (Fig. 19K-L) which, up to a diameter of 35 mm, shows all the typical characters of this species. Immediately beyond this diameter, without any transitional stage, all ribbing and much of the tuberculation is entirely lost. The upper ventro-lateral clavi, the ventral sulci and the siphonal keel are all lost, leaving a smooth, broad, slightly convex venter. All ribbing on the flanks is lost, save fine growth lines, and the ornament comprises paired umbilical tubercles, which have now moved up to mid-flank, and ventro-lateral spines. In another specimen, SAM—PCA2738, at about the same growth stage as the previous example, all the typical Kanabiceras features are retained. SAM-—PCA2736 differs from the other specimens from these beds in having its greatest width at the umbilical shoulder. In this specimen the upper ventro-lateral clavi have become very irregular, with some representing swollen spine bases, whilst others have been suppressed to fine rib proportions. The ventral sulci are absent and the serrated keel has broken up into a row of irregular tubercles. Measurements No. D H Wi Wi/H Uo Ui SAM-—PCA2797 40 16 (40) 19 (48) 1,18 19 14 (35) y 20 9 (45) 11 (55) oe 9 8 (40) SAM-PCA2827 50 18@6) 2560) 1,39 26 18 (36) : 28 11 (39) 14 (50) ty IG 12 (43) SAM-PCA2738 46 19(41) 26 (56) 136 D5 16 (35) 40 16 (40) 18 (45) 2 ? 14 (35) SAM-PCA281 1 25 9 (36) 12 (48) 133° 0713 10 (40) SAM-PCA2835 D) 9 (41) 12 (55) 1,33 ? 7 (32) SAM-PCA2761 31 12 (39) 15 (48) [5 12 10 (32) : 16 7 (44) 9 (56) 129 0 7 (44) SAM-PCA2785 22 7 (32) 11 (50) lead ? 8 (36) - 18 6 (33) 8 (44) 1,33 8 6 (33) SAM-PCA2742 5] 17 (33) 19 (37) 1,12 ? ? . 37 14 (38) 17 (46) |, 22 ? 13 (35) SAM-PCA2756 ? 13 15 1,15 ? ? SAM-PCA2826 ? 2 17 1,42 ? ? SAM-PCA2829 18 8 (44) 11 (61) Lai ? ? 110 ANNALS OF THE SOUTH AFRICAN MUSEUM Discussion As noted by previous authors (Matsumoto 19595; Powell 1963), this is a highly variable species in which no two specimens are alike. The whorl section varies from almost quadrate to strongly depressed. Ribbing may be dominant to almost absent. Tuberculation may be very weak to extremely pronounced. The ventral sulci may be very deep to almost absent. The serrated siphonal keel 1s the most persistent character but it too may sometimes be represented by only a weakly connected row of siphonal tubercles. In a single (pathological ?) specimen it is entirely lost. Matsumoto (1959b) and Cobban & Scott (1972) have already shown that the American species Kanabiceras kanabense (Stanton) and Lyelliceras stanis- lausense Anderson do not bear separation from K. septemseriatum. The present study also confirms Prionotropis echinatus Douvillé to be a junior subjective synonym of K. septemseriatum. Kanabiceras wyomingense Reeside & Weymouth (1931: 12, pl. 1 (fig. 14)) is a crushed Neogastroplites cornutum (Whiteaves) (Reeside & Cobban 1960: 67). The Lower Turonian Kanabiceras puebloense Cobban & Scott (1972: 73, pl. 15 (figs 8-9), pl. 37 (figs 1-8), pl. 38 (fig. 1)) differs from K. septemseriatum in having more prominent uniform ribbing which, on the outer whorls at least, dominates the sculpture. It may be a Kamerunoceras. The Kanabiceras sp. of Freund & Raab (1969: 9, pl. 1 (figs 3-6)) may or may not belong to this genus. The figured specimens are preserved as internal moulds and show prominent constrictions. In view of the range of variation shown by E. (K.) septemseriatum, together with the fact that the Israeli material comes from more or less the same stratigraphical level, they might be conspecific. Alterna- tively, they might possibly belong to Euomphaloceras s.s. or even represent juveniles of Schindewolfites or Kamerunoceras. Occurrence Euomphaloceras (Kanabiceras) septemseriatum (Cragin) is a widespread species, diagnostic of uppermost Cenomanian strata, and recorded from North America (Texas, Colorado, Montana, Kansas, Mexico, Utah, Arizona, Cali- fornia), Japan, Angola, Nigeria, France and England. Subfamily Metoicoceratinae Hyatt, 1903 Genus Metoicoceras Hyatt, 1903 Type species Ammonites swallovi Shumard, 1861 Discussion Metoicoceras is a typically uppermost Cenomanian genus (Kennedy & Juignet 1973), derived from the slightly earlier Thomelites (Wright & Kennedy in Juignet et al. 1973), and giving rise to the Lower Turonian Jeanrogericeras. Jeanrogericeras in turn gave rise to the middle and upper Turonian Spathites from which was derived (according to C. W. Wright, in /itt.) the lower Coniacian UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 111 Buchiceras. From the host of species described, the majority of which have until now been regarded as endemic, there can be little doubt that species populations show a wide range of morphological variation, and that former narrowly defined species limits will break down when large collections are studied. At present, Metoicoceras is known abundantly from the Western Interior of North America (Shumard 1861; Hyatt 1903; Moreman 1927, 1942; Haas 1949; Cobban 1953; Stephenson 1955; Young 1957, 1958), from Europe (d’Orbigny 1842; Petrascheck 1902; Leriche 1905; de Grossouvre 1912; Karrenberg 1935), Israel (Avnimelech & Shoresh 1962), Nigeria (Reyment 1955), Angola (Douvillé Fig. 29. Metoicoceras geslinianum (d’Orbigny). x 1. Lateral view of the crushed ‘neotype’, preserved as a composite internal mould. x 1. i ANNALS OF THE SOUTH AFRICAN MUSEUM 1931; Cooper 1972, 1974), Morocco (Collignon 1963), Brazil (Reyment & Tait 1972a), and Madagascar (Collignon 1964). In the Montana-Wyoming area of the Western Interior, Cobban (1953) recognized four levels of Metoicoceras: M. whitei (youngest zone) M. mosbyense M. n. sp. (densely ribbed form) M. praecox (oldest zone) Of these four levels only the youngest, the level of M. whitei, corresponds to the Sciponoceras gracile Zone. It seems likely therefore, that Metoicoceras first appears at a somewhat earlier level in the Upper Cenomanian of North America than elsewhere, and that this region acted as an evolutionary pool prior to the somewhat later world-wide dispersal of the genus, a dispersal probably related to the late Cenomanian eustatic transgression (Cooper 1974). As herein interpreted, the Metoicoceratinae includes the following genera: Thomelites, Metoicoceras, Jeanrogericeras, Spathites and Buchiceras. Metoicoceras gibbosum Hyatt, 1903 Figs 21A-C, 30 Metoicoceras gibbosum Hyatt, 1903: 121, pl. 15 (figs 5-8). Moreman, 1927: 96, pl. 14 (fig. 4). Adkins, 1928: 248. Moreman, 1942: 211. Mammites gibbosum (Hyatt) de Grossouvre, 1912: 25. Buchiceras swallovi (Shumard) White, 1876: 202, pl. 20 (fig. la—c). Stanton 1893: 168, pl. 37 (fig. 1), pl. 38 (figs 1-3). Hill, 1901: pl. 40 (fig. 2). Metoicoceras whitei Hyatt, 1903: 122, pl. 13 (figs 3-5), pl. 14 (figs 1-10, 15). Grabau & Shimer, 1910: 197, text-fig. 1457c-e. Moreman, 1927: 94, pl. 15 (fig. 1). Adkins, 1928: 249, pl. 26 (figs 1-2). Moreman, 1942: 210. Shimer & Shrock, 1944: 591, pl. 245 (figs 8-10). Cobban & Scott, 1972: 74, pl. 14 (figs 3-4, 9-11), pl. 16 (figs 1-2), text-fig. 34. Metoicoceras kanabense Hyatt, 1903: pl. 15 (figs 9-11). Mammites kanabensis (Hyatt) de Grossouvre, 1912: 24. Metoicoceras acceleratum Hyatt, 1903: 127, pl. 14 (figs 11-14). Adkins, 1928: 249. Mammites acceleratum (Hyatt) de Grossouvre, 1912: 25. Metoicoceras ornatum Moreman, 1942: 211, pl. 32 (fig. 4). Metoicoceras aff. ornatum Moreman, Reyment, 1955: 47, pl. 9 (figs 6-7). Collignon 1963: 34, pl. 19 (figs 1-2). Metoicoceras aff. mosbyense Cobban, Collignon, 1963: 35, pl. 19 (fig. 3). Metoicoceras cf. whitei Hyatt, Cooper, 1972: 248; 1974: 82, 86. ? Metoicoceras aff. whitei Hyatt, Bose, 1918: 203, pl. 12 (figs 4-7). Pulchellia caicedoi Douvillé (non Karsten), 1931: 26, pl. 2 (fig. 1a—b). ? Metoicoceras besairei Collignon, 1964: 149, figs 1628-1629. Barroisiceras trinodosum Moreman, 1927: 212, pl. 33 (figs 1-2). ? Metoicoceras bései Jones, 1938: 127, pl. 10 (figs 1-3). ? Metoicoceras swallovi macrum Stephenson, 1952: 209, pl. 51 (figs 4-7). Description Only one example of this genus was collected from Salinas, and it may now be added to the ‘Pulchellia caicedoi’ described by Douvillé (1931), as the only two representatives of Metoicoceras from the Salinas beds. The specimen is a wholly septate phragmocone with the recrystallized test preserved. It is very involute, compressed, with broad convex flanks converging UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 113 towards the very narrow venter (Fig. 30). On the flanks, initially straight, later slightly flexuous, ribs arise at the umbilicus from very weak umbilical bullae and broaden markedly towards the venter. The ribbing on the inner portion of the flanks, closest to the umbilicus, is very faint and indistinct on the adoral half of the outer whorl. Between all long ribs there are almost invariably 1-2 ribs inter- calated at about mid-flank. At the venter there are about 30 ribs on the outer whorl, with rib terminations broader than the interspaces. All ribs bear strong upper and weak lower ventro-lateral clavi, the latter weakening noticeably with age. Broad low ribs connect the upper ventro-lateral tubercles across the venter on the adoral portion of the outer whorl. The narrow venter is distinctly concave on the adapical portion of the outer whorl, but as tuberculation weakens it becomes tabulate. The greatest width is at mid-flank. Measurements No. D H W H/W U SAM-—PCA2714 84 51 (61) 29 (35) 1,76 7 (8,3) . 57 28 (49) 20 (35) 1,40 ? Discussion The following ontogenetic features are apparent in the Angolan example before the writer: the shell becomes more compressed with age, the tuberculation weakens (especially the lower ventro-lateral clavi) considerably, the venter is showing the tendency, so typical amongst described forms, to round with Fig. 30. Metoicoceras gibbosum Hyatt, SAM—PCA2714. Whorl section. x 1. 114 ANNALS OF THE SOUTH AFRICAN MUSEUM maturity, and flank ribbing weakens and becomes flexuous. A score of ‘species’ have been described from western Europe and the Western Interior of North America. Casts of much of the North American type material, as well as photographs of the European types, kindly shown to the writer by W. J. Kennedy, leave no doubt whatsoever that the species of this genus are highly variable, and that the majority of described forms are merely nominal species. Metoicoceras whitei Hyatt (1903: 122, pl. 13 (figs 3-5), pl. 14 (figs 1-10, 15)) was erected for one of Shumard’s (1861) syntypes of Ammonites swallovi (Stephenson 1952), the types of which are now lost. Hyatt’s (1903, pl. 13 (figs 3—-5)) paratype still remains (USNM 29408) and shows the following features: shell compressed, involute on the inner whorls, with weak, but distinct, umbilical bullae giving rise to prorsiradiate ribs which broaden ventrally. There are about 9 such umbilical bullae (and main ribs) at a diameter of about 85 mm. Between main ribs there are generally 2 intercalatories. All ribs are ornamented with upper and lower ventro-lateral tubercles. On the body chamber, tubercula- tion weakens considerably, the venter rounds, the umbilical bullae are lost, and ribbing coarsens so that there is only a single intercalatory between main ribs. There are 30 ribs on the outer whorl. Metoicoceras gibbosum Hyatt (1903: 121, pl. 15 (figs 5—8)) was founded ona single specimen differing from M. whitei only in being slightly more inflated, apparently lacking umbilical bullae and in having fewer (25) ribs on the outer whorl. Cobban & Scott (1972) regarded M. gibbosum and M. whitei as con- specific, placing the former into the synonymy of the latter. The writer concurs with these authors in regarding M. whitei and M. gibbosum as synonyms. How- ever, contrary to Cobban & Scott (1972), the name gibbosum has page priority over whitei, and is thus the valid name. Metoicoceras kanabense Hyatt (1903, pl. 15 (figs 9-11)) was figured but not described. It is a juvenile, strongly compressed and very involute, with about 28 prorsiradiate ribs on the outer whorl. Umbilical bullae are very weak or absent. All ribs are ornamented with upper ventro-lateral clavi, although lower ventro- lateral tuberculation appears to be absent. This species comes from the same horizon as M. gibbosum and is probably a synonym. Metoicoceras acceleratum Hyatt (1903: 127, pl. 14 (figs 11-14)) is another species based on a juvenile. It is not, at present, directly comparable with the other species but, so far as can be judged from the photograph, is probably not distinct from M. gibbosum. Stephenson (1952) has discussed Metoicoceras swallovi (Shumard); Shumard (1861) failed to illustrate his material, which is now lost. Stephenson (1952: 208) considered M. swallovi to show the following features: shell compressed, relatively evolute, with a moderately deep umbilicus; umbilical seam egresses markedly on the body chamber; outer whorl with 28 ribs, fewer on inner whorls, which are slightly prorsiradiate on the inner whorls but recurve on the body chamber so as to become convex adorally; at a diameter of about 60 mm there are 7 promi- UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA pis nent swollen umbilical tubercles per whorl which on the final whorl become more numerous and bullate; each tubercle gives rise to a single main rib, with 1-2 intercalatories on the inner whorls. Ribbing weakens towards the aperture; all ribs are ornamented with upper and lower ventro-lateral clavi, although the latter become obsolete on the body chamber; the narrow venter remains tabulate on the body chamber, while the upper ventro-lateral clavi remain strong through- out ontogeny. Metoicoceras swallovi differs from M. gibbosum in being relatively more evolute, with more prominent, rounded umbilical tubercles, and in that the ribbing recurves on the body chamber. The strength of the umbilical tubercles has long been regarded as a diagnostic characteristic of many Metoicoceras species. So far as the writer 1s able to judge, this character is highly variable and its value in taxonomy has been overestimated. Metoicoceras swallovi macrum Stephenson (1952: 209, pl. 51 (figs 4-7)) was said to differ from M. swallovi in being more compressed and with bullate umbilical tubercles in the early growth stages. This subspecies would seem merely to point to transitional forms between M. gibbosum and M. swallovi. Metoicoceras mosbyense Cobban (1953: 48, pl. 6 (figs 1-14), pl. 7 (figs 1-3)) is a large species (? macroconch), very involute on the inner whorls but becoming rather evolute at large diameters. The inner whorls show prominent umbilical bullae, about 10 per whorl, giving rise to slightly prorsiradiate ribs, between which are |—2 intercalatories. At the venter, at about 40 mm diameter, there are about 30 ribs per whorl. All ribs are ornamented with upper and lower ventro- lateral tubercles up to a diameter of 45 mm when the lower ventro-lateral tubercles become obsolete. At about 110 mm diameter the upper ventro-lateral clavi are lost and the venter becomes rounded. Also at this stage the ribs coarsen and become more robust, numbering only about 24 per whorl. This species was distinguished from M. whitei (= M. gibbosum) in its loss of lower ventro-lateral tubercles at small diameters. Metoicoceras muelleri Cobban (1953: 49, pl. 6 (figs 15-16), pl. 8 (figs 1-7), pl. 9) is a compressed species with about 32 sigmoidal ribs per whorl at small growth stages, but ribbing coarsens on the outer whorl when there may only be 20. The lower ventro-lateral tubercles are lost at only 11 mm diameter. Cobban (1953) noted transitions between this species and the contemporaneous M. mos- byense, suggesting that the differences are not specific. The large size attained by M. mosbyense suggests that the differences might be sexual. Metoicoceras defordi Young (1957: 1169, pl. 149 (figs 1-8), text-fig. la, e, g, 1) is a compressed species, very involute on the inner whorls but becoming rather evolute on the outer whorls as the umbilical seam egresses. There are about 30 prorsiradiate ribs per whorl at 70 mm diameter, of which about one- quarter are long ribs ornamented with prominent umbilical bullae. There are generally 2 intercalatories between main ribs. The venter remains concave until about 100 mm diameter, after which it becomes tabulate, finally becoming rounded at diameters of about 150 mm. According to Young (1957: 1171) 116 ANNALS OF THE SOUTH AFRICAN MUSEUM ‘Metoicoceras defordi, n. sp., in whorl section and ornamentation appears to be related to M. whitei Hyatt, but the species is older than M. whitei, and the earlier disappearance of one pair of shoulder nodes and especially the flat topped ribs in the penultimate whorls and on the body whorls serve to separate the two species. The flat-topped ribs would appear to be a specific specialization not transmitted to any younger known species of the genus.’ Metoicoceras irwini Moreman (1927: 92, pl. 13 (figs 3—4)) is a compressed, very involute species with about 32 prorsiradiate ribs, broadening ventrally on the outer whorl, half of which is body chamber. Prior to the start of the body chamber all ribs are ornamented with weak lower ventro-lateral tubercles and prominent upper ventro-lateral clavi. On the adapical half of the outer whorl the venter is tabulate, presumably becoming rounded on the body chamber. The inner parts of the flanks are almost smooth and appear to be lacking in ornament. On the early parts of the body chamber the ribs are distinctly convex towards the aperture, but they straighten out adorally. This species is almost identical to M. dumasi (Pervinquiére) which is probably not separable from M. geslinianum. Metoicoceras boesei Jones (1938: 127, pl. 10 (figs 1-3)) is based on a poorly preserved holotype which shows the following features: shell compressed, rela- tively evolute. Fairly prominent umbilical bullae gives rise to one, rarely two, prorsiradiate main ribs which broaden ventrally. There are 2-3 intercalatories between main ribs. All ribs are ornamented with weak upper and lower ventro- lateral clavi. Ribs pass weakly across the narrow, tabulate venter. There were probably about 28 ribs on the outer whorl. This species differs from M. gibbosum and the Angolan example only in being more evolute. Metoicoceras ornatum Moreman (1942: 211, pl. 32 (fig. 4)) is a compressed species, involute on the inner whorls but becoming rather evolute as the umbilical seam egresses. Long ribs arise from rather prominent umbilical bullae and are separated by 1-2 intercalatories. There are 21 ribs on the outer whorl, all of which broaden ventrally and are ornamented with prominent upper and lower ventro-lateral clavi to large growth stages. In the paratype, the ribbing becomes very coarse on the body chamber and forms prominent folds across the venter. This species occurs side by side with M. whitei (Moreman 1927: 93) (= M. gib- bosum) and the writer does not regard the differences as of specific status. ‘Barroisiceras’ trinodosum Moreman (1927: 212, pl. 33 (figs 1—2)) is con- sidered by Cobban & Scott (1972: 63) to be a pathological example of M. whitei. ‘Barroisiceras’ brittonense Moreman (1927: 212, pl. 33 (fig. 3)) is possibly also a pathological Metoicoceras. Metoicoceras latoventer Stephenson (1952: 209, pl. 51 (figs 4-7)) is an involute, compressed form in which the umbilical seam egresses markedly on the outer whorl. The final whorl bears 19 slightly sinuous, prorsiradiate ribs generally alternating long and short. Long ribs (13 to 15 per whorl) bear prominent conical umbilical tubercles up to 25 mm diameter, whereafter they become bullate. On the inner whorls all ribs are ornamented with lower ventro-lateral tubercles and upper ventro-lateral clavi, which become obsolete on the body chamber. Up to UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 117 about 25 mm diameter the juvenile shell shows a weak row of siphonal clavi. This species differs from M. gibbosum in being more evolute, with more numerous umbilical bullae and coarser, more distant flank ribbing. Metoicoceras crassicostae Stephenson (1952: 210, pl. 58 (figs 6-8)) was based on a single specimen showing the following features: shell moderately inflated and rather evolute, with umbilical seam egressing markedly on the outer whorl; there are about 11 long ribs on the outer whorl which arise from prominent umbilical bullae; there are sporadic intercalatories between long ribs, with about 18 ribs on the outer whorl; the venter is ‘considerably worn’, but shows signs of upper and lower ventro-lateral tubercles. Stephenson (1952: 210) distinguished M. crassicostae from M. latoventer in that ‘the venter of this species is broader and not quite so flat at a corresponding stage of growth, the ribs are fewer, the nodes on the umbilical shoulder are much stronger, and the nodes on and near the ventral angles are weaker’. So far as can be judged, M. crassicostae differs from M. latoventer in having fewer, more robust umbilical bullae on the inner whorls. It is thus close to M. swallovi, differing only in its more distant ribbing. The North American species of Metoicoceras are at present difficult to evaluate, since they occur at different stratigraphic horizons and the population structures appear to differ with time. Thus while the inner whorls of M. defordi are very similar to M. whitei, Cobban & Scott (1972) consider Young’s slightly older species to be distinctive. Metoicoceras whitei is itself very close to M. ges- linianum, in fact it is doubtful whether the juvenile whorls can be distinguished. Until the American populations are objectively revised and compared with European material the writer prefers to refer the Angolan material to Metoico- ceras gibbosum. ‘Metoicoceras’ antiquum (Karrenberg 1935: 139, pl. 31 (fig. 13)) does not appear to belong to this genus nor indeed the Metoicoceratinae. It is a Lower Cenomanian species better referred to the Mantelliceratinae. Metoicoceras bethlehemense (Avnimelech & Shoresh) (1962: 533, pl. 15 (fig. 3)) is a strongly compressed (crushed ?) form with prorsiradiate ribs through- out ontogeny. Main ribs arise singly or in pairs from fairly prominent umbilical bullae, with 1—2 intercalatories between long ribs. On the inner whorls the flank ribs are distinctly flexuous. Upper ventro-lateral clavi are retained to maturity, although the lower ventro-lateral tubercles are lost on the body chamber. This species is doubtfully separable from M. gibbosum. Ammonites geslinianus d’Orbigny (1850: 146) (Fig. 29) was proposed for Ammonites catillus d’Orbigny (non Sowerby) (1840: 325, pl. 97 (figs 1-2)) from the uppermost Cenomanian of Lamnay, Sarthe. D’Orbigny’s synthetograph, highly idealized as was frequently the case, shows a very compressed form, very involute (although the umbilical seam egresses markedly on the final whorl), and with flexuous ribs (about 30 per whorl) broadening rapidly towards the venter. Ribs arise singly or in pairs from fairly distinct umbilical bullae, with 1-2 inter- calatories between main ribs. On the final whorl all ribs are ornamented with 118 ANNALS OF THE SOUTH AFRICAN MUSEUM upper ventro-lateral] clavi only, the lower ventro-lateral tubercles having become obsolete at an early growth stage. Sornay (1955, fiche 11, figs 3-4) refigured d’Orbigny’s example, showing the ribbing to become very slightly convex adorally on the body chamber, at which stage, also, the venter becomes rounded. This adorally convex ribbing is well shown by Metoicoceras pontieri Leriche (1905: 120, pl. 2) which Wright & Kennedy (in Juignet et a/. 1973) consider a junior subjective synonym of M. geslinianum. Metoicoceras geslinianum (d’Orbigny) appears to differ from M. gibbosum and the Angolan example, only in losing its lower ventro-lateral tubercles at small sizes, and in the recurvature of the flanks ribs on the body chamber. Wright & Kennedy (in Juignet et al. 1973) considered Metoicoceras petraschecki (de Grossouvre) (1912: 22, pl. 2 (fig. 2)), M. bureaui (de Grossouvre) (1912: 22, pl. 1 (fig. 2)) and M. dumasi (de Grossouvre) (1912: 23, pl. 2 (fig. 1)) as probable synonyms of M. geslinianum. Metoicoceras gourdoni (de Grossouvre) (1912: 20, pl. 1 (fig. 1)) is a relatively evolute species with 6 prominent umbilical bullae on the penultimate whorl, from which arise coarse, robust, almost rectiradiate ribs which broaden ventrally. On the adoral portion of the outer whorl the ribbing becomes convex adorally. There is a single intercalatory between main ribs on the outer whorl, with 21 ribs per whorl. Unless there are two or more intercalatories between main ribs on the inner whorls, which seems likely, juveniles are equally distantly ribbed. At the largest growth size, which includes about a half-whorl of body chamber, there are prominent upper and lower ventro-lateral tubercles. The most distinctive features of MM. gourdoni are, therefore, its relatively evolute form, distant, robust ribbing and the persistence of the ventro-lateral tubercles to large diameters. These features serve to distinguish the holotype from M. geslinianum. Metoicoceras stoliczkai Sastry & Matsumoto (1967: 2, pl. 1 (fig. 1)) from the topmost beds of the Utatur Group of southern India differs from all other Metoicoceras species in having a deep, crater-like umbilicus, with steep umbilical walls. This feature, together with the suture-line, suggests it is better referred to the Lower Turonian Jeanrogericeras. Indeed, it would appear to be conspecific with J. binicostatum (Petrascheck) (cf. Wiedmann 1964: 126, figs 10-11). Collignon (1964) described a number of species of Metoicoceras, supposedly from the ‘Lower’ Cenomanian of Madagascar. Almost all the typical Upper Cenomanian species recorded from Madagascar are reported from this “Zone a Mantelliceras mantelli et Calycoceras newboldi’, and leave little doubt that the faunas of this zone represent collecting from different palaeontological horizons. However, it is difficult to ascertain whether they represent true Metoicoceras or are homoeomorphous mantelliceratinids allied to Utaturiceras and Graysonites. Metoicoceras swalloviforme Collignon (1964: 149, fig. 1627) is a moderately evolute species with about 11 fairly prominent umbilical bullae on the outer whorl, from which arise slightly prorsiradiate ribs. Between long ribs there is generally an intercalatory, so that there are 22 ribs on the outer whorl. All ribs are ornamented with small, pointed lower ventro-lateral tubercles and prominent UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 119 upper ventro-lateral clavi. This species was said to differ from M. swallovi in whorl section and details of the suture-line. As noted by Hyatt (1903), however, the suture-line is highly variable. The main difference from M. swallovi appears to be in the more robust, distant ribbing of M. swalloviforme, and in being some- what more inflated. It is thus close to M. crassicostae Stephenson. Metoicoceras besairiei Collignon (1964: 149, figs 1628-1629) is a compressed species with flat flanks, a very narrow umbilicus and a concave venter. Ornament comprises 10-12 long ribs, with weak umbilical bullae, between which are 1-2 intercalatories. The ribs are prorsiradiate, and number about 17 ribs per half whorl. All ribs are ornamented with lower ventro-lateral tubercles and upper ventro-lateral clavi. Metoicoceras besairiei was said to differ from M. whitei in having more flexous ribbing, maximum width near the umbilicus and in differ- ences in the suture-line. The differences are slight. Metoicoceras sakarahense Collignon (1964: 150, figs 1630-1632) shows the following features: the ribbing is very fine; main ribs are slightly more prominent than the intercalatories, and are slightly flexous, prorsiradiate, with weak umbilical bullae; between 2 and 4 intercalatories between main ribs; all ribs are ornamented with upper and lower ventro-lateral tubercles; on the adoral portion of the outer whorl, the ribbing forms a chevron across the venter; there are at least 42 ribs per whorl. Metoicoceras fasciculatum Collignon (1964: 151, fig. 1633) was charac- terized by the sinuous, fasciculate nature of the ribbing. It is a small species, very involute, with flat, parallel flanks. There are about 23 very sinuous main ribs which arise at the umbilicus. Of these, some are simple while others give rise, at various levels on the flanks, to 2—3 secondaries, so that there are 40 ribs at the venter of the outer whorl. Umbilical tubercles appear to be lacking, although some of the ribs may be weakly flared near the umbilicus. All ribs are ornamented with upper and lower ventro-lateral tubercles. Ribbing forms a weak chevron across the venter. Thus, of 27 described ‘species’ of Metoicoceras, two (M. antiquum Karren- berg and M. stoliczkai Sastry & Matsumoto) do not belong to this genus, while M. gibbosum Hyatt includes amongst its synonyms M. whitei Hyatt, M. kana- bense Hyatt, M. acceleratum Hyatt, M. ornatum Moreman and possibly also M. boesei Jones, M. bethlehemense (Avnimelech & Shoresh) and M. besairiei Collignon. Metoicoceras muelleri Cobban and M. defordi Young are possible synonyms of M. mosbyense, while probable synonyms of M. geslinianum include M. pontieri Leriche, M. bureaui (de Grossouvre), M. petraschecki (de Gros- souvre), M. dumasi (de Grossouvre), M. gourdoni (de Grossouvre) and M. irwini Moreman. Metoicoceras crassicostae Stephenson and M. latoventer Stephenson are doubtfully separable contemporaneous species, while M. swallovi is not well . known and its affinities are unclear. The difficulties associated with the Madagas- can species have already been noted; M. swalloviforme appears to belong to the swallovi-crassicostae group, while M. sakarahense Collignon and M. fasciculatum Collignon are doubtfully separable. 120 ANNALS OF THE SOUTH AFRICAN MUSEUM Occurrence Metoicoceras gibbosum is recorded from the uppermost Cenomanian of Texas, Mexico, the Western Interior, Morocco, Nigeria and Angola, and is possibly also present in the Middle East and Madagascar. Subfamily Mammitinae Hyatt, 1900 Genus Watinoceras Warren, 1930 Type species Watinoceras reesidei Warren, 1930 (= Acanthoceras amudariense Arkhangel’skii, 1916) 1930 Watinoceras Warren 1954 Benueites Reyment Discussion As diagnosed by Wright (in Arkell et al. 1957: L416), Watinoceras shows the following features: ‘Early whorls compressed, finely ribbed, with inner and outer ventro-lateral and siphonal tubercles as in Neocardioceras, but siphonal row soon lost; later, venter may be concave between rows of ventro-lateral clavi or rounded, with ribs passing over in chevrons; ornament usually coarsens with age.’ Wright’s (in Arkell et al. 1957) diagnosis was based upon ‘Acanthoceras’ amudariense Arkhangel’skii (1916: 48, pl. 7 (figs 8-13)) and undescribed forms from Devonshire said to be transitional from Neocardioceras to Watinoceras (cf. Reyment 1955: 55). Siphonal tubercles are not known in the holotype of W. reesidei, nor have they ever been recorded from undoubted species of this genus. Consequently, contrary to Reyment (1955), any forms possessing siphonal tubercles are best retained in the genus Neocardioceras, and Watinoceras is restricted for those species with a quadrituberculate venter at some or all growth Stages. On the basis of the North American material, Cobban & Scott (1972: 75) gave the following, somewhat contrasting diagnosis of Watinoceras: “This is a moderately evolute genus that has somewhat compressed whorls, narrow but conspicuous ribs, and umbilical, lower ventro-lateral, and upper ventro-lateral tubercles of which the upper ventro-lateral ones are the strongest. The siphonal area is narrow, somewhat flattened, and bordered by the high closely spaced upper ventro-lateral tubercles. Ribs are prorsiradiate and, on the inner whorls, sigmoidal. The suture is rather simple and has a wide slightly incised first lateral saddle and much narrower lateral lobe.’ This conflicting diagnosis is due to the fact that Cobban & Scott (1972) based their diagnosis of Watinoceras on the apparently immature W. reesidei Warren and the adult W. coloradoense (Henderson), neither of which shows the change in ornament exhibited by W. amudariense (Arkhangel’skii). UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 121 Callomon (1963), in his important work on sexual dimorphism in ammonites, considered three features to be diagnostic of the attainment of maturity by an ammonite. These were: (i) A change in the style of ornament. (ii) Uncoiling of the umbilical seam. (iii) Sutural approximation. The inner whorls of Watinoceras amudariense (Arkhangel’skii) are orna- mented with upper and lower ventro-lateral tubercles, between which the venter appears flattish. On the adoral half of the outer whorl, however, the ventro- lateral tuberculation is lost, the venter becomes rounded, and the ribs form prominent chevrons across the venter. Moreover, between 15 and 21 mm dia- meter W. amudariense has an umbilical ratio of 38,8—-39,5 per cent, whereas at the adoral end of the outer whorl the umbilical ratio is 41,5 per cent. Arkhangel’skii’s (1916) material would seem to retain recrystallized test, and hence the nature of the suture-line is unknown. The fact that Watinoceras amudariense shows two of the features considered diagnostic of maturity (Callomon 1963) is significant, and the writer regards W. amudariense as mature. A number of authors (Reyment 1955; Collignon 1963; Cobban & Scott 1972) have regarded ‘Acanthoceras’ amudariense var. horridum Arkhangel’skii (1916: 49, pl. 8 (figs 8-10, 14-15)) as conspecific with Watinoceras coloradoense (Henderson), an assignment with which the writer concurs. The latter species, however, attains diameters of 110 mm (Cobban & Scott 1972), and rather than lose tuberculation in maturity, the ventro-lateral tubercles become very pro- nounced, while ribbing coarsens. Cobban & Scott (1972) noted, however, that an impression of the inner whorls of the holotype of W. coloradoense (Fig. 27) ‘shows strong narrow rectiradiate ribs numbering 15 per half whorl at a diameter of approximately 24 mm’. The inner whorls thus bear a close resemblance to both W. amudariense and Watinoceras reesidei Warren. It is thus significant that Cobban & Scott (1972) record W. reesidei from the same horizon as W. coloradoense. Cobban & Scott (1972: 76) considered Warren’s (1930, 1947) type and topo- type material of Watinoceras reesidei to show the following features: “(The shells) are moderately evolute, and their umbilical ratio is about 40%. The whorls, which are higher than wide, are ornamented with numerous narrow slightly signoidal prorsiradiate ribs of which about every other one extends to the umbilicus. Each rib bears a small sharp lower ventro-lateral tubercle and a slightly larger upper ventro-lateral tubercle. Most of the ribs that extend to the umbilicus have a low bullate umbilical tubercle. The last half whorl of the holotype has 22 ribs.’ These are the exact features which characterize Watinoceras amudariense (Arkhangel’skii), except that in the latter species tuberculation is lost at diameters of the order of 25 mm, whereafter the ribs form chevrons across the venter. Since none of Warren’s (1930, 1947) material is over 20 mm in diameter the differences 122 ANNALS OF THE SOUTH AFRICAN MUSEUM are not considered significant, and Watinoceras reesidei Warren is considered a junior subjective synonym of W. amudariense (Arkhangel’skii). The interesting situation is now arrived at that the small Watinoceras amudariense occurs side by side with the large Watinoceras coloradoense from areas as far afield as Alaska (Cobban & Gryc 1961), Alberta (Warren 1930), Colorado (Cobban & Scott 1972), Turkestan (Arkhangel’skii 1916) and possibly Morocco (Collignon 1966). As has already been noted, the inner whorls of W. coloradoense are finely ribbed and thus close to W. amudariense. Indeed, the rib density of W. reesidei and W. amudariense falls within the range of variation observed for juvenile W. coloradoense (Fig. 29), and it seems likely that at this growth stage the two species cannot be separated. The two species W. amudariense (Arkhangel’skii) and W. coloradoense (Henderson) thus show all the prerequisites of sexual dimorphism, viz. the inner whorls appear to be indistinguishable; they attain maturity at greatly differing diameters; the microconch shows a marked change of ornament on the body chamber and they occur side by side, and con- sequently are herein interpreted as microconch and macroconch respectively. The oldest available name is W. coloradoense (Henderson). The genus Benueites Reyment is very close to Watinoceras, the main points of difference being that the former has a sulcate venter, finer ribbing and less regular tuberculation. The differences are merely ones of degrees, and are herein considered as of only subgeneric status. Of interest is the fact that Reyment (19715) has recently recorded dimorphism within Benueites, dimorphs being separated on the basis that ‘one of the pair is ornamented with dense ribs which are feebly adorned with tubercles, while the other member has much coarser rib- bing, and usually three rows of tubercles, namely, umbilical, upper and lower ventro-lateral, the former being mostly more numerous and stronger than the umbilical row. The ventral furrow of the first group is deep and narrow and seldom cut by ribs, whereas that of the second category is broad and shallow and frequently crossed by ribs.’ Whilst Reyment (19715) considered this to be a novel type of dimorphism, it appears that the differences are largely due to a com- parison of microconch dimorphs with the middle whorls of the corresponding macroconch. The genus may now be divided as follows: W. (Watinoceras). Shell evolute, with prominent ribbing bearing umbilical and upper and lower ventro-lateral tubercles. Dimorphic; microconchs small with ventro-lateral tubercles lost on the body chamber, when the ribs form a chevron across the venter. Macroconch relatively large, tubercles and ribbing becoming coarser and more prominent with age. Age: Basal Lower Turonian. W. (Benueites). Similar to W. (Watinoceras), but microconch with very reduced tuberculation and a prominent siphonal sulcus across which ribbing is effaced. Macroconch with coarser ribbing and tuberculation and a weakly developed ventral sinus across which ribs are weakly con- nected. Age: Upper Lower Turonian. UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 123 Watinoceras (Watinoceras) coloradoense (Henderson, 1908) (9) Figs 18C—D, 19E-F, 31-34 Microconch (3) Acanthoceras amudariense Arkhangel’skii, 1916: 48, pl. 7 (figs 8-13). Watinoceras reesidei Warren, 1930: 67, pl. 3 (fig. 2), pl. 4 (figs 9-12). Cobban & Gryc, 1961: 186, pl. 38 (figs 44-49). Hattin, 1965: text-fig. 3. Cobban & Scott, 1972: 75, pl. 27 (figs 7-10), pl. 28 (fig. 4). ? Watinoceras nov. sp. aff. reesidei Warren, Collignon, 1966: 37, pl. 19 (figs 14-15). Sumitomoceras amudariense (Arkhangel’skii) Matsumoto et al., 1969: 282. Macroconch (@) Acanthoceras coloradoense Henderson 1908: 259, pl. 13, figs 10-11. Acanthoceras amudariense var. horridum Arkhangel’skii, 1916: 49, pl. 8 (figs 8-10). Gauthiericeras aff. bravaisi (d’Orbigny) Moreman, 1927: 96, pl. 14 (fig. 2). Watinoceras coloradoense: (Henderson) Reyment, 1955: 57; Collignon 1963: 37. Cobban & Scott, 1972: 76, pl. 27 (figs 11-19), pl. 28 (figs 1-3, 5-9). ? Watinoceras nov. sp. aff. coloradoense (Henderson) Collignon, 1966: 37, pl. 20 (figs 1-3). Watinoceras horridum (Arkhangel’skii) Matsumoto et al., 1969: 282. Description Two specimens, one beautifully preserved, the other rather crushed, appear to belong here. Both are entirely septate and have the recrystallized test preserved. SAM-PCA2730 (Fig. 18C—D) is rather evolute, with about three-quarters of the penultimate whorl visible, and has a compressed whorl section, polygonal costally but rounded intercostally (Fig. 32). The flanks are slightly convex with maximum width at mid-flank. The umbilicus is moderately wide and rather deep, with a steep umbilical wall, slightly undercut at the umbilical seam. The umbilical shoulder is abruptly rounded. At a diameter of 48 mm there are 41 ribs per whorl, with only 19 arising from bullae on the umbilical shoulder. These bullae are rather prominent to a diameter of 38 mm, after which they weaken considerably. Between each main rib there is invariably a shorter intercalated rib, occasionally two. Ribbing is strong and sharp, prorsiradiate across the flanks. Up to 25 mm diameter the ribs are rather rigid and straight, beyond which they become flexuous and convex forwards. At 38 mm diameter, when the umbilical bullae become obsolete, the main ribs are seen to continue across the umbilical wall to the umbilical seam. Ribs are narrowest near the umbilicus and broaden ventrally. At the lower ventro- lateral clavi the ribs bend forwards to the upper ventro-lateral clavi, crossing the venter with a slight adoral convexity. Siphonal tubercles are lacking at all visible growth stages. The ventro-lateral tubercles only become clavate at about the Same stage as the umbilical tuberculation is lost (38 mm diameter). Prior to this, prominent bubble-like lower ventro-lateral tubercles occur only on the main ribs, while similar bubble-like upper ventro-lateral tubercles occur on all ribs. Up to 38 mm diameter the upper ventro-lateral tubercles are distinctly higher than the ribbing across the venter, so that the venter has a concave aspect. Beyond this 124 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 31. Watinoceras coloradoense (Henderson). Lateral and ventral views of the holotype. Note fine ribbing of the inner whorls on the external mould. Photo: W. J. Kennedy. x 1. Fig. 32. Watinoceras coloradoense (Henderson) (9). Whorl sections of SAM—PCA2730. x 1. diameter, the upper ventro-lateral clavi are no higher than the ribbing, giving the venter a tabulate appearance. The suture-line (Fig. 33) closely resembles that of Watinoceras coloradoense (Henderson) as figured by Cobban & Scott (1972). The shell of SAM-—PCA2753 (Fig. 19E-F) is rather evolute, with a moderately wide umbilicus. The umbilical wall is steep and slightly undercut at the umbilical seam. The umbilical shoulder is well rounded with broad, flattish flanks. UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 125 Ornament comprises rather weakly developed umbilical bullae from which arise slightly prorsiradiate main ribs, separated by 1-2 intercalatories. There are 18 ribs per half whorl, generally with upper and lower ventro-lateral clavi. The Fig. 33. Watinoceras coloradoense (Henderson) (2). Suture-line of SAM—PCA2730 at about 48 mm diameter. upper ventro-lateral clavi are very close together and stand out above the ribbing, giving the venter a concave aspect. There is no sign of siphonal tubercles at any stage. Measurements No. D H W W/H U SAM-PCA2730 48 21 (44) 17 (35) 0,81 ls) I) . 35) 16 (46) 12 (34) 0,75 9 (26) SAM-—PCA2753 4l 17 (41) 13 (Gv) 0,76 14 (34) Discussion As shown above, Watinoceras coloradoense (Henderson), W. reesidei Warren and W. amudariense (Arkhangel’skii) are conspecific, the oldest available name being W. coloradoense. There appears to be a wide range of variation, both ontogenetic and intra- specific, within Henderson’s species. The variation in rib density within the macroconch is well shown by Cobban & Scott (1972) (see also Fig. 34), while these authors record 1-3 intercalatories between main ribs in this species. Arkhangel’skii’s (1916, pl. 8 (figs 8, 14)) juvenile macroconchs, however, show only simple, main ribs. Moreover, the diameter at which the coarse macroconch ornament first appears is also variable. Thus, typical ‘Watinoceras coloradoense- type’ ornament appears at only 25 mm diameter in ‘Acanthoceras’ amudariense var. horridum, whereas the Angolan example still retains ‘Watinoceras reesidei- type’ ornament at 48 mm diameter. A similar amount of variation would also 126 ANNALS OF THE SOUTH AFRICAN MUSEUM seem to occur in the first appearance of adult ornament in the microconch. Thus, ‘Acanthoceras’ amudariense is adult at 30 mm diameter, whereas Cobban & Scott (1972) note that ‘impressions of Watinoceras reesidei from north-western Montana (Cobban, 1956a, p. 1003-1004) suggest that the adults attain diameters of 70-80 mm and that the adult whorls are about as densely ribbed as the juvenile whorls’. It seems likely, however, that some of the specimens interpreted by Cobban & Scott (1972) as juvenile W. coloradoense are in fact microconchs and, conversely, some of those interpreted as adult W. reesidei are merely densely ribbed, immature variants of the macroconch. Consequently, previous records of these species now need reinvestigation under this new light. Matsumoto (in Matsumoto et al. 1969) assigned ‘Acanthoceras’ amudariense to Sumitomoceras, a genus to which it most certainly does not belong. None the less, it is significant that the Angolan example closely resembles certain species of Tarrantoceras (= Sumitomoceras), e.g. T. rotatile Stephenson (Fig. 20). However, in Tarrantoceras, a siphonal row of tubercles persists to large diameters, and is only lost on the final whorl, while siphonal tubercles are unrecorded from undoubted Watinoceras. Family Vascoceratidae Spath, 1925 Discussion Wright (in Arkell et al. 1957: L418) gave the features of this family as: ‘Derivatives of Acanthoceratinae which rapidly lose ribbing and tuberculation of that subfamily and are either smooth or bluntly tuberculate or have sparse coarse ribs. Suture may comprise shallow, irregular and slightly indented or deep and much indented elements. The whorl section and degree of involution is variable, even within species. Typically Tethyan in occurrence.’ Within the family Vascoceratidae, Wright (in Arkell et al. 1957) included the following genera: Nigericeras, Spathites, Gombeoceras, Ezilloella, Paravasco- ceras, Pachyvascoceras, Vascoceras, Paramammites, Plesiovascoceras, Fagesia, Thomasites and Neoptychites. Wiedmann (1959) subdivided the family into two subfamilies, the Vasco- ceratinae and the Fallotitinae, based on the presence or absence of siphonal tubercles. The subfamily Fallotitinae, which lacks siphonal tubercles, was thought to be derived from the Mammitinae, while the Vascoceratinae, with siphonal tubercles, was considered to have evolved from the Acanthoceratinae. Within the Fallotitinae, Wiedmann (1964: 127) placed the genera Plesiovasco- ceras, Fallotites, Paramammites, Ingridella, Spathitoides, Neoptychites and doubtfully Metasigaloceras. This leaves Nigericeras, Spathites, Gombeoceras, Ezilloella, Paravascoceras, Pachyvascoceras, Vascoceras, Fagesia and Thomasites within the Vascoceratinae. Of all the species of Vascoceras s.s. described by Choffat (1898), Pervin- quiere (1907), Reyment (19545, 1955), Barber (1957), and Freund & Raab (1969), no mention is made of siphonal tubercles at any growth stage. Moreover, through the kindness of P.-Y. Berthou, the writer obtained excellent photographs 127 UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA (CLOT) HOSS 2 URQgoD Joye Jed Url Byeq *(Sso1d) o[duiexo uejosuy sy} pue ‘(S}Op) (UOSIopudH) asuaopos0jos “yy “(so[>II9) (IS, JosueYyyIY) Wnplisoy “Ted asualappniup - fy ‘(sorenbs) (Lys,JasueyyIV) asuatuppnuup -yy ‘(So[SUPII}) UdLIe AA Japisaas SpsavoulD YY JO SUBUUISOdS IOJ JoJOWILIP ysuIeSe AjISUOP QII JO 10]g “PE ‘SIq sajoweig Www O21 ole Ol Jsoya yey /squs ‘oN 128 ANNALS OF THE SOUTH AFRICAN MUSEUM of all of Choffat’s (1898) specimens, none of which shows siphonal tubercles. This is confirmed by an examination of the type material (P.-Y. Berthou, pers. comm.) and would appear to invalidate Wiedmann’s subdivision of the family Vascoceratidae. Genus Vascoceras Choffat, 1898 Type species Vascoceras gamai Choffat, 1898 The genus Vascoceras Choffat (type species Vascoceras gamai Choffat, 1898: 54, pl. 7, pl. 8 (fig. 1), pl. 10 (fig. 2), pl. 21) was diagnosed by Barber (1957: 13) as follows: ‘Shell discoidal, or subglobose; whorls normally largely embracing. Venter sub-acute, rounded or flattened. Umbilicus steep sided. Early whorls ornamented with strong umbilical tubercles and fold-like costae which cross the venter. With age the ornament is lost and the last whorl is normally smooth. Suture line: three incised broad saddles and three bifid or denticulate lobes.’ The genus Paravascoceras Furon (type species Vascoceras cauvini Chudeau, 1909: 67, pls 1-3) was considered by Freund & Raab (1969) to include the synonyms Pachyvascoceras (Furon 1935: 58), Paracanthoceras (Furon 1935: 59), Broggiiceras (Benavides-Cacéres 1956: 409) and Discovascoceras (Collignon 1957: 11). These authors diagnosed the genus as follows: “Vascoceratids lacking umbilical tubercles. Compressed and involute at early stages; less involute later. During growth, there is either a short or a very long stage without any ornament. Lateral or marginal bulges sometimes appear; prorsiradiate ribs, uninterrupted crossing the venter, usually appear at a late stage. They rarely attain the umbilical border. The suture is simple and typical for the family.’ Freund & Raab (1969: 20) considered the most diagnostic feature of Para- vascoceras to be the absence of umbilical tubercles at early growth stages. Greenhornoceras (type species Vascoceras (Greenhornoceras) birchbyi Cobban & Scott 1972: 85, pl. 22, pl. 23 (figs 1-13), pl. 24 (figs 1-12), pl. 25, pl. 26 (figs 5-8, 11-12), pl. 27 (figs 1-6)) was erected as a subgenus of Vascoceras for those forms differing in being more involute and in maintaining a squarish or rectangular whorl section. In view of the present systematic chaos within the Vascoceratidae, the value of this subgenus is questionable. Plesiovascoceras Spath (type species Ammonites catinus Mantell 1822: 195, pl. 22 (fig. 10)) was considered by Wright (in Arkell et al. 1957: L419) to show the following characters: ‘Evolute, with very depressed whorl section; early whorls with coarse ribs in pairs, interrupted on venter; later whorls almost smooth apart from very large, blunt umbilical or lateral tubercles. Suture as in Vascoceras.’ Reeside (1923) described a rich Plesiovascoceras fauna from the upper part of the Thermopolis Shale Formation of the Colorado Group in southern Montana. With regard to his material, Reeside (1923: 29) wrote, “The specimens in hand from Montana are a bit confusing in their relations. The general form, except perhaps for the width of the umbilicus, would permit an assignment to UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 129 Fagesia, and the proportions and degree of incision of the suture fit those of Fagesia better than those of Vascoceras. Moreover, the umbilical tubercles are prominent, and there are traces of the ventral ribs in even the largest specimens. The young whorls, however, show distinctly the umbilical and ventral tubercles of Vascoceras, the umbilicus is wider than shown in most of the figured specimens of Fagesia, and the suture in every specimen examined shows a pointed first and second lateral lobe, not bifid. The writer therefore assigns the specimens to Vascoceras rather than Fagesia.’ Freund & Raab (1969: 33) provided the following diagnosis for Fagesia: ‘Inflated cadicones with large umbilicus. Two to three strong prorsiradiate ribs which cross the venter uninterruptedly, split from each strong umbilical tubercle. Ornament preserved until a late stage. In most species the ribs disappear before the tubercles, and in others, the reverse is true. Outermost whorls usually smooth. Suture deeply incised and rather complicated relative to the common one of the family. Three or four high saddles, the first and second often divided into three branches; lobes narrow, deep, and bifid.’ According to Barber (1957: 131), however, “Comparison of the diagnoses of Vascoceras s.s., Fagesia and Plesiovascoceras reveals that the differences are of degree rather than of kind, and no single character may be used to separate the genera. The shell-shape and the stage at which the ornament is lost are perhaps the most diagnostic of the characters and therefore young specimens cannot be definitely assigned to a genus. The suture-line is useful in placing doubtful inter- mediate forms.’ As such, therefore, Fagesia and Plesiovascoceras are separated only on sutural details, a difference which is surely not generic. The genus Gombeoceras Reyment, considered a probable junior synonym of Thomasites by Basse (1940) and Freund & Raab (1969), differs from Vascoceras in the possession of a row of siphonal tubercles. Ezilloella Reyment differs from Vascoceras in having a sharply arched to subcarinate venter on the inner whorls and regularly situated bulges on the flanks of the outer whorls. Neoptychites Kossmat differs from Vascoceras in its extreme involution, and in lacking tubercles at any stage, while Spathitoides Wiedmann, considered a synonym of Vascoceras s.s. by Freund & Raab (1969), is probably best treated as a subgenus of Neoptychites. Fallotites Wiedmann is clearly distinguished from Vascoceras in the presence of distinct upper and lower ventro-lateral tubercles, while in Nigericeras there are also siphonal tubercles. At present the following subgenera may be recognized within the genus Vascoceras: V. (Vascoceras). Compressed to strongly inflated, globose forms in which all ornament is lost on the outer whorls. Inner whorls with prominent swollen umbilical tubercles. Earliest whorls with fold-like ribs which cross the venter. Ribbing quickly becomes effaced across the siphonal line, leaving only ventro-lateral folds. V. (Greenhornoceras). As for V. (Vascoceras), but more involute and main- taining a quadrate whorl section to large diameters. 130 ANNALS OF THE SOUTH AFRICAN MUSEUM V. (Paravascoceras). As for V. (Vascoceras), but umbilical tubercles are lacking at all growth stages. Inner whorls compressed and moderately involute, later becoming somewhat more evolute. Prorsiradiate ribs, uninterrupted across the venter, usually appear at a late growth stage and are generally confined to the ventral region. V. (Plesiovascoceras). Similar to V. (Vascoceras), but more evolute and with a strongly depressed whorl section. Prominent swollen umbilical tubercles retained to maturity. Early whorls with coarse robust ribbing which weakens, and at later growth stages is entirely effaced across the siphonal line. Ribbing may persist to maturity. Genus Vascoceras Choffat, 1898 Subgenus Paravascoceras Furon, 1935 Type species Vascoceras cauvini Chudeau, 1909 Vascoceras (Paravascoceras) cf. cauvini Chudeau, 1909 Figs 6C-H, 35-37 Compare Vascoceras cauvini Chudeau, 1909: 68, pls 1-3. Furon, 1935: 268, pl. 9 Gig, 17). Thomasites cauvini (Chudeau) Chudeau, 1921: 463, fig. 1. Thomasites meslei Chudeau (non Pervinquiere), 1921: 465, pl 1 (figs 1-2), text-fig. 2. Vascoceras (Paravascoceras) cauvini Chudeau, Furon, 1935: 60, pl. 4 (fig. 2), pl. 5 (fig. 1). Vascoceras (Paravascoceras) cauyini var. semiglabra Furon, 1935: 61, pl. 4 (fig. 3). Vascoceras (Pachyvascoceras) crassus Furon, 1935: 58, pl. 3 (fig. 2), text- im, 7, Vascoceras (Paracanthoceras) chevalieri Furon, 1935: 59, pl. 4 (fig. 1), text- fig. 18. Nigericeras jacqueti Schneegans, 1943: 125, pl. 7 (fig. 1). Paravascoceras cauvini (Chudeau) Schneegans, 1943: 128, pl. 5 (fig. 2). Freund & Raab 1969: 20, pl. 3 (figs 1-3). Schobel, 1975: 119, pl. 4 (figs 1-3), pl. 5 (figs 1-4). Paravascoceras cauvini var. inflata Schneegans, 1943: 131. Paravascoceras cauvini var. evoluta Schneegans, 1943: 130, pl. 8 (fig. 2). Paravascoceras crassus var. bullata Schneegans, 1943: 131, pl. 8 (figs 3-4). Paravascoceras chevalieri (Furon) Schneegans, 1943: 132, pl. 4 (fig.-7). Broggiiceras humboldti Benavides-Cacéres, 1956: 471, pl. 56 (figs 1-3). Broggiiceras olssoni Benavides-Cacéres, 1956: 471, pl. 55 (figs 1-4). Paravascoceras rumeaui Collignon, 1957: 10, pl. 1 (fig. 2). Freund & Raab, 1969: 21, pl. 3 (figs 4-5), text-fig. 5. UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA By Paravascoceras tayense Freund & Raab (non Faraud), 1969: 23, pl. 2 (fig. 9), text-fig. 5. Paravascoceras crassum (Furon) Freund & Raab, 1969: 24, text-fig. 5. Paravascoceras cf. baroicensis Choffat, Schneegans, 1943: 134, pl. 8 (fig. 1). Paravascoceras aff. cauvini (Chudeau) Barber, 1957: 37, pl. 14 (figs 2-3), pl. 32 (figs 8-9). Collignon, 1965: 21. Paravascoceras aff. chevalieri (Furon) Reyment, 1955: 63, pl. 14 (fig. 1). Stoliczkaia dispar var. attenuata Douvillé, 1931: 29, pl. 2 (fig. 2). Gombeoceras sp. Cooper, 1972: 248. Description Three rather poorly preserved examples, with the recrystallized test pre- served, appear to belong here. They show some variation in ornament, but appear to belong to a single species. Fig. 35. Vascoceras (Paravascoceras) cf. cauvini Chudeau. The holotype of Stoliczkaia dispar var. attenuata Douvillé (after Douvillé 1931). x 1. SAM-PCA2816 (Fig. 6C—D) is fragmentary but the shell appears to have been moderately involute, with a quadrate, compressed whorl section and an evenly rounded venter. The umbilical wall is steep, with a subangular umbilical shoulder. The flanks are flattened, converging slightly towards the venter, with maximum width just above the umbilical shoulder. On the outer whorl, the ornament consists of broad fold-like ribs, strongest on the ventro-lateral shoul- ders but only weakly joined across the venter by faint ribs and growth lines which are convex adorally. SAM-PCA2727 (Fig. 6E-F) and PCA2796 (Fig. 6G—H) differ from the 132 ANNALS OF THE SOUTH AFRICAN MUSEUM above in lacking distinct ventro-lateral folds. Both specimens are moderately involute, slightly less than half the previous whorl being covered, with a com- pressed, oval whorl section. The umbilicus is rather narrow and fairly deep. The umbilical wall is steep and slightly undercut at the umbilical seam, with a well- rounded shoulder. The flanks are broad and flat, converging slightly towards the evenly rounded venter. Besides very faint growth striae, ribbing is virtually absent. However, SAM—PCA2727 shows a trituberculate venter (on the internal mould only) where the outer shell layer has been removed. The suture line is illustrated in Figure 37. Measurements No. D H W W/H U SAM-PCA2816 45 20 (44) 16 (36) 0,80 8 (0,18) Pa 19 7 (37) LCD 1,00 3 (0,16) SAM-—PCA2727 + 42 + 19 (45) + 14 (33) 0,74 + 10 (0,24) . 33 15 (45) 10 (30) 0,67 tO) SAM-PCA2796 37/ 19 61) 13565) 0,68 u Discussion Although there are obvious differences between specimens which lack ventro- lateral folds (SAM-—PCA2727, 2796) and the ribbed forms (SAM—PCA2816 and Douvillés holotype (Fig. 35)), they are grouped together because of their close similarity in all other respects. This view is supported by Schneegans’s (1943) observation that Paravascoceras passes through an unornamented stage between Fig. 36. Vascoceras (Paravascoceras) cf. cauvini Chudeau. Whaorl section of SAM-—PCA2816. x 1. 20 and 55 mm diameter. The trituberculate venter, on the internal mould only of SAM-PCA2727, probably merely reflects its ancestry (Pseudocalycoceras?). The Angolan material most closely resembles V. (P.) cauvini Chudeau (1909: 67, pl. 1-3), and may possibly be conspecific. Chudeau’s holotype, however, is 107 mm in diameter, while Schneegans’s (1943) material is of similar size, and consequently they are not directly comparable. Furthermore, the significant ontogenetic changes in morphology undergone by Paravascoceras make com- parisons even more difficult. Schdbel (1975) has recently demonstrated the extreme morphological varia- tion to be found within topotype material of V. (P.) cauvini, and the Angolan material easily falls within the intraspecific limits of Chudeau’s species. Reserva- UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 133 ap * Fig. 37. Vascoceras (Paravascoceras) cf. cauvini Chudeau. Suture-line of SAM—PCA2727 at about 48 mm diameter. tion as to the identity of the Salinas material stems from the fact that it was collected from surface scree and thus its exact stratigraphic horizon is unknown, together with the acanthoceratid suture-line (Fig. 37) with a broad, bifid lateral saddle. AGE OF THE FAUNA The Salinas fauna has yielded many elements in common with both western Europe and North America, and correlation is clearly possible. In Europe, Juignet & Kennedy (1977) have adopted the following biostrati- graphic subdivision of the Cenomanian of southern England and northern France, including the type area of the Cenomanian, Le Mans (Sarthe): Eucalycoceras pentagonum Zone Acanthoceras jukesbrowni Zone Middle Turrilites acutus Zone Turrilites costatus Zone Mantelliceras dixoni Zone Lower Mantelliceras saxbii Zone Hypoturrilites carcitanensis Zone pect ‘ Sciponoceras gracile Zone At Salinas there is no sign of the characteristic Upper Albian association of mortoniceratids— Stoliczkaia—Anisoceras, nor of the typical Lower Cenomanian faunal assemblage of Mantelliceras—Sharpeiceras—Hypoturrilites. Furthermore, the Middle Cenomanian Acanthoceras—Calycoceras—Turrilites faunal association recorded from Novo Redondo (Cooper 1973) is also missing at Salinas. In England, the zone of Eucalycoceras pentagonum is characterized by the appearance of Calycoceras naviculare while Acanthoceras hippocastanum (J. de C. Sowerby), Eucalycoceras pentagonum (Jukes-Browne) and Schloenbachia lymense Spath are typical. Other important elements of the fauna are Puzosia (Austini- ceras) austeni (Sharpe), Calycoceras gr. newboldi (Kossmat), Scaphites equalis J. Sowerby and Protacanthoceras spp. Rare components of the fauna include Calycoceras (Lotzeites) aberrans (Kossmat), Carthaginites cf. inornatus (Collig- non), Euomphaloceras euomphalum (Sharpe), Turrilites costatus Lamarck, Thomelites sornayi (Thomel), and T. prerusticum (Thomel). 134 ANNALS OF THE SOUTH AFRICAN MUSEUM The first appearance of Metoicoceras geslinianum (d’Orbigny) marks the base of the Sciponoceras gracile Zone. Still present are Calycoceras naviculare and Scaphites equalis, while Puzosia (Anapuzosia) dibleyi (Spath) appears for the first time. Other forms include “Pachydiscus’ sp. and Sciponoceras sp. In the ‘Metoicoceras gourdoni’ Zone, M. geslinianum still occurs, while also present are “M. gourdoni’ (de Grossouvre), Allocrioceras annulatum (Shumard), E. (Kanabiceras) septemseriatum (Cragin), Tarrantoceras faustum (Matsumoto & Muramoto), Sciponoceras gracile (Shumard), Pseudocalycoceras angolaense (Spath) and ?Puebloites sp. (W. J. Kennedy pers. comm.). Horizon A, which in northern France has as its equivalent the Craie a Terebratella carantonensis (d’Orbigny), was introduced to cover a poorly known part of the sequence yielding Sciponoceras of the gracile group and Neocardio- ceras sp., but few other ammonites. The Mammites nodosoides Zone at the base of the classical Turonian (Juignet et al. 1973; Kennedy & Juignet 1973) is characterized by the occurrence of Mammites nodosoides (Schlotheim) and its varieties, together with the bivalve Inoceramus (Mytiloides) labiatus labiatus (Schlotheim). Other ammonites include Vascoceras (Plesiovascoceras) catinum (Mantell), Fagesia pachydiscoides (Spath), F. cf. superstes (Kossmat), Metasigaloceras rusticum (J. Sowerby), Lewesiceras peramplum (Mantell) and Watinoceras? spp. There can be little doubt that the bulk of the Salinas fauna finds its equiva- lents in the fauna of the Actinocamax plenus Marls (Jefferies 1962, 1963) and Horizon A (Kennedy & Juignet 1973) in southern England, and in the Sables a Catopygus obtusus in northern France. Uppermost Cenomanian strata are exposed over wide areas of the Western Interior of North America. In Colorado, top Cenomanian strata, referred to a Zone of Sciponoceras gracile, are represented by the basal beds of the Bridge Creek Limestone Member. This unit has yielded a rich ammonite fauna (Cobban 1971; Cobban & Scott 1972) comprising the following forms: Metoicoceras whitei Hyatt (= M. gibbosum Hyatt), Calycoceras naviculare (Mantell), Pseudo- calycoceras dentonense (Moreman) (= P. angolaense (Spath)), Sciponoceras gracile (Shumard), Euomphaloceras (Kanabiceras) septemseriatum (Cragin), Allocrioceras annulatum (Shumard), Worthoceras gibbosum Moreman, W. yer- miculum (Shumard), Puebloites corrugatus (Stanton), Anisoceras _plicatile (J. Sowerby), Hemiptychoceras reesidei Cobban & Scott, Desmoceras (Moremano- ceras) scotti (Moreman) and the bivalve Inoceramus pictus Sowerby. A limestone unit some 6 m higher up has yielded a fauna including Watinoceras coloradoense (Henderson), Ampakabites collignoni Cobban & Scott, Neoptychites xetriformis Pervinquiére, N. cf. cephalotus (Courtiller), Fagesia sp., Mammites sp., Baculites cf. yokoyamai Tokunaga & Shimizu, Puebloites spiralis Cobban & Scott, Vasco- ceras (Greenhornoceras) birchbyi Cobban & Scott and the bivalve Jnoceramus labiatus. The close similarity with the Angolan fauna leaves little doubt as to their contemporaneity. In Texas, the zone of Metoicoceras whitei Hyatt is represented in the Britton UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 135 Member of the Eagle Ford Group. Moreman (1942) has listed the following ammonites from this unit: Tarrantoceras bentonianum (Cragin), Pseudocalyco- ceras angolaense (Spath) (including Eucalycoceras dentonense Moreman, E. lewis- villense Moreman, E. indianense Moreman and E. underwoodi Powell), Desmo- ceras (Moremanoceras) scotti (Moreman), Allocrioceras annulatum (Shumard) (including A. pariense (White)), A. larvatum (Conrad), A. dentonense Moreman, A.? rotundatum (Conrad), Metoicoceras gibbosum Hyatt (including M. whitei Hyatt, M. acceleratum Hyatt, M. kanabense Hyatt, and M. ornatum Moreman), M. irwini Moreman, Worthoceras vermiculum (Shumard), W. gibbosum More- man, Euomphaloceras (Kanabiceras) septemseriatum (Cragin), Sciponoceras gracile (Shumard), Scaphites brittonense Moreman, S. minutus Moreman, Proten- gonoceras planum Hyatt, Epengonoceras acutum Hyatt, Proplacenticeras cum- minsi (Cragin), P. pseudoplacenta occidentale (Hyatt) and P. stantoni bolli (Hyatt). Although this list needs revision, and there are a number of apparently endemic taxa within the Texas fauna, there can be no question about its contemporaneity with the Angolan fauna. A typical uppermost Cenomanian fauna is known (Matsumoto ef al. 1957; Matsumoto 19595; Matsumoto et al. 1969; Matsumoto & Kawano 1975) from the top of unit IIc, middle member of the Mikasa Formation, central Hokkaido, Japan, including FE. (Kanabiceras) septemseriatum (Cragin), Sciponoceras gracile (= S. kossmati (Nowak)), Pseudocalycoceras angolaense (Spath) (= Eucalyco- ceras sp., Matsumoto ef al. 1957: 38, pl. 11 (fig. 2)) and Tarrantoceras faustum (Matsumoto & Muramoto). In both north-western Europe (Kennedy 1971; Kennedy & Juignet 1973; Juignet et al. 1973) and North America (Cobban & Scott 1972) the faunal associa- tion Sciponoceras gracile—Kanabiceras septemseriatum—Metoicoceras spp. is con- sidered diagnostic of the uppermost Cenomanian. The writer concurs with these opinions. Records of Metoicoceras from the Lower Cenomanian of Madagascar (Collignon 1964) clearly need reinvestigation, while the appearance of this genus in the Lower Turonian of Israel (Freund & Raab 1969) is possibly based upon misidentification. Collignon (1966) recorded Metoicoceras from the Lower Turonian of Morocco, but these faunas are almost certainly contemporaneous with the Angolan assemblage. The occurrence at Salinas of Watinoceras coloradoense (Henderson) and Vascoceras (Paravascoceras) cf. cauvini Chudeau suggests a slightly higher faunal horizon (the basal Turonian zone of Watinoceras coloradoense) to be present, although as these forms were picked up in surface scree this needs substantiating. THE CENOMANIAN-TURONIAN BOUNDARY PROBLEM The exact position of the Cenomanian—Turonian boundary has long been a point of contention. In north-western Europe, it has been variously drawn, on both litho- and biostratigraphic grounds, at below the plenus Marls (= geslinia- num + gourdoni Zones) (Thomel 1972, 1973; Jefferies 1962, 1963), some way above the plenus Zone (Kennedy 1971; Kennedy & Juignet 1973), and at a still 136 ANNALS OF THE SOUTH AFRICAN MUSEUM higher horizon (Berthou & Lauverjat 1974), so as to include a zone of Vascoceras gamai-mundae within the Cenomanian. The type area of the Cenomanian is in the vicinity of Le Mans, Sarthe, north-western France (d’Orbigny 1847), while ‘l’etage Turonien’ was first recog- nized (d’Orbigny 1842) along the Cher Valley between Montrichard and Tours, Touraine. This stage was later restricted (d’Orbigny 1847: 270) to those beds containing Ammonites lewesiensis (= Lewesiceras), A. peramplus (= Lewesiceras) A. vielbanci (= Mammites of authors), A. woollgari (= Collignoniceras), A. fleuri- ausianus (= Prionotropis of authors), A. deverianus (= Romaniceras). Kennedy & Juignet (1973) have recently shown there to be a considerable overlap between the succession of the type Cenomanian and that of the Turonian stratotype. These authors recognized the following lithostratigraphic and ammonite succession across the Cenomanian—Turonian boundary. From above downwards they are: (iv) Craie a Inoceramus labiatus—characterized by the appearance of I. labiatus labiatus Schlotheim and Mammites nodosoides (Schlotheim). (iii) Craie a Terebratella carantonensis—in places Inoceramus gr. pictus J. Sowerby occurs in situ, but the only known ammonites are derived Upper Cenomanian forms. (ii) Sables a Catopygus obtusus—yielding a fauna characterized by Metoico- ceras gourdoni (de Grossouvre), M. geslinianum (d’Orbigny). Evompha- loceras (Kanabiceras) septemseriatum (Cragin), Sciponoceras gracile (Shumard), Calycoceras naviculare (Mantell) and Watinoceras? sp. (i) Marnes a Ostrea biauriculata—with Calycoceras naviculare (Mantell), Thomelites sornayi (Thomel), Eucalycoceras rowei Spath and Calyco- ceras gr. newboldi (Kossmat). Kennedy & Juignet (1973: 199) were led to conclude that since “the Metoico- ceras gourdoni Zone is present within the type Cenomanian, where it is repre- sented by the Sables a Catopygus obtusus. Their fauna, including key forms such as M. geslinianum and T. carantonensis were always regarded by d’Orbigny as Cenomanian’, that this Zone should be placed in the Cenomanian, and that ‘the most convenient level for the base of the Turonian would thus appear to be the base of the classic Mammites nodosoides|Inoceramus labiatus Zone, defined on its fauna and not on lithology’. Unfortunately, they do not specify what this fauna is, and undoubted Turonian ammonites occur below either of these taxa. In order to objectively review the Cenomanian—Turonian boundary prob- lem, it is necessary to look at the present biostratigraphical zonation through these two stages. After Wright (in Arkell et al. 1957) and Juignet & Kennedy (1977), these are: Upper Turonian Subprionocyclus neptuni Middle Turonian Collignoniceras woollgari Lower Turonian Mammites nodosoides Sciponoceras gracile Upper Cenomanian { Eucalycoceras pentagonum UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 137 In the classical Turonian of Le Mans, Sarthe, Kennedy & Juignet (1973) recognized a zone of Mammites nodosoides only in the Lower Turonian. How- ever, in those parts of the world from where rich and diversified Lower Turonian faunas are known, the base of the Turonian is characterized by strata rich in Vascoceras s.s., together with forms of V. (Paravascoceras), Nigericeras, Gombeo- ceras, Watinoceras, Fallotites and Fagesia, while Mammites appears for the first time only well above the Vascoceras faunas in Israel (Freund & Raab 1969), Nigeria (Reyment 1955; Barber 1957), northern Spain and Portugal (Wiedmann 1959, 1964), and the Western Interior of North America (Cobban & Scott 1972). In Madagascar, M. nodosoides is known only from the Middle Turonian ‘Zone a Pseudaspidoceras conciliatum (Stoliczka)’ (Collignon 1965). The relative paucity of fossils within the early Turonian chalk environment of north-western Europe does not allow for the recognition of a detailed biostratigraphical zonation, and it becomes necessary to look elsewhere in the world. Choffat (1900) subdivided the Upper Cenomanian-Turonian of Portugal into 12 units, numbered alphabetically from C to N. From above downwards they were: M, N' Limestones with gasteropods IL Beds with Fagesia superstes, Choffaticeras barjonai, Vascoceras durandi and Inoceramus labiatus I-K __Unfossiliferous limestones and dolomites H Limestones which at their summit yield Fallotites subconciliatus, Vascoceras durandi and Pseudaspidoceras footeanum G Passage beds lacking in fossils E-F Beds with Vascoceras gamai and V. mundae D Beds with the echinoids Anorthopygus michelini and A. orbicularis, but lacking in ammonites. C Beds with Neolobites vibrayeanus (= N. choffati Hyatt) and Calyco- ceras naviculare A supposedly characteristic Cenomanian microfauna from Beds E-F led Berthou & Lauveryjat (1974) to include a zone of Vascoceras gamai and V. mundae at the top of the Cenomanian. This was supported by Thomel’s (1972) assignment of Choffat’s C. naviculare to C. stoliczkai (Collignon), a species considered by him to be restricted to his zone of Calycoceras robustum Thomel, at the base of the Upper Cenomanian. As shown by Cobban (1971) and Kennedy (1971), how- ever, the specimens figured by Choffat (1898: 72, pl. 4 (fig. 6a—b), pl. 6 (figs 1—2)) are true C. naviculare (Mantell), of which C. stoliczkai is a junior subjective synonym. Consequently, Calycoceras naviculare is a relatively long-ranging species which is known from many levels in the Upper Cenomanian and its occurrence some way below strata with Vascoceras is not considered significant. The problem of whether or not to include a zone of Vascoceras gamai Choffat at the top of the Cenomanian is difficult. Like Kennedy & Juignet (1973), the writer prefers to regard the genus Metoicoceras as being restricted to an 138 ANNALS OF THE SOUTH AFRICAN MUSEUM horizon very high in the Cenomanian. Its relationship to the vascoceratid faunas is thus critical. In Israel, Freund & Raab (1969: 78) record Metoicoceras cf. whitei Hyatt, unfortunately without figure or proper description, from their Zone 4 of Choffati- ceras securiforme (Eck). If correctly identified, this could provide strong sup- porting evidence for the inclusion of at least part of the vascoceratid faunas within the Cenomanian. It seems more likely, however, that the Israeli form may be a misidentification of a typically Lower to Middle Turonian Spathites (Spathites) or S. (Jeanrogericeras). From the Puentedey valley, near Soncillo, Villamartin, Turzo (Burgos Province), northern Spain, Wiedmann (1964) gave a detailed succession across the Cenomanian—Turonian boundary. From above downwards this was: (vii) 6 m of blue marls with Fallotites UIngridella) cf. malladae (Fallot), Wrightoceras mirabile (Pervinquiere), W. llarenai (Karrenberg), Hopli- toides sp., Proromaniceras pseudodeverianum (Jimbo), Vascoceras (Para- vascoceras) grossouvrei (Choffat), Spathites laevis (Karrenberg), Jean- rogericeras revelieranum (Courtiller), Schindewolfites inaequicostatus Wiedmann. (vi) 10 m of blue marls with Choffaticeras quaasi (Peron), C. pavillieri (Per- vinquiere), Vascoceras durandi (Peron), Spathites cf. laevis (Karrenberg), Fallotites (Fallotites) sp. (v) 2 m of blue marls with F. (Fallotites) cf. subconciliatus (Choffat). (iv) 5 m of blue marls with Vascoceras (Plesiovascoceras) fagesioides Wied- mann, Vascoceras cf. gamai Choffat, Watinoceras sp. and Metoicoceras cf. swallovi (Shumard). (ii) 4 m of marls with M. swalloyvi (Shumard), M. cf. swallovi (Shumard), M. cf. whitei Hyatt, ‘Parapuzosia’ cf. gaudama (Forbes). (ii) 2 m of marls and limestones with Metoicoceras muelleri Cobban and Rhynchostreon suborbiculatum (Lamarck). (i) 2mof marls with Neolobites brancai Eck, N. cf. vibrayeanus (d’Orbigny), N. choffati Hyatt, N. cf. schweinfurthi Eck, Calycoceras naviculare (Mantell), Pseudocalycoceras haugi (Pervinquiere). The importance of the Spanish sections cannot be overestimated, since this is one of the few areas where rich Metoicoceras and vascoceratid faunas have been described from the same section. It is also very significant that the vast majority of Metoicoceras occur immediately below the vascoceratids. However, from his Lower Turonian Zone 2, Wiedmann (1964) records Metoicoceras cf. swalloyi (Shumard) in association with Vascoceras (Vascoceras) cf. gamai Choffat and Vascoceras (Plesiovascoceras) fagesioides Wiedmann. It is, unfortunately, not clear whether Metoicoceras occurs side by side with the vascoceratids or at a slightly lower level in the 5 m section. Consequently, once again unequivocal evidence for the overlap of the Metoicoceras and Vascoceras faunas is at present lacking. Cobban & Scott (1972) have recently recorded faunas across the UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 139 Cenomanian—Turonian boundary from the Bridge Creek Limestone Member near Pueblo, Colorado. A typical Sciponoceras gracile faunal assemblage, placed as the uppermost zone in the Cenomanian, occurs associated with the bivalve Inoceramus pictus Sowerby. Some 3 m higher up, a 15 cm limestone bed has yielded Vascoceras (Greenhornoceras) birchbyi Cobban & Scott, Watinoceras coloradoense (Henderson), Puebloites spiralis Cobban & Scott and a Fagesia sp., in association with the typically Turonian Jnoceramus labiatus Schlotheim. Also significant is the fact that Mammites nodosoides (? non Schlotheim) first appears some 1,5 m above strata with Vascoceras. The only continuous section through the lower part of the Lower Turonian of Nigeria is at Pindiga (Barber 1957) where some 30 m of shales, sandy mud- stones and sandstones, lacking in ammonites, rest conformably upon terrestrial beds and are overlain by about 80 m of fossiliferous strata, Barber (1957: 60) gave the following succession from above downwards: (viii) About 40 m of shales with thin nodular horizons yielding the ammonites Pseudotissotia (Bauchioceras) nigeriensis nigeriensis (Woods), P. (B.) n. tabulata Barber and Eotissotia simplex Barber. (vii) 0,8 m of rubbly limestone with Vascoceras globosum (Reyment), Gombeoceras gongilense gongilense (Woods), G. g. lautum Barber, G. g. tectiforme Barber, Paramammites sp. and Nigericeras sp. (vi) 4 m of barren shale. (v) 0,22 m of crystalline limestone with Vascoceras nigeriense Woods, V. ellipticum Barber, V. polygonum Barber, V. (Paravascoceras) costa- tum (Reyment), Gombeoceras gongilense (Woods), G. g. costatum Barber. (iv) About 60 m of unfossiliferous shales. (iii) 2 m of nodular limestone with Vascoceras bulbosum (Reyment) and V. depressum Barber. (ii) Some 12 m of barren shale. (i) 0,3 m of crystalline limestone with Vascoceras bulbosum (Reyment) and Epengonoceras dumbli (Cragin). It is of interest to note Barber’s (1957: 61) observation that “The fauna associated with the Salmurian ammonites here described has many members which may occur in the Cenomanian. It would appear therefore that the Nigerian Salmurian occurs close to the Cenomanian—Turonian boundary.’ Moreover, Epengonoceras dumbli is known only from the Middle and Upper Cenomanian in Texas and the Western Interior. From the Cerro del Macho in Coahuila, Mexico, Bése (1918) described a rich upper Cenomanian—Lower Turonian fauna. Bose (1918: 183) gave the following section, from above downwards: (iii) 5-6 m of hard gray limestone with Vascoceras (Paravascoceras) anger- manni Bose, V. ex aff. gamai Choffat, Neoptychites aff. xetriformis Pervinquiere, Hoplitoides aff. mirabilis Pervinquiere and Inoceramus labiatus Schlotheim. 140 ANNALS OF THE SOUTH AFRICAN MUSEUM (11) 2,5 m of bluish-gray marls with Fallotites mohovanensis (Bose), Pseuda- spidoceras flexuosum Powell, Pseudaspidoceras aff. pedroanum White, Vascoceras aff. adonense Choffat, Fagesia haarmanni Bose and Fagesia pervinquieri Bose. (i) 2 m of yellow and reddish marls and limestones with Metoicoceras aff. whitei Hyatt, M. boesei Jones and Exogyra (Costagyra) cfr. olisiponensis Sharpe. The above section once again clearly shows Metoicoceras to occur immedi- ately below what have always been regarded as typical Lower Turonian ammonites. It becomes obvious, therefore, that for both biostratigraphic correlation and in order to clarify the Cenomanian—Turonian boundary problem, it is important to introduce a zone beneath the classical Lower Turonian zone of Mammites nodosoides. In the Western Interior, Cobban & Scott (1972) have recognized a basal Turonian zone of Watinoceras coloradoense. The wide geographic distribu- tion of this species, together with the fact that Watinoceras also appears to mark the base of the Turonian in Angola, Spain and possibly the Anglo-Paris basin, all seem to support its usage as the basal Turonian index fossil. There can be little doubt that with the rich faunas available, together with a better under- standing of vascoceratid taxonomy and stratigraphy, a far more refined zonal scheme for the Lower Turonian will eventually be possible. At present only the following Upper Cenomanian—Lower Turonian ammonite zones appear to be of world-wide importance: Lower Turonian Mammites nodosoides Watinoceras coloradoense Upper Cenomanian Sciponoceras gracile Eucalycoceras pentagonum These are assemblage zones, and may be briefly diagnosed as follows: Eucalycoceras pentagonum Zone—typical elements of the fauna include E. penta- gonum (Jukes-Browne), Acanthoceras hippocastanum (J. de C. Sowerby), Protacanthoceras of the bunburianum-compressum group, Thomelites spp., Pseudocalycoceras harpax (Stoliczka), Euomphaloceras euomphalum (Sharpe), Calycoceras (Newboldiceras) spp., C. (Calycoceras) boulei Collignon and C. (C.) naviculare (Mantell). Sciponoceras gracile Zone—characterized by S. gracile (Shumard), Metoicoceras spp., Euomphaloceras (Kanabiceras) septemseriatum (Cragin), Pseudocalycoceras angolaense (Spath), and Calycoceras naviculare (Mantell), together with Puzosia (Austiniceras) austeni (Sharpe), P. (Anapuzosia) dibleyi (Spath), Desmoceras (Moremanoceras) scotti Moreman, Allocrioceras spp., and Tarrantoceras faustum (Matsumoto & Muramoto). Watinoceras coloradoense Zone—the faunas of this zone are not yet well differen- tiated from those of the overlying nodosoides Zone. However, Watinoceras 141 UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA “SOLUDYOS pd}de19 A[SNOIAoId YM poreduos ‘ulosoy posodoid uoneuoz [eorydessyessoig uULIUOIN], JoMOT-URIUeUIOUSD Joddy “g¢ ‘317 wunsSogng svdav0IsvA WINJDISOI SDAIIOISDADAD Sisualsasiu Spsavo1yonvg NYyISa SpAaJoUnABUIDY, WNISNGOA SDAIIOIA]DI WINSSDAI SDABIOIAIDI adAD[NIIADU SDABIOIAIDI Sajsdadns visaso.. SOP1OSOPOU SajiMuopy 1q09vf SDAQIOYJUDIDIOAd, 1D ffoys saj1qojoan’ 1da]JaNUL SD419IOINOJAW 1AO]JOMS SD1BIOINOJAW 1DWUDS SDAIIOISDA SNIDY1IUOIGNS $aJ1J0]]D-] 1ZUADS 4, SAJUMUDUIDADg ADpDIJDUL DjJaplAsuy 1AAIUNUL SDAZIOIY SIA (661) UUPUIPSI A, AADI[NIIADU SDAIIOIA]DID WNUDIUYSAS SDAQIONOJAW 1UOpANOs SD1IIONOJaW V UOZLIOH (€L61) yousiIng 2 Apouudy ‘ds spsav0ndjoD ‘ds sajiqojoan ‘ds spsaoiqouvy 1UIANDI SDAIIOISDADIDY 1101d SDAQIOISDY , aUsOf1AnJas spsanijoffoy) isponb spszayoffoy) av1In] SDABIIJO JOY) (6961) qeey 2 punory SNUDAADAGIA SaJ1GOJOaN HCH Cd snsddoyjsoup D0 nvaAIN aDpunuU —1DUDS sD12ad0ISD A aspsspd ap ayamnoy 1pup.inp —lapjlanop spsav0asvadyovg Sajsdadns pisasvq (SL61) yelIoaAne’yT 2 noy.og 1puod spsavouvsaaunq WNJIPuoD SDAQIOUDBAAUNG aSuajsaqg]|D SDAIIOUDSaAUNG aj1avAs SpsvIOUOdIIG ISUBOPDAOJOI SDADIOULID A SAPlOSOpou sSajluUD Ww (ZL61) 09g 2 ueqqoD winuosvJUuad SpsaI0IA]VINT aplIvAs SDsaIoUOdIIG SOPlOSOpOU Sajid py (LL61) Apouusy 2 Jousine wNnuosvDJUad SDABIOIA]DIN] ISUIOPDAOJOID SDADIOUIID SaploOsOpou sajluUDw UIdIOH] WNnUOsvDJUad SDABIOIAIDINT AlIDAS SDAIIOUOAIIS ISUIOPDAOJOI SDAIIOUIID A SOP1OSOPOU SAJIMUD UIdIOH] NVINVWONd) wddddn NVINOYWNL YUAMOT NVINOUNL NVINVWONA) YUAMOT ddddn 142 ANNALS OF THE SOUTH AFRICAN MUSEUM coloradoense (Henderson) and Vascoceras (Paravascoceras) cauvini Chudeau appear to be typical, possibly also with species of Pseudaspidoceras, Nigericeras, V. (Vascoceras) of the gamai-mundae group, V. (Plesiovascoceras), Fagesia, Neoptychites and Fallotites. Mammites nodosoides Zone—it is from this zone that most of the classical Lower Turonian ammonites have been collected, and further subdivision will undoubt- edly prove possible. Typical elements of the fauna include M. nodosoides (Schlot- heim), together with species of Choffaticeras, Fallotites, Hoplitoides, Neoptychites, Kamerunoceras, Schindewolfites, Pseudotissotia (Wrightoceras), Thomasites, Vascoceras, Fagesia, Spathites (Spathites), S. (Jeanrogericeras) and Donenriquo- ceras. As can be seen from Figure 38, there has been considerable debate as to the position of the Cenomanian—Turonian boundary. The writer prefers to include the Sciponoceras gracile Zone as the uppermost biostratigraphical unit in the Cenomanian for the following reasons: (i) A typical Cenomanian acanthoceratinid fauna, including Calycoceras, Pseudocalycoceras, Tarrantoceras, Metoicoceras and possibly Protacan- thoceras persist into this zone. (11) The diagnostic Upper Cenomanian Calycoceras naviculare (Mantell) is abundant in the gracile Zone. (iii) The persistence of the typically Cenomanian Jnoceramus pictus into this zone. (iv) The association of the diagnostic Lower Turonian Jnoceramus labiatus with most vascoceratid faunas, and its absence from the gracile Zone. PALAEOGEOGRAPHIC SIGNIFICANCE The Salinas fauna is part of the characteristic, world-wide (Fig. 39), upper- most Cenomanian Sciponoceras gracile faunal assemblage. Members of this fauna are widely distributed in the Western Interior of North America (Cobban & Scott 1972), extending as far north as Montana (Cobban 1953) and as far south as Texas (Moreman 1942; Powell 19635) and Mexico (Bése 1918; Jones 1938). In the Anglo-Paris basin, the association of Sciponoceras gracile—Kanabiceras septemseriatum—Metoicoceras spp. is known over wide areas (Jefferies 1962, 1963; Juignet et al. 1973). In Spain and Portugal (Karrenberg 1935; Wiedmann 1959, 1964) a rich Metoicoceras fauna is reported from the top of the Cenomanian, but the exact constituents are at present poorly known. Metoicoceras is also known from Morocco (Collignon 1966), Israel (Avnime- lech & Shoresh 1962), Nigeria (Reyment 1955), Brazil (Reyment & Tait 1972a) and Madagascar (Collignon 1964), but the stratigraphy of these areas is poorly understood, and the associated ammonite faunas are not known. The record of this genus from southern India (Sastry & Matsumoto 1967) is based upon the closely allied Lower Turonian Spathites (Jeanrogericeras). Enigmatic is the almost complete absence of Metoicoceras, save for the Madagascar occurrence, ee UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 143 m4 Fig. 39. Palaeogeographic reconstruction of Laurasia and the southern continents during the latest Cenomanian-earliest Turonian showing distribution of Sciponoceras gracile Zone and Watinoceras coloradoense Zone fauna. m = Metoicoceras spp., k = Euomphaloceras (Kanabi- ceras) septemseriatum (Cragin), c = Calycoceras naviculare (Mantell), s = Sciponoceras gracile (Shumard), p = Pseudocalycoceras angolaense (Spath), d = Puzosia (Anapuzosia) dibleyi (Spath), w = Watinoceras spp. 144 ANNALS OF THE SOUTH AFRICAN MUSEUM from the Indo-Pacific realm. Its absence along the east coast of Africa, however, is related to the absence of strata of this age (Cooper 1974). Sciponoceras gracile (Shumard) (including S. kossmati) is recorded from California (Anderson 1958; Matsumoto 1959a) in association with E. (K.) septem- seriatum and C. naviculare, as also in Japan (Matsumoto & Obata 1963) where other elements of the fauna include Tarrantoceras and Pseudocalycoceras. Elements of the gracile Zone fauna recorded from southern India (Stoliczka 1865; Kossmat 1895) include the relatively long-ranging C. naviculare and S. gracile, but the exact associations are not known; Metoicoceras and Kanabi- ceras are unrecorded and it is possible that uppermost Cenomanian strata are absent. Thus, a typical gracile Zone fauna is known from as far afield as North America (California, Western Interior, Texas, Mexico), South America (Brazil), Africa (North Africa, Morocco, Nigeria, Angola), the Middle East, France, England, India? and Japan. Its absence from the Arctic and austral realms may be due to absence of strata of this age, although it is more likely that there was an ecological barrier (? cold water). A gracile Zone fauna appears, therefore, to have attained a virtually cosmopolitan distribution within the equatorial and temperate waters of the late Cenomanian oceans. A number of authors (Reyment 1969, 1971la, 1972, 1973; Reyment & Tait 1972a, 19726) have concluded from a study of Cretaceous ammonite distributions in the sedimentary basins surrounding the South Atlantic that a continuous con- nection between the North and South Atlantic Oceans was not established until the end of the Lower Turonian. In this respect the position of Angola on the initial line of rifting of the. Africa-South America plates is critical. The occurrence of a typical uppermost Cenomanian faunal assemblage from Salinas, virtually identical to similar faunas in North America and western Europe, indicates this seaway to have been open prior to the beginning of the Turonian (Kennedy & Cooper 1975). However, since the uppermost Cenomanian represents a period of eustatic transgression (Cooper 1974), faunal interchange could have occurred merely by flooding of the hypothetical Nigeria—Pernambuco connection. In this respect, the Middle Cenomanian (Turrilites costatus and T. acutus Zones) fauna recorded from Novo Redondo (Cooper 1973) is also of little help, since this period may also have been an episode of eustatic trans- gression (Hart & Tarling 1974). To determine the dating of the final break between Africa and South America, i.e. the establishment of a continuous marine connection, it is important to study the faunas of regressive episodes, when exposure of land-bridges, and hence barriers to faunal migration, are most likely to have occurred. The uppermost Albian (dispar Zone) faunas provide this oppor- tunity. Whilst the dispar Zone fauna of Angola is poorly documented, it is extremely rich and includes the following European species: Anisoceras armatum (J. Sowerby), A. perarmatum Pictet & Campiche, Stoliczkaia clavigera Neumayr, Mortoniceras (Durnovarites) perinflatum (Spath), M. (D.) subquadratum Spath, Cantabrigites curvatum Renz, Mariella gresslyi (Pictet & Campiche), and possibly UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 145 also the Texas mortoniceratinid genus Drakeoceras (Cooper 1977). The latest Albian faunas of Angola are, therefore, virtually identical to similar faunas in western Europe at a time of world-wide eustatic regression, which would seem to indicate continuous faunal interchange between the two areas. This suggests that the final rifting between the African and South American plates took place sometime prior to the end of the Albian. SUMMARY The type locality of the classic Salinas fauna described by Douvillé was revisited and new material collected. The very high Cenomanian age of the fauna, suspected by previous authors, was confirmed and shown to form part of the world-wide Sciponoceras gracile faunal assemblage. The presence of Watinoceras and V. (Paravascoceras) in surface scree at this locality suggests the presence of basal Turonian strata. The geological section was measured and the succession is interpreted as a typical transgressive sequence. The problem of the Cenomanian—Turonian boundary is reviewed, and Cobban & Scott (1972) are followed in drawing it between the zones of Watino- ceras coloradoense above and Euomphaloceras (Kanabiceras) septemseriatum below. Within the family Acanthoceratidae, the new subfamily Euomphalocera- tinae is erected to incorporate the following taxa: E. (Euomphaloceras), E. (Kana- biceras), Schindewolfites, Kamerunoceras, Yubariceras, Romaniceras, Obiraceras, Shuparoceras and tentatively also Tunesites. The full faunal list from Salinas now reads: Calycoceras (Calycoceras) naviculare (Mantell) (= Acanthoceras borgesi Douvillé) Pseudocalycoceras angolaense (Spath) (= Lyelliceras lyelli Douvillé non Leymerie) Pseudocalycoceras aff. haugi (Pervinquiére) ?Protacanthoceras spp. Tarrantoceras sp. juv. indet. Euomphaloceras (Kanabiceras) septemseriatum (Cragin) (= Prionotropis echinatus Douvillé) Watinoceras coloradoense (Henderson) Metoicoceras gibbosum Hyatt (= Pulchellia caicedoi Douvillé non Karsten) Gaudryceras (Gaudryceras) isovokyense Collignon Anagaudryceras cf. cassisianum (d’Orbigny) (= Gaudryceras salinarium Douvillé) Tetragonites aff. blaisoni Collignon Desmoceras (Pseudouhligella) aff. ezoanum Matsumoto (= Desmoceras toucasi Douvillé non Jacob) ?Proplacenticeras sp. 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Some acanthoceratids from Hokkaido. Mem. Fac. Sci. Kyushu Univ. (D) Geol. 6: 1-45. McLearn, D. 1972. Ammonoids of the Lower Cretaceous sandstone member of the Haida Formation, Skidegate Inlet, Queen Charlotte Islands, western British Columbia. Bull. geol. Surv. Canada 188: 1-78. MoreEMAN, W. L. 1927. Fossil zones of the Eagle Ford Group of north and central Texas. J. Paleont. 1: 89-101. MoreMAaNn, W. L. 1942. Paleontology of the Eagle Ford Group of north and central Texas. J. Paleont. 16: 192-220. Murphy, M. A. 1967a. The Aptian—Cenomanian members of the ammonite genus Tetragonites. Univ. Calif. Publs geol. Sci. 69: 1-78. Murpny, M. A. 19676. The ammonoid subfamily Gabbioceratinae Breistroffer. J. Paleont. 41: 595-607. Nowak, J. 1908. Untersuchungen iiber die Cephalopoden der obéren kreiden in Polen. Bull. Acad. Sci. Cracovie 1908: 327-353. 150 ANNALS OF THE SOUTH AFRICAN MUSEUM OrBiGny, A. D’. 1840-1842. Paléontologie Francaise; Terrains crétacés. I, Céphalopodes: 1-662. Paris: Masson. OrsBIGNy, A. D’. 1847. Paléontologie francaise; Terrains crétacés IV. Brachiopodes: 1-390. Paris: Masson. OrBIGNY, A. D’. 1850. Prodrome de paléontologie stratigraphique universelle des animaux mol- lusques et rayonnés. Paris: Masson. PERVINQUIERE, L. 1907. Etudes de paléontologie tunisienne. 1. Céphalopodes des terrians secon- daires. (Régence de Tunis . . . Carte géologique de la Tunisie.) Paris: De Rudeval. PETRASCHECK, W. 1902. Die Ammoniten der sachsischen Kreideformation. Paldont. Oest-Ung. Orients 14: 131-162. PicteT, F. J. & CAMPICHE, G. 1858-1864. Matériaux pour la paléontologie Suisse. Description des fossiles du terrain crétacé des environs de Sainte-Croix. Geneva. Popovici-HATZEG, V. 1899. Contributions a l’étude de la faune du crétacé supérieur de Roumanie. Environs de Campulung et de Sinaia. Mem. Soc. géol. Fr. (Paléont.) 8: 1-20. POWELL, J. D. 1963a. Cenomanian—Turonian (Cretaceous) ammonites from trans-Pecos Texas and north-eastern Chihuahua, Mexico. J. Paleont. 37: 309-322. PowELL, J. D. 19635. Turonian (Cretaceous) ammonites from north-eastern Chihuahua, Mexico. J. Paleont. 37: 1217-1232. REESIDE, J. B. 1923. A new fauna from the Colorado Group of southern Montana. Prof. Pap. U.S. geol. Sury. 132-B: 25-33. REESIDE, J. B. & COBBAN, W. A. 1960. Studies of the Mowry Shale (Cretaceous) and contem- porary formations in the United States and Canada. Prof. Pap. U.S. geol. Surv. 355: 1-126. REESIDE, J. B. & WeymouTH, A. A. 1931. Molluscs from the Aspen Shale (Cretaceous) of south-western Wyoming. Proc. U.S. natn. Mus. 78: 1-24. RENZ, O. 1972. Die Gattungen Puzosia Bayle, Bhimaites Matsumoto und Desmoceras Zittel (Ammonoidea) im oberen Albien Venezuelas. Eclogae geol. Helv. 65: 701-724. REYMENT, R. A. 1954a. New Turonian (Cretaceous) ammonite genera from Nigeria. Colon. Geol. Miner. Resour. 4: 149-164. REYMENT, R. A. 19546. Some new Upper Cretaceous ammonites from Nigeria. Colon. Geol. Miner. Resour. 4: 248-270. REYMENT, R. A. 1955. The Cretaceous Ammonoidea of southern Nigeria and the southern Cameroons. Bull. geol. Surv. Nigeria. 25: 1-112. REYMENT, R. A. 1969. Ammonite biostratigraphy, continental drift and oscillatory transgres- sions. Nature, Lond. 224: 137-140. REYMENT, R. A. 1971la. Experimental studies of Cretaceous transgression for Africa. Bull. geol. Soc. Am. 82: 1063-1072. REYMENT, R. A. 19715. Vermeteter Dimorphismus bei der Ammonitengattung Benueites. Bull. geol. Instn Univ. Upsala (n.s.) 3: 1-18. REYMENT, R. A. 1972. Cretaceous (Albian-Turonian) geology of the South Atlantic. Proc. Conf. Afr. Geol., Dec. 1970. Nigeria: Ibadan University Press. REYMENT, R. A. 1973. Paléontologie evolutive et nouvelle tectonique. Publ. Palaeont. Inst. Univ. Upsala 135: 1-18. REYMENT, R. A. & Tart, E. A. 1972a. Biostratigraphical dating of the early history of the south Atlantic Ocean. Phil. Trans. R. Soc. Lond. (B) 264: 55-95. REYMENT, R. A. & Tait, E. A. 19726. Faunal evidence for the origin of the South Atlantic. Publ. Palaeont. Inst. Univ. Upsala 131: 316-323. SAsTRY, M. V. A. & Matsumoto, T. 1967. Notes on some Cretaceous ammonites from southern India. Part II. Occurrence of Metoicoceras in Trichinopoly Cretaceous. Mem. Fac. Sci. Kyushu Univ. (D) Geol. 18: 1-5. SAYN, G. 1890. Description des ammonitides du Barrémien du Djebel-Ouach, prés Constantine. Annls Soc. Agric. Hist. nat. Lyon (6) 3: 135-208. SCHLUTER, C. 1871-76. Die Cephalopoden der oberen deutschen Kreide. Palaeontographica 24: 1-24 (1871); 24: 25-120 (1872); 24: 121-264 (1876). SCHNEEGANS, D. 1943. Invertébrés du Crétacé supérieur du Damergou (Terretoire du Niger). In: Etudes stratigraphiques et paléontologiques sur le Bassin du Niger. Bull. Fr. W. Afr. Direction Mines 7: 87-150. SCHOBEL, J. 1975. Ammoniten der familie Vascoceratidae aus den unteren Unterturon des Damergoi-gebietes, Republique du Niger. Publ. palaeont. inst. Univ. Upsala 3: 1-136. UPPERMOST CENOMANIAN-BASAL TURONIAN AMMONITES FROM SALINAS, ANGOLA 151 SHARPE, D. 1853-1857. Description of the fossil remains of Mollusca found in the Chalk of England. I. Cephalopoda. Palaeontogr. Soc. (Monogr.): 1-68. Sumer, H. W. & SHROCK, R. R. 1944. Index fossils of North America. New York: Wiley & Sons. SHIMIZU, S. 1934. Ammonites. In: SHimi1zu, S. & OBATA, T. Cephalopoda. Twanami’s series of geology and palaeontology. Tokyo. (In Japanese.) SHIMIZU, S. 1935. The Upper Cretaceous Cephalopoda of Japan. J. Shanghai Sci. Inst. (2) 1: 159-226. SHUMARD, B. F. 1861. Description of new Cretaceous fossils from Texas. Trans. Acad. Sci. St. Louis 1: 590-610. Sornay, J. 1955. Ammonites (Mammites) geslinianus d’Orbigny. Palaeont. Universalis, n.s., 11. SpATH, L. F. 1922. On the Senonian ammonite fauna from Pondoland. Trans. R. Soc. S. Afr. 10: 113-147. SPATH, L. F. 1923. On the ammonite horizons of the Gault and contiguous deposits. Mem. geol. Sury. Summ. Prog. (1922): 139-149. SpATH, L. F. 1925. On Senonian Ammonoidea from Jamaica. Geol. Mag. 62: 28-32. SpATH, L. F. 1926. On the zones of the Cenomanian and the uppermost Albian. Proc. Geol. Assoc. 37: 420-432. SPATH, L. F. 1927. Revision of the Jurassic cephalopod fauna of Kutch. Mem. geol. Surv. India Palaeont. indica n.s. 9: 1-84. SPATH, L. F. 1931. A monograph of the Ammonoidea of the Gault. Palaeontogr. Soc. (Monogr.): 313-378. SPATH, L. F. 1937. The nomenclature of some Lower Chalk ammonites. Geol. Mag. 74: 277-281. STANTON, T. W. 1893. The Colorado Formation and its invertebrate fauna. Bull. U.S. geol. Surv. 106: 1-288. STEPHENSON, L. W. 1952. Larger invertebrate fossils of the Woodbine Formation (Cenomanian) of Texas. Prof. Pap. U.S. geol. Sury. 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: 56-67. 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. TAUBENHAUS, H. 1920. Die Ammoneen der Kreideformation Palastinas und syriens. Z. dt. PalastVer. 43: 1-58. THOMEL, G. 1969. Reflexions sur les genres Eucalycoceras et Protacanthoceras (Ammonoidea). C.R. hebd. Séanc. Acad. Sci., Paris 268: 649-652. THOMEL, G. 1972. Les Acanthoceratidae Cenomaniens des chaines supalpines Meridionales. Mem. Soc. géol. Fr. n.s. 51: 1-204. THOMEL, G. 1973. A propos de la limite entre les étages Cenomanien et Turonien. C.R. hebd. Séanc. Acad. Sci., Paris 277: 761-764. UsHer, J. L. 1952. Ammonite faunas of the Upper Cretaceous rocks of Vancouver Island, British Columbia. Bull. geol. Surv. Can. 21: 1-182. VENZO, S. 1936. Cefalopodi del Cretacea medio-superiore dello Zululand. Palaeontogr. ital. 36: 59-133. WARREN, P. S. 1930. New species of fossils from Smokey River and Dunvegan Formations, Alberta. Rep. Alberta Res. Council geol. Sury. 21: 57-68. WARREN, P. S. 1947. Cretaceous fossil horizons in the Mackenzie River Valley (Canada). J. Paleont. 21: 118-123. Wuite, C. A. 1876. Report upon the invertebrate fossils, collected in portions of Nevada, Utah, New Mexico and Arizona, by parties of the expedition of 1871, 1872 and 1873, with descriptions of new species. Rep. U.S. geogr. geol. Explor. Surveys W. 100th Mer. 4: 1-219. WHITEAVES, J. F. 1884. On the fossils of the coal-bearing deposits of the Queen Charlotte Islands collected by Dr. G. M. Dawson in 1878. Geol. Surv. Can., Mesozoic fossils 1: 191-262. WHITEAVES, J. F. 1900. On some additional or imperfectly understood fossils from the Creta- ceous rocks of Queen Charlotte Islands, with a revised list of the species from these rocks. Geol. Sury. Can., Mesozoic Fossils 1: 263-307. 52 ANNALS OF THE SOUTH AFRICAN MUSEUM WIEDMANN, J. 1959. Le Crétacé supérieur de l’Espagne et du Portugal et ses Céphalopodes. In: Colloque Crétacé supérieur francais. Soc. Savantes Paris, 84th Cong., Dijon, Sec. Sci., Comptes Rendus Colloque Crétacé: 709-764. WIEDMANN, J. 1962. Ammoniten aus der vascogotischen Kreide (Nordspanien). I. Phyllocera- tina, Lytoceratina. Palaeontographica A118: 119-237. WIEDMANN, J. 1964. Le Crétacé Supérieur de l’Espagne et du Portugal et ses céphalopodes. Estud. geol. Inst. Invest. geol. Lucas Mallada: 107-148. WIEDMANN, J. 1973. The Albian and Cenomanian Tetragonitidae (Cretaceous Ammonoidea), with special reference to the circum-Indic species. Eclogae geol. Helv. 66: 585-616. WIEDMANN, J. & DIENI, I. 1968. Die Kreide Sardiniens und ihre Cephalopoden. Palaeontogr. ital. n.s. 34: 1-171. Woops, H. 1911. The palaeontology of the Upper Cretaceous deposits of northern Nigeria (Appendix). Jn: FALCONER, J. D. The geology and geography of northern Nigeria. London: Macmillan. WRIGHT, C. W. 1963. Cretaceous ammonites from Bathurst Island, northern Australia. Palaeontology 6: 597-614. WRIGHT, C. W. & Matsumoto, T. 1954. Some doubtful Cretaceous ammonite genera from Japan and Saghalien. Mem. Fac. Sci. Kyushu Univ. (D) Geol. 4: 107-134. WRIGHT, C. W. & WRIGHT, E. V. 1951. A survey of the fossil Cephalopoda of the Chalk of Great Britain. Palaeont. Soc. (Monogr.): 1-40. YABE, H. 1902. Cretaceous Cephalopoda from Hokkaido. J. Coll. Sci. Imp. Univ. Tokyo 20: 1-45. YOuNG, K. 1957. Cretaceous ammonites from eastern Apache County, Arizona. J. Paleont. 31: 1167-1174. Youna, K. 1958. Cenomanian (Cretaceous) ammonites from Trans-Pecos Texas. J. Paleont. 32: 286-294. 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. 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Spawning behaviour, ese masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51. THIELE, J. 1910. Mollusca: B. Peircincaouees 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 75 Band March 1978 Maart Part 6 Deel NEW RECORDS OF CRABEATER SEALS (LOBODON CARCINOPHAGUS) FROM SOUTH AFRICA By GRAHAM J. B. ROSS PETER D. SHAUGHNESSY & PETER B. BEST 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.), 7-4), 8, 911-2, 7), 10(1-3), 11(1-2, 5, 7, t.—p.i.), 15(4-5), 24(2), 27, 311-3), 32(5), 33 Price of this part/Prys van hierdie deel R1,60 ISBN 0 908407 40 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 NEW RECORDS OF CRABEATER SEALS (LOBODON CARCINOPHAGUS) FROM SOUTH AFRICA By GRAHAM J. B. Ross Port Elizabeth Museum, Port Elizabeth PETER D. SHAUGHNESSY & PETER B. BEST Sea Fisheries Branch, Cape Town (With | figure and 1 table) [MS. accepted 5 January 1978] ABSTRACT Information is provided on nine records of the crabeater seal in South Africa since 1968 and one previously recorded stranding in 1957. They occurred on the south and east coasts between False Bay and East London. Most of the strandings occurred during summer months, and most were of seals in their first year. It is suggested that these seals come from the western Atlantic sector of the Antarctic pack-ice, south-west of South Africa. CONTENTS PAGE Introduction . : : : 153 Records . : : ; : 154 Discussion ; : 5 A 156 Acknowledgements . : sey eS, References ; 4 5 : 158 INTRODUCTION The crabeater seal Lobodon carcinophagus is a pelagic species with a circum- polar distribution in antarctic waters associated with the drifting pack-ice. On several occasions, however, vagrants have been recorded from islands in the Southern Ocean, Australia, Tasmania, New Zealand, South Africa and the Atlantic coast of South America. These records have been mapped by Erickson & Hofman (1974) as far north as latitude 30°S, though the locality of the northernmost record is Pontal, Rio de Janeiro, at latitude 23°S (Vaz Ferreira 1965). Since Courtenay-Latimer (1961) reported the first South African specimen of a crabeater seal, nine more animals have been recorded on this coast. Details of all these records are reported here and their localities shown in Figure 1. IDs Ann. S. Afr. Mus. 75 (6), 1978: 153-158, 1 fig., 1 table. 154 ANNALS OF THE SOUTH AFRICAN MUSEUM SOUTH AFRICA Gamtoos R. Van Staden’s R. 35°S 20° E 25° 30° Fig. 1. Map showing the localities of crabeater seal strandings in South Africa. RECORDS On 22 July 1957 a female seal hauled out on the beach at Hamburg (33°17’S 27°30’E) (Courtenay-Latimer 1961). The authors have determined the condylo- basal length (CBL) of the skull as 241 mm, and the standard length of the mounted specimen on display at the East London Museum (ELM 572) as 1,69 m. On 17 March 1968 the decomposed carcass of a crabeater seal was collected between the Gamtoos and Van Staden’s River mouths (33°57’S 25°08’E). The skeleton was collected for the Port Elizabeth Museum (PEM 1513/101), but the skull has subsequently disappeared. The seal was not measured, but comparison of the length of its articulated vertebral column with that of another specimen (PEM 1519/25; 1,23 m) suggested that the standard length of this animal was about 1,9 m. A male crabeater seal hauled out at The Strand, False Bay (34°07’S 18°50’E) on 26 December 1971, and was killed by bystanders the following morning. External measurements of this animal (and ancther three animals that were measured in a standard manner) are provided in Table 1. Each testis weighed 7,5 g, and measured 45 x 17 mm and 40 x 18 mm. The skin and skeleton are in the collections of the South African Museum, Cape Town (SAM-36357). The viscera and eyes are in the collections of the Sea Fisheries Branch. The stomach of this seal contained approximately 150 g of gravel and sand, and 15 g of algae identified by R. H. Simons (Sea Fisheries Branch) as Dictyota intricata, Ploca- NEW RECORDS OF CRABEATER SEALS FROM SOUTH AFRICA 155 TABLE | Measurements of four crabeater seals that stranded on the coast of South Africa Flipper length Standard Nov it. Axillary Thickness? Skull Specimen Sex Mass length front hind girth blubberand CBL number (kg) (m) (m) (m) (m) skin(mm) (mm) SAM-36357 M 77,3 1,66 0533) 0332 1,01 24 fragments SAM-36358 F 99,1 lv 0,33 0,33 1,10 19 248 3PEM 1518/42 F 64 1,67 0,33 0,33 _ — 247 83PEM 1519/25 + F 148 2,08 — — -- — 2 1 Measured from anterior insertion of flipper to tip of first claw. 2 Measured at the posterior end of the sternum. 3 Measured at time of death. On 6 February 1974 PEM 1519/25 measured 1,88 m in standard length. mium corallorhiza and P. rigidum, all of which are common sublittorally in False Bay. On 19 January 1972 a female crabeater seal came ashore at Kalk Bay (34°07’'S 18°27’E) and was euthanized. External measurements are provided in Table 1. The skin and skeleton are in the collections of the South African Museum (SAM-36358). The stomach contained 30 small stones with a total mass of 5 g and a large number of tiny stones, sand and shells totalling 170 g. The shells have been identified by B. Kensley (South African Museum) as Oxystele variegata, Turritella sp. and Burnupena sp., all of which are common in South African waters. A female crabeater seal was collected alive on 28 January 1973 at Jeffreys Bay (34°04'S 24°56’E) and taken to the Port Elizabeth Oceanarium where it was maintained on a fish diet (Ross et al. 1976). The seal was measured (Table 1) when it died on 25 April 1973 in an emaciated condition. The skull and ovaries are preserved in the Port Elizabeth Museum (PEM 1518/42). On 26 January 1974 a female crabeater seal was collected at Seaview (34°03’S 25°30’E), near Port Elizabeth. It was also kept at the Port Elizabeth Oceanarium (Ross e¢ al. 1976). It died on 15 April 1974 when it was measured (Table 1). The complete skeleton and ovaries are preserved in the Port Elizabeth Museum (PEM 1519/25). A crabeater seal died on 15 April 1974 some 36 hours after it had been first reported at Nature’s Valley (33°58’S 23°33’E). The total length including the hind flippers was 2,11 m and the maximum girth round the foreflippers and chest was 1,06 m. The seal had a mass of 85,2 kg. The fragmented skull and some vertebrae are preserved in the collections of the Tsitsikama Coastal National Park (TNP/B/1974/1). Comparison with the measurements of the captive animal that stranded at Seaview in 1974, which on 6 February 1974 had a standard length of 1,88 m and a total length including hind flippers of 2,10 m, suggests that the standard length of this seal was approximately 1,9 m. A crabeater seal, thought to be a female, hauled out at Igoda (33°06’S 27°47'E), 17 km south-west of East London, on 11 January 1975. It survived In captivity for two days. The mounted skin with skull inside has been prepared 156 ANNALS OF THE SOUTH AFRICAN MUSEUM for display at the East London Museum (ELM 916). The standard length of the mounted animal is 1,68 m. On 26 January 1975 a crabeater seal of unknown length and sex hauled out at Jeffreys Bay and died after several hours. Colour photographs in the files of the Port Elizabeth Museum show that the animal was unscarred. It was estimated by observers and from the photographs to be less than 2 m in standard length. No parts of the carcass were saved. On 28 January 1975 a crabeater seal was found on Leisure Island, in Knysna Lagoon (34°05’S 23°03’E) where it remained for the day before swimming away in the late afternoon. The animal was approximately 2 m in total length and unscarred (M. J. Clarke in litt.). Two photographs of it are on file in the Port Elizabeth Museum. DISCUSSION Though few observations have been made of new-born crabeater seals, available records indicate that pups are born between the middle of September and early November at approximately 1,35—1,5 m in standard length (King 1957; @ritsland 1970b; Corner 1972). From an analysis of the standard length frequen- cies of 292 crabeater seals collected in the Antarctic between the months of January and April, Laws (1958) concluded that peaks of 1,88 m and 2,03 m represented the modal lengths of seals aged 4 year and 14 years, respectively. Further, sexual maturity in females is attained at a standard length of 2,06 m and physical maturity in both sexes at 2,26 m. By comparison, the standard lengths of eight of the South African specimens for which reliable data are available (ranging from 1,66 to 1,9 m) indicate they were in their first year of life. Because the other two animals were not measured, their age-class cannot be determined, but it seems likely that they were considerably smaller than physically mature animals. Laws (1958) determined the ages of crabeater seals from the number of dentine layers seen in a transverse section of a tooth when viewed by reflected light in water. Sections of upper canines of PEM 1518/42, SAM —36357 and SAM-36358, and lower canines of ELM 572 and PEM 1519/25 all showed a single, incomplete postnatal dentine layer, confirming that these five animals were less than | year old. The origin of these seals is unknown. It is possible that they came from the nearest pack-ice some 2 000 km to the south of South Africa, though the West Wind Drift would have tended to drift the animals eastwards as they moved northwards from this sector of the Antarctic. It seems more likely that they came from west of the longitude of South Africa, possibly from the western Atlantic sector of the Antarctic, where crabeater seals are particularly abundant (Erickson & Hofman 1974). There the northern part of the pack-ice lies within the influence of the West Wind Drift, which would assist seals in their passage. Even so, the current could provide a drift of only some 2000 km over a period of four - NEW RECORDS OF CRABEATER SEALS FROM SOUTH AFRICA 157 months, for its speed is about 0,7 km/h (Hydrographic Department 1961). Thus it seems that a recently weaned crabeater seal moving from the pack-ice to South African waters would have to swim actively in order to complete the journey in three or four months. The South African records are markedly seasonal, with eight of them occurring between late December and early March, while six of these occur in January alone. The seasonal occurrence in the summer months contrasts with the statement by Scheffer (1958) that crabeater seals move outward from Antarctica in autumn. It also contrasts with the seasonal occurrence of records in Australia, where only two of the six animals hauled out during summer (one each in December and January) and the other four hauled out between June and September (Anon. 1946; Hall 1903; Ingham 1960; Le Soeuf 1929; Troughton 1965). The only dated strandings of crabeater seals in the available literature for New Zealand and South America were for the months of April (Oliver 1921) and June (Berg 1898), respectively. By comparison it is interesting to note that a seasonal influx of crabeater seal pups was observed in December and January at the Bay of Whales in Antarctica by Lindsey (1938), and several other authors have recorded the Seasonal movement of crabeater seals towards the continent in midsummer (see @Mritsland 1970a), though these seals are not always young animals (Bertram 1940). The timing of these two movements suggests that they are related: as seals begin to move southward with the retreat of the pack-ice in summer, an unknown proportion moves northward and eastward with the West Wind Drift. The few South African records must be a small proportion of the number of crabeater seals reaching these waters, for the geographical distribution of the present records reflects that of interested biologists. Further, as South Africa lies on the northern edge of the influence of the West Wind Drift, those animals reaching South Africa can be only a fraction of those that do not land, but pass to the south. It is likely that few of these animals survive. The presence of stones, sand and other indigestible matter in the stomachs of two animals suggests that the seals were unable to fend for themselves in South African waters. They may con- stitute a small but potentially significant part of total pup mortality. Studies of the movements of pups using radio telemetry during the break-up of the pack-ice to determine whether pup dispersal occurs randomly or in a fixed direction would assist in assessing the potential size of emigration from the population to the north and the east. ACKNOWLEDGEMENTS For permission to examine and publish data on specimens in their collec- tions, we thank the Directors of the East London, Port Elizabeth and South African Museums and the Warden of the Tsitsikama Coastal National Park. 158 ANNALS OF THE SOUTH AFRICAN MUSEUM We also thank Dr B. Kensley and Mr R. H. Simons for identifying stomach contents of seals. REFERENCES ANoNn. 1946. [Lobodon carcinophagus in Tasmania.] Pap. Proc. R. Soc. Tasm. 1945: 165. BERG, C. 1898. Lobodon carcinophagus (H.J.) Gr. en el Rio de la Plata. Comun. Mus. nac. B. Aires 1: 15. BERTRAM, G. C. L. 1940. The biology of the Weddell and crabeater seals with a study of the comparative behaviour of the Pinnipedia. Scient. Rep. Br. Graham Ld Exped. 1: 1-139. CorRNER, R. W. M. 1972. Observations-on a small crabeater seal breeding group. Bull. Br. Antarct. Surv. 30: 104-106. COURTENAY-LATIMER, M. 1961. Two rare seal records for South Africa. Ann. Cape Prov. Mus. (nat. Hist.) 1: 102. ERIcKSON, A. W. & HOFMAN, R. J. 1974. Antarctic seals. Am. Geogr! Soc., Antarct. Map Folio Ser. 18: 4-13. Hatt, T. S. 1903. [Crabeater seals in Australian waters.] Nature, Lond. 67: 327-328. HYDROGRAPHIC DEPARTMENT. 1961. The Antarctic pilot. 3rd ed. London: Her Majesty’s Stationery Office. INGHAM, S. E. 1960. The status of seals (Pinnipedia) at Australian Antarctic stations. Mammalia 24: 422-430. Kina, J. E. 1957. On a pup of the crabeater seal Lobodon carcinophagus. Ann. Mag. nat. Hist. (12) 10: 619-624. Laws, R. M. 1958. Growth rates and ages of crabeater seals, Lobodon carcinophagus Jacquinot & Pucheran. Proc. zool. Soc. Lond. 130: 275-288. Le Souer, A. S. 1929. Occurrence of the crab-eating seal Lobodon carcinophaga Hombron and Jacuinot [sic], in New South Wales. Aust. Zool. 6: 99. Linpsey, A. A. 1938. Notes on the crab-eater seal. J. Mammal. 19: 456-461. OLIveER, W. R. B. 1921. The crab-eating seal in New Zealand. Trans. Proc. N.Z. Inst. 53: 360, pl. 56. QMRITSLAND, T. 1970a. Biology and population dynamics of Antarctic seals. In: HOLDGATE, M. W., ed. Antarctic ecology 1: 361-366. London: Academic Press. QMRITSLAND, T. 19705. Sealing and seal research in the south-west Atlantic pack ice, Sept.—Oct., 1964. In: HoLpGaTE, M. W., ed. Antarctic ecology 1: 367-376. London: Academic Press. Ross, G. J. B., RYAN, F., SAAYMAN, G. S. & SKINNER, J. 1976. Observations on two captive crabeater seals Lobodon carcinophagus at the Port Elizabeth Oceanarium. Int. Zoo Yb. 16: 160-164. SCHEFFER, V. B. 1958. Seals, sea lions, and walruses. A review of the Pinnipedia. Stanford, California: Stanford University Press. TROUGHTON, E. 1965. Furred animals of Australia. 8th ed. Sydney: Angus & Robertson. VAZ FERREIRA, R. 1965. Ecologia terrestre y marina de los pinnipedios del Atlantico sudocci- dental. Anais Acad. bras. Cienc. 37 (Suppl.): 179-191. (In Spanish, English summary.) 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. GRAHAM J. B. ROSS PETER D. SHAUGHNESSY & PETER B. BEST NEW RECORDS OF CRABEATER SEALS (LOBODON CARCINOPHAGUS) FROM SOUTH AFRICA OLUME 75 PART 7 JUNE 1978 ISSN 0303-2515 07:63 3 ace 4 j rs vy a ANNALS OF THE SOUTH AFRICAN ee ae AMIEL IVE CAPE TOWN INSTRUCTIONS TO AUTHORS 1. MATERIAL should be original and not published elsewhere, in whole or in part. 2. LAYOUT should be as follows: (a) Centred masthead to consist of Title: informative but concise, without abbreviations and not including the names of new genera or species Author’s(s’) name(s) Address(es) of author(s) (institution where work was carried out) Number of illustrations (figures, enumerated maps and tables, in this order) (b) Abstract of not more than 200 words, intelligible to the reader without reference to the text (c) Table of contents giving hierarchy of headings and subheadings (d) Introduction (e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents (f) Summary, if paper is lengthy (g) Acknowledgements (h) References (i) Abbreviations, where these are numerous 3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced with 2,5 cm margins all round. First lines of paragraphs should be indented. Tables and a list of legends for illustrations should be typed separately, their positions indicated in the text. All pages should be numbered consecutively. Major headings of the paper are centred capitals; first subheadings are shouldered small capitals; second subheadings are shouldered italics; third subheadings are indented, shouldered italics. Further subdivisions should be avoided, as also enumeration (never roman numerals) of headings and abbreviations. Footnotes should be avoided unless they are short and essential. Only generic and specific names should be underlined to indicate italics; all other marking up should be left to editor and publisher. 4. ILLUSTRATIONS should be reducible to a size not exceeding 12 x 18 cm (19 cm including legend); the reduction or enlargement required should be indicated; originals larger than 35 x 47 cm should not be submitted; photographs should be rectangular in shape and final size. 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REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: ‘Smith (1969) describes. . ‘Smith (1969: 36, fig. 16) describes...’ “As described (Smith 1969a, 19695; 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, 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. FIsCHER, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. gén. 74: 627-634. \ Koun, A. J. 19604. 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) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 75 Band June 1978 Junie Part a Deel SOUTHERN AFRICAN CUMACEA EIT 2 FAMILY BODOTRIIDAE, SUBFAMILY BODOTRIINAE By JENNIFER DAY 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 45 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 SOUTHERN AFRICAN CUMACEA PART 2 FAMILY BODOTRIIDAE, SUBFAMILY BODOTRIINAE By JENNIFER DAY Zoology Department, University of Cape Town (With 48 figures and 3 tables) [MS. accepted 18 January 1978] ABSTRACT The Bodotriinae in southern Africa are represented by 34 species in 9 genera. 31 of these species are described and figured. 3 of the genera (A/ticuma, Austrocuma and Mossambicuma) are new, as are 16 of the species (Eocuma foveolatum, E. winri, E. aculeatum, Cyclaspoides pellucidus, Mossambicuma elongatum, Austrocuma platyceps, Alticuma bellum, Iphinoe producta, Cyclaspis scissa, C. australora, Bodotria clara, B. nitida, B. tenuis, B. falsinus, B. serica and B. vertebrata). B. vertebrata is divided into two subspecies. The females of J. dayi and Iphinoe ? zimmeri are described for the first time, as are the males of A. carinata, C. spectabilis, B. magna and B. montagui. Alticuma carinatum and Iphinoe capensis are allocated to different genera and Iphinoe ? zimmeri is considered to be incertae sedis. Keys are given to the genera of the Bodotriinae, the southern African species in the sub- family, the world species of Eocuma and the African and European species of Iphinoe and Bodotria. The general distribution of the Bodotriinae is discussed and a more detailed account is given of the subfamily in southern African waters. It is concluded that the genera Bodotria and Iphinoe are the most successful in this region, contributing more than half of the species and 93 per cent of the individuals. The Bodotriidae in general and the Bodotriinae in particular are the most successful of the cumacean families in these waters. CONTENTS PAGE Introduction . : : f : , 160 Material and station data ; : 5 : 160 Methods : : , : : : § GO Systematics . : : : : 161 Key to the genera of Bodotrinae | 4 : 64 Key to the southern African Bodotriinae : = 166 Eocuma . : : : ; : 3 ‘ 168 Upselaspis : : ; : , 5 Se) Cyclaspoides . ; : : ‘ . 184 Mossambicuma gen. nov. . : : : . 188 Austrocuma gen. nov. ‘ ; : : : 192 Alticuma gen. nov. . : : : : > Is Iphinoe . : ; : : : : 208 Cyclaspis : : 5 : : ‘ 5 ZS Bodotria ; ; , ‘ . 246 Distribution of the Bodotriinae ‘ . 284 Distribution of the southern African Bedorinae 5 RSS) Acknowledgements : : : : ; 5 Peis) References . : , : ; : : 5 RSS) 159 Ann. S. Afr. Mus. 75 (7), 1978: 159-290, 48 figs, 3 tables. 160 ANNALS OF THE SOUTH AFRICAN MUSEUM INTRODUCTION This is the second in a series of papers on the Cumacea of Africa south of 20°S. The reader is referred to the first in the series (Day 1975) for a discussion of the structure and terminology of Cumacea in general, as well as a report on the taxonomy and distribution of the subfamily Vaunthompsoniinae in these waters. There has been little previous work on the southern African Bodotriinae, and the cumacean fauna of the region is generally poorly known. The few earlier descriptions are to be found in: Zimmer’s (1908) paper on the material from the Deutsches Tiefsee-Expedition and his report on the collection in the Berlin Zoologisches Museum (Zimmer 1921); two papers by Stebbing (1910, 1912); Fage’s (1951) report on material from the Belgian Oceanographic Expedition; Jones’s (1956) report on material from the Atlantide and Galathea Expeditions, and two papers on material collected by the Zoology Department of the University of Cape Town (Hale 1953; Jones 1960). MATERIAL AND STATION DATA The vast majority of samples used in this study was collected by the Zoology Department of the University of Cape Town (UCT) during a benthic survey round the South African coast, the programme being funded by the Oceanographic Research Institute of the Council for Scientific and Industrial Research (CSIR) and headed by J. H. Day. Other material was obtained from: the South African Museum, mostly collected by the S.S. Pieter Faure in 1898— 1907 and the R.V. Meiring Naude in 1976-1977; the National Institute for Water Research of the CSIR; the Sea Fisheries Branch in Cape Town; a survey of Richards Bay conducted by the Port Elizabeth Museum; a survey of Lake St Lucia conducted by the Zoology Department of Rhodes University. Due to the numerous samples and sources of material it is unpractical to list exact station data for each species. Thus in the distribution records the area of collection and the institution providing the material are designated by code letters and only extremities of range and depth are given. Table | lists the code letters and their geographical positions. These are also shown graphically in Figure 1. METHODS Collections: estuarine material was collected by means of plankton nets of various kinds and most benthic samples by grabs or dredges. A few of the UCT samples from the shallower stations at Lambert’s Bay, Saldanha Bay, Still Bay and Langebaan Lagoon were obtained by means of a diver-operated suction-sampling device. Length measurements were taken from the anterior tip of the carapace to the posterior edge of the telsonic somite. Exhalant siphons and uropods were excluded in every case. SOUTHERN AFRICAN CUMACEA: PART 2 161 TABLE | Code letters of the survey programmes and their geographical ranges. Institute Area Explanation Geographical position UCT University of Cape Town SWD_ South West Africa benthic survey Cape Cross (21°S 13°E) to Orange River Mouth (28°S 16°E) WCD West coast benthic survey Orange River Mouth (28°S 16°E) to Cape Agulhas (34°S 20°E) LBT Lambert’s Bay benthic transect Lambert’s Bay, shore to 800 m (32°S 18°E) SB Saldanha Bay benthic survey Saldanha Bay (32°S 17°E) LB Langebaan Lagoon benthic survey Langebaan Lagoon (33°S 18°E) FAL False Bay benthic survey False Bay (34°S 18°E) FBY False Bay benthic transect False Bay, shore to 84 m (34°S 18°E) SST Still Bay transect Still Bay, shore to 200 m (34°S 21°E) SCD South coast benthic survey Cape Agulhas (34°S 20°E) to Natal border (31°S 30°E) KNY Knysna estuarine survey 34°S 23°E (plankton) CP Cape Peninsula shore survey 34°S 18°E CPR _ Cape Province shore survey Orange River Mouth (28°S 16°E) to Umtamvuna River Mouth (31°S 30°E) NIWR National Institute for Water Research COD benthiccoastal survey near Durban 30°S 30°E BLL grid of benthicstations off Durban 29°S 31°E CON Morrumbene estuarine survey Morrumbene estuary, Mozambique (23°S 35°E: plankton) SAM South African Museum various (see text for details) RU Rhodes University (Zoology Dept.) Lake St Lucia, Zululand (28°S 32°E: plankton) PEM Port Elizabeth Museum Richards Bay, Zululand (28°S 32°E: plankton) FISH Sea Fisheries Branch South West Africa (20°S 12°E: plankton) BMNH British Museum (Natural History) Hermanus, shore (34°S 19°E) SYSTEMATICS Family Bodotriidae Scott, 1901 Diagnosis No free telson. Pleopods (males only) with an outer process to the inner ramus—usually five pairs, but may be two or three. Mandibles narrow at base. Endopod of uropod 1- or 2-segmented. Branchial apparatus without gill-plates or supports. Remarks The family was divided by Hale (19445) into two subfamilies according to the number of thoracic limbs bearing exopods. The Vaunthompsoniinae are characterized by having exopods on pereiopods other than the first pair and the southern African representatives of this subfamily were dealt with in the first paper in this series (Day 1975). ANNALS OF THE SOUTH AFRICAN MUSEUM 162 ‘ode UIO}SOM-YINOS :4OSUT “SBOIL POD JO SIOPIOG OYVOIPUI SOUT] o|QnOC *s19}}0] Opodo SuIpuodsoar10S pue svore 3UT}D9];09 (GOD 8 118) ueqing £ (Wad) 40g SP10>4>!4 (Nd) DIIN] IS sonbupyw ~os5uainoy _eUsqUIN dO Ww :$.07 JO YINOS voLIpY UJOYyINOY ‘| “BLy 4.9¢ 4.0¢ YI2@qQDZI}9 14046 (15S) Avg JIS (AN) Puskuy UOPpUO] JsDgz O (181) ADg sysequin] © 4, Zl (dD) bjNsuluag adp> Zjuapny Q (a1) uoobp] unpqe6un7—~ Dg SIAIDM (as) Avg DYUDPIDS —— ssOuy adn ° <2 EN NE OO eA ae SOUTHERN AFRICAN CUMACEA: PART 2 163 Subfamily Bodotriinae Hale, 1944 Diagnosis Bodotriidae with exopods only on the third maxilliped and first pereiopod in both sexes. Remarks The genera of the Bodotriinae form a fairly homogeneous and apparently natural group. The subfamily up to now has consisted of eight genera, all related morphologically and similar in general appearance. Stephanomma is a monotypic genus, known only from a single specimen in the West Indies, characterized by the absence of pseudorostral lobes. Zygosiphon, also mono- typic, is known from a few specimens from Indo-China and is characterized by the pseudorostral lobes being produced laterally to accommodate the widely divergent branchial siphons. Representatives of all the other genera occur in the present collection, as well as specimens which do not fit into any known genus. Since generic boundaries are already tenuous in most cases, it is felt that less confusion will result from the erection of new genera than from the expansion of the definitions of existing ones. A small number of individuals from Morrumbene estuary in Mozambique are in some respects similar to Eocuma, while in others they are quite distinct. They have been placed, rather hesitantly, in a new genus, Mossambicuma. There are also some specimens of a single species in which the male has only three pairs of pleopods. Since this character is unique in the subfamily they too must be separated, and are placed in the new genus Austrocuma. Representatives of Cyclaspis carinata Zimmer, 1921 occur in the collection. Further examination shows them to be quite atypical for the genus: the second pereiopod is 7-segmented, while the first pedigerous somite is quite clearly visible in both sexes and the endopod of the uropod is 2-segmented. A new genus, Alticuma, has therefore been created for them and for a second similar species found in deep water in the southern Mozambique Channel. The Bodotria—Iphinoe-Cyclaspis group consists of a continuum of species which are separated into three genera on the basis of the number of free pedige- rous somites and of segments of pereiopod 2 and of the endopod of the uropod. Thus Iphinoe has five free pedigerous somites, the second pereiopod is 6-segmented and the endopod of the uropod 2-segmented. Cyclaspis has four visible pedigerous somites, the second pereiopod is 7-segmented and the endopod of the uropod l-segmented. Bodotria has four visible pedigerous somites, the second pereiopod is 6-segmented and the endopod of the uropod 1- or 2-segmented. But generic distinctions are not absolute, for in the females of some species of Cyclaspis the first pedigerous somite may be visible and in Bodotria the fusion of the ischium of pereiopod 2 with the basis is not always complete, so that it is sometimes difficult to say with certainty whether the limb is 6- or 7-segmented. ANNALS OF THE SOUTH AFRICAN MUSEUM 164 A][@19}"] JSvoy ye soedeivS Jopun 9}1WOs snosesiped pany} BUIIvS [eSIOPpIW YIM Pessoiduros Aj[v19ye] sovdevavo SUuO] ATOAT}E[OI DDedeIKO “j10ys usuopqe suo] Joy10ys yonur Jo}10ys (Gua | 9 PP SUOS UT ATUO AT[eI0}RT SIGISIA “S Ul sovdeieo rspun ou saplodspjac9a S9qO] [v1jso1opnosd ou suou ouou 9} e1opow J10ys Jasuoy| [enboqns 4 9 © Ul oovdvied Jopun 1O MOJIeU AIOA ‘4 UI OPI ou Sidspjasdy (panuyjuoD) seunnopog 9} JO B19U93 OY} JO s1o}OVIeYO OsOusvIq *7 ATV], ouou OS AJoJe19poUur IO 97eUIO ouou 9} ¥1opour ou0ou Josuo] yonut Josuoy yonur L o7e19pour Sok Duuouvnydajs Ayysry ‘ureyd — opquizea 9[QeIvA AIOA }A0YS AIDA 0} yOYsS O[Quiiea g]quiiea L MOLICU (66 *SIAO ouIOS ul }d99x9) ou sidspjacy 9PIM AIOA Jey 10ys A][¥iSIp Julod v 0} psonpold | podotoiod jo stseq suIOY [e107e] YM JO/pue pousyjey A][e1yUsAOSIOp ATjensn 9}eSuOClO 9}P.1OPOUL 0} 7104S Joy10Ys yonur JoJ1oYys yonur 9 (s0"deie9 YIM osny ABU) MOIIeU ou pUuNndoy spodogjd jo sired 9914} YIM P JUOSIOAIP ATOPIM suoydis jerysueiq auou 9uoUu oyeIOpoU yAoYs AOA JUSSIOAIP “Buoy Jesuo] yonur Josuo] [enbo Josuoy I 6 9 L (&) Opi AIOA 0} (2) o7e10pour 97e1OpoUr Ajuo & utr SoA “AOU “U83 DUUNIOAISNY uoydisosd 7 9su0ou 9BUTIvO [¥19}¥] JO sired S10U IO 9UO YIM UaIyzo qnO}s OF 9}e1OpoUulL WOYsS 419A 0} 0Ys Jesuo] yonur Josuo] ct 9 & Ul OpIM AIOA “P Ul 9}¥10pour ou DIAJOopog ouou O[QUIIvA 9}819POU 0} JOpudTs 9}.1OPOW 0} J10Ys Iasuo] Jasuoy] (4 L 97e.10pour SoA ‘AOU "W903 DIUNoIY py SeUNNOPOY oY} JO viouss 94} Jo siajovIKYO oNsousvIq “7 ATAV I, on]eA S1jsouseIp suou JO soinjzesy I0yj}0 ouou UuOol}e}JUSWIeUIO o}esu0O wa0j Apoq soqoy| 9}e1opour [®1jsor1opnesd Jo y3uey S}UIOS SIUOS]oz 0} UOl}v]Jo1 UT podoin JO 9[sunped jo yysue] TWeI 0} UOT}e[O1 UI podoin JO 9founped jo yI3uey] podoin jo podopus jo I SyusWIdes JO Joquinu 7 podotsied jo 9 sjuouses JO Joquinu OUIOS sno1osIpod Puodes JO YIpPIM E9QISIA O}1TUIOS ou SnoJosiped ysi1q [enboqns Jo 19}10y4s Jov0Ys o71eIopow “AOU "Uds DUNIIGuDSSOp x9[dur09 Ajyuenbery © JO soyui9}s SIOVIOY} [VIQUSA OJVIIOS UDO ‘eure On[VA o1souseIp JO Soinjzesy 19q10 [estopprur AyyUenbaary UOT}e}USWIeUIO Jopuoys wo} Apoq soqoy] SUO] 0} 91¥10pour [®1jso1opnesd jo yysuey 9}TWIOS STUOS]O} 0} WOT}ejo1 UL podoin JO 9jounped jo yy3u9y TMI O} UOT} ¥91 UT podoin JO spounped jo y)8u9] podoin jo podopuo jo Z S}UOWUd9S JO JoquInu Z podotesiod yo 9 S}U9WUS9s JO JIoquINnU (d) opr 0} 9}IWOS sno1osipod (2) 271e.10pour Puoses JO YIpPIM é9[QISIA O}1WWIOS Sok SNOJOSIpod JsIny aoulydy IOSUuo] Iasuo] 10 [enbo SOUTHERN AFRICAN CUMACEA: PART 2 165 However, the species of both Iphinoe and Bodotria are generally rather characteristic of their genera, those of Jphinoe usually being slender and elongate with long pseudorostral lobes and of Bodotria being more compact and frequently bearing one or more pairs of lateral carinae and very short pseudorostral lobes. One of the species occurring in considerable numbers in the present collection was named Jphinoe zimmeri by Stebbing (1910) on the basis of a single adult male. Although in external appearance and in details of the limbs it quite clearly belongs to Iphinoe, since the first pedigerous somite is invisible it should by rights be placed in Bodotria, which is plainly unsatis- factory. It is therefore considered to be genus incertus and is called ‘Iphinoe ? zimmer. The main problem in Bodotria seems to be less a matter of generic than of specific boundaries. Many of the numerous species are very similar to one another and may well prove to be genetic morphs rather than valid species. Cyclaspis is a very variable genus, the carapace of some species being highly ornamented and of others quite unadorned, while the proportions of the segments of the uropods to the telsonic somite vary far more than they do in other genera. Added to this, Cyclaspis has a large number of species (about 80) and could profitably be split into two or more genera of roughly equal numbers. There does not, however, seem to be any satisfactory means of doing so since the ornamentation, which is the most striking polymorphic feature, does not correspond uniformly with any other obvious distinguishing characters. Within Cyclaspis there is one small group of species (subgrandis, tasmanica, longicaudata, gigas and spectabilis) which are clearly related in a number of features, notably the unadorned, more or less spherical carapace, the long telsonic somite, the short peduncle of the uropods and the large basis of maxilliped 3. They also tend to be deep-water forms. However, these characters, which are quite distinctive in the species mentioned, are to be found to a lesser degree in some of the more typical species. As a result it would be necessary to examine representatives of a large number of species quite closely in order to decide whether, in fact, those mentioned above could satisfactorily be removed to a new genus. This could well be one of the more useful applications of numerical taxonomy. Table 2 (see page 164) lists the diagnostic characters of the genera of the Bodotriinae, which are also keyed below. KEY TO THE GENERA OF BODOTRIINAE 1 Second pereiopod 7-segmented (ischium not at all fused with basis)...................- 2 — Second pereiopod 6-segmented (ischium at least partly fused with basis)................ 4 DRINOMPSEUdOKOStraliODES se hue-chrnieac eh Bo es setae Kou mue eee Stephanomma Sars, 1871 —sescudorostralslobes widely diversemt. «2.2. so: . ac ss-ces seme Zygosiphon Calman, 1905 —sESeuadonostralulobessnonmalnantenorhyadirected....- 42 sue dee one aie ee eee ne ee 3 3 First pedigerous somite visible at least dorsally in male, dorsally and laterally in female; EHCOPOdEOMmUFO pod 2-seemented.. eon. ace oe eae Alticuma gen. nov. First pedigerous somite never visible in male, sometimes visible dorsally in ovigerous female send @poG onuropod |-segmented.s.....-....206656. 40a a: Cyclaspis Sars, 1865 166 ANNALS OF THE SOUTH AFRICAN MUSEUM 4 First pedigerous somite visible in both sexes; endopod of uropod 2-segmented; male with fivespairs'of pleopods.ck snc ots Oh os Oe ee eee Iphinoe Bate, 1856 — First pedigerous somite visible in female only; endopod of uropod 1-segmented; male With bhiiee ipalrs Ol PlEODOCSeryaeeeieier ei ere aera eT ee Austrocuma gen. nov. — First pedigerous somite visible in neither sex; endopod of uropod 1- or 2-segmented; male with-five pairs:of pleopods...: \. os4:..s5s2.des ose douse on atoe: oo eee 5 5 Gut coiled; pedigerous somites 2 and 3 incorporated under carapace at least dorsally in Oth: SEXES: ce Wics ier ores PS a ts Cyclaspoides Bonnier, 1896 — Gut straight; pedigerous somite 3 always free and visible in both sexes; pedigerous somite 2 visible in female, sometimes fused with or incorporated under carapace in male........ 6 6 Peduncle of uropods nommore thanthalflenethiot rani sano. eee nee 7 =. Peduncleoruropods equal im lengthito, orlonger thans ramicn een eeenereene 8 7 Third segment of antenna 1 longest; basis of pereiopod 1 not distally produced Mossambicuma gen. nov. — Third segment of antenna 1 no longer than first; basis of pereiopod 1 distally produced COVA DOMME eitisqodc tenes CBee Fe eee ee eT ne ee Eocuma Marcussen, 1894 8 Lateral carinae frequently present on carapace; second pedigerous somite always visible in both sexes (often as wide as deep in female, half as wide as deep in male); peduncle of ureoyoxoral sonore ny Ikoraveeir Won TENT. ooo codons bboebo obo OOO UOC ODO Bodotria Goodsir, 1843 — Carapace without lateral carinae; second pedigerous somite free in female (about half as wide as deep), incorporated under carapace in male or else very narrow; peduncle of uropod Subequaliim: lene thutomaniipes sana nen mre cree Upselaspis Jones, 1955 *Peduncle of uropod subequal in length to rami in Jphinoe ? zimmeri. KEY TO THE SOUTHERN AFRICAN BODOTRIINAE Taxonomically accurate keys to both genera and species are often difficult for the inexperienced to follow. Since such keys also frequently require exami- nation of adult animals of both sexes and details of appendages which may be missing, the following key is provided for convenience. It should be noted, however, that although this key will separate all species found to date in southern African waters, it will mot necessarily distinguish them from species in other areas. The key is based almost entirely on characters of those parts of the body that are least likely to be damaged or missing, so that although it can be used to identify damaged animals, it is not as rigorous as the keys to individual genera and species, which should always be consulted for final identification. 1 Carapace with one or more pairs of longitudinal ridges (carinae) or depressions lateral to midline (Figs 4A; 26A;'4 0A). baci Re eds te re ho eo a eee 2, — Carapace with paired carinae (Fig. 15A), horns (Fig. 5B) or depressions (Fig. 6A) lateral to midinesbut none Gunning lonsitudinallysanns meee ane nee 15 — Carapace without paired carinae, horns or depressions lateral to midline (Figs 2A, 13A), although) dorsaloutlnemmay,undulater(hics37/ a cere a eon ern orn nee 18 2 Carapace very strongly depressed dorsoventrally, lateral carina forming plate-like edge and produced anterolaterally to form a pair of rounded projections visible in dorsal view Eocuma winri (Fig. 4) — Carapace not strongly depressed dorsoventrally; carinae neither platelike nor forming 1110) db ee nr ee Rea mmr HM Cnn kG ak ois a'v'diso.000 000000 3 3 Carapace with one or more pairs of shallow longitudinal grooves or depressions but lacking latétal:-carinae. :4 rocco eee eee Oe cee Pas sO Sc 4 — Carapace with no more than one pair of longitudinal depressions; at least one pair of lateral carinae present (Pig. I2ZA)io0.0 02 a 4c os oe nace ee eee ae Oe Seer 5 4 All five pedigerous somites visible dorsally; a number of shallow longitudinal grooves Presentonicithensideolcarapacenaen ee eee eee Iphinoe crassipes (Figs 26-27) SOUTHERN AFRICAN CUMACEA: PART 2 167 — Only four pedigerous somites visible dorsally; a single shallow longitudinal groove present Onecithers(dexolucakapace saan re eae aes Mossambicuma elongatum (Figs 10-11) 5 Two pairs of lateral carinae on at least part of carapace (Fig. 43A, N) (lower one may be secondary, forming ventral edge of continuous midlateral depression (Fig. 43A) at least UENCC THO Lely) aesee cee or oh ok Ds poten Sut ap etee s S Une sep nia oltre Loy tauaee Dat taente asain 2 6 — A single pair of lateral carinae on carapace (Fig 45A, E); midlateral depression, if present, not bounded anteroventrally by continuous secondary carina (Fig. 46A,I).............. 9 6 Dorsal parts of free pedigerous somites laterally compressed, forming narrow plates Bodotria clara (Figs 34— = — Free pedigerous somites not compressed, not forming narrow plates dorsally........... (eCarapaceimorethanitwicerds long dsideepe. sme sean ee Bodotria tenuis (Fig. 2 SROakapaccuessaumanitwicerasn ONS asideCephavan aeons ici ie ce ise rie mie ecient = 8 Carapace of female wider than long, of male less than one and a quarter times as long as wide; carpus of pereiopod 1 less than three times as long as wide. . Bodotria falsinus (Fig. 43) — Carapace of both male and female at least one and a half times as long as wide; carpus of pereiopod 1 at least four times as long as wide......................-- Bodotria australis 9 Carapace about as wide as long and more than one and a half times as wide as deep, making animal conspicuously flat above; male with three pairs of pleopods........... Austrocuma platyceps (Fig. 12) — Carapace longer than wide and less than one and a half times as wide as deep; male with HIV SRDAINSHOLEO| CO POG Sen Mem iey ae Meet oh eo iam cue aoe ir an TAU A. eres cpheembee taer ere, 10 10 Second pedigerous somite carinate laterally................. Bodotria montagui (Fig. 41) = SECOMG! jOSGhiSroUS SOMA wVOKs CRnaTORNI JENIN. 56 050n050000000000000000000000008 11 11 Distal prolongation of basis of maxilliped 3 pointed, reaching well beyond insertion of carpus on merus; carapace distinctly more than twice as long as deep; lateral carina PLESCHMOMFANCeHOnRpant Of carapace omly.cews) she ce aoe oe Bodotria glabra — Distal prolongation of basis of maxilliped 3 rounded, hardly or not reaching insertion of carpus on merus (Fig. 39E); carapace no more than twice as long as deep; lateral carinae WRITS. oasigch whens ein Seles Cece RUS ad AER RROD NEG tema PON 7 efor eT en Re a 12 12 Pedigerous somites 4 and 5 elevated to a point dorsally (more distinct in female than BUY rea aa GAC AWARE) Ree ae re etek IN cena cane -F Dica cto CaO | SR Darl ARR Nl aN Os Vranas ya) oh 13 — Pedigerous somites 4 and 5 not elevated dorsally in either sex (Fig. 46A,I)............. 14 13 Lateral carina reaching posterior border of carapace with a longitudinal row of rounded WEPIESSIONSIDElOW. ns tie. Ck eee ee Bodotria vertebrata vertebrata (Fig. 44) — Lateral carina not reaching posterior border of carapace; no rounded depressions below Bodotria vertebrata semicarinata (Fig. 45) 14 Second pedigerous somite strongly elevated to a point dorsally in female and juvenile; lateral carina sinuous, most evident along midregion of carapace; integument often SHhOMelyaCal Cikle digesta As hone ie oh, Sen AD als eee Bodotria elevata (Figs 39-40) — Second pedigerous somite never elevated dorsally in either sex; lateral carina straight, evident along almost entire length of carapace; integument usually silky and never strongly Call Cili Colima ter I th ee ee ea ele Nt oe nite Teen eM TER Newsies Bodotria serica (Fig. 46) 15 Carapace with a pair of oblique ridges, depressions or slashes running from middorsal linestowanrdsaventrolateraledger(bigs) LOAN oD A) nasser eee aati nite eee cae 16 aa Cakapace withoutobliqueinmesulanities (RigOA) a ae oe caine aera cane 17 16 Caraplace slashed by transverse groove..............0..0eeeee Cyclaspis scissa (Fig. 32) — Carapace with raised transverse ridge, faint in adult male, very strong in females and YOU SENNal CSeen even ee wtel oon, Mie: ae Hehe N Sie Ok ei oe Rae ese Alticuma bellum: (Figs 15-16) 17 Single pair of acutely pointed lateral horns on carapace; integument tuberculate and hairy Eocuma aculeatum (Fig. 5) — Anterolateral edges of carapace produced to level of pseudorostrum to form blunted, horn-like projections; integument slightly wrinkled.................. Eocuma sp (Fig. 6) iS Puc SumMeEntstrongly:pilted Ok TUSOSeH ays asso soe ee eee eee eee ede eae: 19 — Integument smooth (faint pitting or reticulations may be visible at high magnifications). . 20 19 Integument, especially of carapace, very rugose; second pedigerous somite not narrower LY NTC NIK nearer yee ON Rated Pte aut at Soa. a einen plea nts es ee Bodotria magna (Fig. 36) — Integument strongly pitted; second pedigerous somite narrower than third.............. Eocuma foveolatum (Figs 2-3) 168 ANNALS OF THE SOUTH AFRICAN MUSEUM 20 Pseudorostral lobes not meeting in front of eyelobe (Fig. 31A).................00e8- 21 — Pseudorostral lobes meeting in front of eyelobe, even if only for a very short distance (Figs TB, 47@) 5.2. S eS es VO a ek oe ik ED RC eT on DD 21 Dorsal outline of carapace undulating; articulatory peg present between carapace and BOSE URS FONAESTOUIS SOMME. bono oaccaosdsceoscasdobdonsaue Bodotria nitida (Figs 37-38) — Dorsal outline of carapace smoothly arched; no articulatory peg between carapace and first ines PediSenous SoMmlen sean ter eer ee Cyclaspis australora (Figs 30-31) 22, €atapace at least two and a half timesivas longyas deep... 4.245405. eee ee 23 = @arapace less) than’ two anda half times as long as deeprn oe ene 10 eee 24 23 Carapace three times as long as deep, circular in cross-section, not serrate middorsally Iphinoe stebbingi (Figs 17-18) — Carapace nearer two and a half times as long as deep, elliptical in cross-section, serrate middorsallly.. 3. Ae Ge eee Oe ee ee ee ae Iphinoe producta (Fig. 21) 24 Pereiopod 2 less than three-quarters length of pereiopod 3... . Iphinoe africana (Figs 19-20) — Pereiopod 2 more than three-quarters length of pereiopod 3 deb le ed See 25 25 Middorsal carina evident; carapace elliptical in cross-section (Figs 9C, 13A, D)........ 26 — Middorsal carina defined poorly or not at all; carapace almost rounded in cross-section (Figs: TASB) sacs csi oa Be, He godess GOW eel wena aL Unig la Geet oe ON 28 DOESECOndepercioOpodm-=sccmentcd nr tn nearer Alticuma carinatum (Figs 13-14) = Second pereiopod G=seamented icc. 2 ok se ieee sae cles ee cn eee Pa] 27 Middorsal carina serrate; first pedigerous somite visible in both sexes..............---- Iphinoe dayi (Figs 22-23) - Middorsal carina not serrate; first pedigerous somite not visible in either sex........... Cyclaspoides pellucidus (Figs 8-9) 28 Carapace globose, vaulted dorsally, less than one and a half times as long as deep....... Cyclaspis spectabilis (Fig. 33) — Carapace not globose, not vaulted dorsally, more than one and a half times as long as deep (Figs TAs. 23A) esd jee sec otal lenestlee wae aaa any Wale eee 29 D9 AY CDSE RI fe sac cece os asec oes eee as ee ae Re ee nan ae Upselaspis caparti (Fig. 7) = LYE PIEESEME s 15 soc Siero oats 8 aicbinena eke Aula d Ree Gee enone: Caemcee, rah SERIALS eer 30 30 Prolongation of basis of maxilliped 3 reaching level of insertion of propodus on carpus 6 isp 12 Gl) ee ree ee ee ne A een ne eRe toto t.o.006 0905026 31 — Prolongation of basis of maxilliped 3 reaching merus (Fig. 28D)...............---++: 32 31 Prolongation of basis of maxilliped 3 a quarter its total length. Iphinoe fagei (Figs 24-25) — Prolongation of basis of maxilliped 3 a third its total length......... Iphinoe senegalensis 32 Merus and carpus of maxilliped 3 strongly flattened, carpus and propodus widely inserted Ono precedingscementS..4.54 ae ee en ee eee eee Iphinoe capensis (Fig. 29) — Merus and carpus of maxilliped 3 not strongly flattened, carpus and propodus inserted over little more than half width of preceding segments (Fig. 28D, L).............---++- 33 33 Basis of maxilliped 3 less than four times as long as wide, one and a half times length of LemMainineses ments tOcelhe heen ater Iphinoe truncata (Fig. 28) — Basis of maxilliped 3 six times as long as wide, twice length of remaining segments together Iphinoe ? zimmeri (Figs 47-48) Eocuma Marcussen, 1894 Generic diagnosis Carapace frequently with lateral horns and/or very distinct lateral carinae, almost always wider than deep. First pedigerous somite always invisible, second frequently fused with carapace. Distal prolongation of maxilliped 3 large and stout, merus frequently greatly expanded. Basis of pereiopod 1 distally pro- duced beyond insertion of ischium. Second pereiopod 6-segmented. Peduncle of uropod much shorter than telsonic somite or rami. Endopod of uropod 1-segmented. SOUTHERN AFRICAN CUMACEA: PART 2 169 Type species E. hilgendorfi Marcussen, 1894, from Japan. Remarks The genus, consisting of 22 species, is well known. In almost all cases its species can readily be distinguished by distinctive lateral horns and/or carinae. However, one of the new species, E. foveolatum, is in most respects quite clearly a member of the genus, yet has neither horns nor carinae in either sex, thus resembling the females of E. dimorphum Fage, 1928. Only a single specimen of Eocuma has previously been described from South Africa (Stebbing 1910). Distribution of Eocuma Members of the genus are confined to the warmer waters of the eastern hemisphere, being distributed mainly round the coast of Africa (12 species) and Indochina (9 species); 1 species occurs in Australian and 2 in Japanese waters; 3 of the African species are here described as new. Most species in the genus are found in shallow waters less than 50 m in depth, and a few are found as deep as about 100 m. The greatest recorded depth for the genus is vastly increased by the presence of EF. aculeatum sp. nov. from 550 m off Natal. KEY TO THE SPECIES OF EOCUMA 1 Carapace dorsoventrally flattened, entire lateral border carinate.................---- yD — Carapace rounded or dorsoventrally flattened, but carinate for less than half its length, OWE TOE: ENE EE eee scar gs ecu Me ABU a oA a la AT Poe Raitt 13 2 Carapace in dorsal view with at least one pair of distinct lateral horns or projections..... 3 — Carapace in dorsal view without distinct lateral horns, but edge may be incised or bear a [OG NUE OLE aT SL Ie se as a ce eam RR ce Ne eed Np ae Re Be caer 10 3 Second pedigerous somite fused with carapace, unsutured dorsally.................... dollfusi Calman, 19075— Mediterranean and Morocco to northern France —BSCCONGE DE CISeLOUS-SOMltenikee OL SULUTECIGOKSallymanh ane ee enone dees anos dee. 4 ABO AapacewAthoutapaired dorsalonidges sn ode ee ee ee ae see oes 5) — Carapace with paired dorsal ridges on posterior half at least..................--2-8- 8 5 Basis of pereiopod 1 almost as long as rest of limb. . productum Calman, 1907a—Indo-China —basisson perciopod 1 about two-thirds length of rest of limbyj..5.0--54.+624500 24> -- 6 6 In dorsal view pseudorostrum narrow, carapace tapering smoothly anteriorly from lateral horns in a straight line; horns laterally directed, tips forming widest part of carapace longicorne Calman, 1907a—Suez — In dorsal view pseudorostrum wide, carapace rounded anterior to lateral horns; horns anteriorly directed, tips slightly anterior to widest part of carapace.............-.++5- 7 7 Pereiopod 2 shorter than basis of pereiopod 3; second and third segments of antenna 1 Suibeciualleimmlemotines em en te ken te ial yan On 1 Jee ee neta een, att on. Weta k Winri Sp. NOV. — Pereiopod 2 longer than basis of pereiopod 3; second segment of antenna 1 about half lengthroteumindtscsiment er rmeneceiee ee oe coe taprobanicum Calman, 1904a—Ceylon Eye with at least three corneal lenses.............. hilgendorfi Marcussen, 1894—Japan Eye without lenses Carpusrolapenicopodmmmorechankewice lenethiot dactylnn 55 aan. see ee ae oe stelliferum Calman, 1907a—Indo-China Carpus of pereiopod 1 much less than twice length of dactyl...................--+-+: latum Calman, 1907a— Mediterranean, Indo-China, Japan oo \o 170 ANNALS OF THE SOUTH AFRICAN MUSEUM 10 Carapace smooth dorsally with no longitudinal ridges....... kempi Kurian, 1954—India = Carapace with at least one pair of longitudinalridges...-... 05. 0000 eo Cee eee il 11 Dactyl and propodus of pereiopod 1 of equal length. . . .cadenati Fage, 1928— West Africa — Dactyl of pereiopod 1 little more than half length of propodus..................... (V2 12 Lateral carina of carapace produced anteriorly to form two obtusely rounded lobes amakuense Gamo, 1967—Japan — Lateral carinae incised anteriorly forming a pair of slightly angular lobes.............. cochlear LeLoeuff & Intes, 1972— West Africa 13 Two pairs of horns forming pointed anterolateral projections..................-+--+-- calmani Fage, 1928—West Africa = One pair of horns laterally or none! .°.) 4c eenc tee ee en ee eee 14 14 Horns short, anteriorly directed, reaching anterior tip of pseudorostrum............. 15 — Horns laterally directed, not reaching level of pseudorostrum anteriorly, or absent...... 16 iS) Elonns welldevelopeds acutely; pointeds eee sarsi (Kossmann, 1880)—Red Sea Ons, poorivadeveloped avehyaSHOllR een ener Eocuma sp.—South Africa 16 Ischium of maxilliped 3 subequal in length to maximal length of merus, or longer..... 17 — Ischium of maxilliped 3 no more than half maximal length of merus................ 18 17 Dactyl of pereiopod 1 half length of propodus; peduncle of uropod nearly a third length OfsaMmite ates eee sr Sale Act atts, SUR Re rey me nS affine Calman, 1904a—India — Dactyl of pereiopod 1 longer than propodus; peduncle of uropod nearly quarter length of AMINE 5 3. de Banas repree sb, So eens Chacon og achs Re Renee mone neo agrion Zimmer, 1914— Australia 18 Basis of pereiopod 1 equal in length to next three segments together; horns present...... travancoricum Kurian, 1951—India — Basis of pereiopod 1 considerably longer than next three segments together; horns present OF ADSENES, 2 5:6. 5 a:e acegh Bais Seapos senescent Oe eR es OA 19 19 Carpus of pereiopod 1 longer than ischium and merus together; horns present........ 20 — Carpus of pereiopod 1 shorter than ischium and merus together; horns present or absent .21 20 Integument smooth; ischium of maxilliped 3 wider than long.................-----+-- lanatum LeLoeuff & Intes, 1972—West Africa — Integument tuberculate; ischium of maxilliped 3 longer than wide..... aculeatum sp. nov. 21 Maximal length of merus of maxilliped 3 little less than twice length of ischium; horns PRESEN se cee eae he se ee eee te eee ferox (Fischer, 1872)— Mediterranean — Maximal length of merus of maxilliped 3 nearly three times length of ischium; horns present. Or A@DSEnts sa2.. 6 se kd Ae ee ee ee Dap 22 Tip of basis of pereiopod 1 reaching end of ischium or just beyond; ¢ with horns, @ with- out: pereiopod 2 of 9 less than’ half leneth of perciopod 3h454-4- 0-4 o eo eee dimorphum Fage, 1928—West Africa — Tip of basis of pereiopod 1 reaching along a third length of merus; ¢ and 2 without horns; pereiopods,2 and) 3 role subequalsimylenethtes nee eee foveolatum sp. nov. Eocuma foveolatum sp. nov. Figs 2-3 Records sub. adult adult ovig. no. of 3 3 S$ ¢ @ juv. total records LB 33°S 18°E 5m 6 5 l “6 2a 24 ies FAL 34°S 18°E 15-60 m 5 5 Au 4) 2 3 21 20 SS 934282188 80 m 1 y 3 6 1 SCD 3458S 2358 -338)25-E 42-44 im 1 2 ML, 1 6 3 *Five samples collected by plankton net. SOUTHERN AFRICAN CUMACEA: PART 2 IFA Holotype Ovigerous female, in the South African Museum, SAM-A15492, collected during the UCT benthic survey, 5 December 1962. Type locality: 44 m, off East London (33°53’S 25°48’E). UCT station number SCD 378L. Description Ovigerous female, holotype, length 4,8 mm. Integument marked by fine reticulations interspersed with deep pits, particularly on sides of carapace, and fine hairs causing small particles of debris to stick to entire animal. Carapace (Fig. 2A) smoothly rounded, one and a half times as long as deep, with no trace of lateral horns. Antennal notch (Fig. 2B) small, anterolateral angle obtuse, defined by a small, sharp tooth. Carapace in dorsal view (Fig. 2C) almost oval, not much longer than wide, lacking middorsal carina; middorsal line marked by a shallow indentation. Pseudorostral lobes short, truncate anteriorly. Eyelobe eyeless, rounded, with a few scattered tubercles on surface. First pedigerous somite invisible. second well defined and not fused with carapace; third and fourth much less elevated dorsally than second, bearing rounded sideplates. Carapace twice length of free pedigerous somites. cephalo- thorax shorter than first five abdominal somites together. Abdomen very elongate, cylindrical. Antenna | (Fig. 2D) relatively short and stout. Flagellum (Fig. 2E) very short, l-segmented, with two aesthetascs; accessory flagellum minute, 1-segmented. Maxilliped 3 (Fig. 2F) stout, basis strongly flexed at mid-point, part distal to point of flexure subequal in length to rest of limb. Distal prolongation long and narrow, reaching articulation of carpus and merus. Merus three times length of ischium, expanded, distal prolongation reaching distal tip of carpus. Carpus, propodus and dactyl subequal in length, cylindrical. Pereiopod | (Fig. 2G) stout, basis slightly longer than remaining segments together, distal projection reaching beyond distal tip of ischium. Distal seg- ments fairly stout, all of similar lengths. Pereiopod 2 (Fig. 2H) long, 6-segmented; basis subequal in length to rest of limb. Merus and carpus subequal, propodus half length of dactyl. Pereiopods 3 (Fig. 21) to 5 similar, dactyl extremely small and unarmed. Telsonic somite two-thirds length of preceding one, not produced between uropods. Peduncle of uropod (Fig. 2J) very short, as wide as long, little more than half length of telsonic somite, unarmed. First segment of exopod expanded dorsally beyond insertion of second, less than half length of second, armed with a small spine on outer edge. Second segment armed with two long, fine spines on inner edge, seven short spines on outer edge and two complex hooked setae terminally. Endopod 1-segmented, bearing six fine plumose setae on inner edge and one very stout spine terminally. Adult male, paratype, length 6,7 mm. As female, except as follows: carapace (Fig. 3A) smoothly ovoid, nearly twice as long as deep, slightly flattened dorso- VD ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 2. Eocuma foveolatum sp. nov. Ovigerous female, holotype. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view of carapace. D. Antenna 1. E. Detail of distal tip of antenna 1. F. Maxilliped 3. G. Pereiopod 1. H. Pereiopod 2. I. Pereiopod 3. J. Uropod. Scale line = 1 mm for A, C; 0,5 mm for B, D, F—J; 0,25 mm for E. SOUTHERN AFRICAN CUMACEA: PART 2 173 wey, ar. « | Lif ¥ Yi YUy VY Fig. 3. Eocuma foveolatum sp. nov. Adult male, paratype. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view of carapace. D. Antenna 1. E. Detail of distal tip of antenna 1. F. Maxilliped 3. G. Pereiopod 1. H. Pereiopod 2. I. Uropod. Scale line = 2 mm for A; 1 mm for C, H; 0,5 mm for B, D, F-G, 1; 0,25 mm for E. 174 ANNALS OF THE SOUTH AFRICAN MUSEUM ventrally, integument with pits even more obvious than in the female. Antennal notch (Fig. 3B) very small, as is anterolateral angle. Carapace narrower in dorsal view (Fig. 3C), one and a half times as long as wide, marked middorsally by faint indentation. with a single pair of rounded depressions on either side about midway along length. Eyelobe narrower, bearing three large, clear lenses. Abdominal sideplates defined ventrally. Flagellum of antenna | (Fig. 3D-E) with two long and four short aesthetascs. Basis of maxilliped 3 (Fig. 3F) not flexed at midpoint, distal pro- longations of basis and merus broader, carpus and propodus also somewhat expanded. Basis of pereiopod | (Fig. 3G) with numerous sharp spines near midventral edge. Basis of pereiopod 2 (Fig. 3H) slightly shorter. Telsonic somite relatively smaller and narrower, uropods (Fig. 31) more slender and much more strongly armoured. Peduncle longer than wide, with three long plumose setae and numerous short, blunt spines on inner edge. Exopod with thirteen plumose setae on outer edge, six fine spines distally on inner edge and a single stout complex seta terminally. Endopod with thirteen long plumose setae and seventeen shorter blunter spines on inner edge, followed by six or seven scale-like serrations distally, and a strong complex spine terminally. Length Adult male 6,7-7,5 mm Ovigerous female 4,8—6,4 mm Remarks E. foveolatum is clearly distinguished from most other members of the genus by the complete absence of lateral horns or flattening of the carapace in both sexes. However, due to the structure of the third maxillipeds, the basis of pereiopod 1, and the uropods, it is readily accommodated in the genus. It is most closely allied to E. dimorphum Fage, 1928 from west Africa, in which the carapace of the female is smooth, although the male possesses a pair of lateral horns. Added to this, in both sexes of E. dimorphum the carapace is longer, the antennal notch poorly defined, the second pereiopod is much smaller than the third, and the merus of maxilliped 3 and the basis of pereiopod | are shorter. Adult females of E. dimorphum reach a size of 8 mm, whereas in E. foveolatum, the largest is 6,4 mm. The variable nature of the lateral horns in E. dimorphum links E. foveolatum with the more typical members of the genus. As a matter of interest, it was probably an individual of this species which Stebbing (1910) referred to as Cyclaspis sp, saying that it was similar in several respects to C. spectabilis, but hairy and ‘clogged with extraneous matter’. Distribution Apparently endemic to the warmer waters of the south and south-western coasts of South Africa, being found between 5 and 80 m from Langebaan Lagoon to East London. A fairly common species in shallow, sheltered waters SOUTHERN AFRICAN CUMACEA: PART 2 175 of the south-western Cape, rarer in other areas. Numerically it constitutes a little over 1 per cent of the individuals in the collection. Eocuma winri sp. nov. Fig. 4 Records NIWR_ 30°S 30°E-27°S 32°E 37-80 m_ 1 adult J, | ovig. 9, 3 29, 1 manca (5 records) Holotype Adult male, in the South African Museum, SAM-A15493, collected by the NIWR, 4 September 1975. Type locality: 50 m, off the Natal coast (27°33’S 32°41’E). NIWR station number MN 75/24 Gg. Description Adult male, holotype, \ength 11,6 mm. Integument white, shiny, faintly reticulate, brittle and opaque on carapace, less so on thorax and abdomen. Carapace (Fig. 4A) strongly depressed dorsoventrally, lateral edges marked by acute carina bearing single pair of forward-pointing blunt horns. No antennal notch or anterolateral angle. In dorsal view (Fig. 4B), lateral horns mark widest part of carapace. Pseudorostral lobes wide, meeting for some distance in front of bluntly rounded eyelobe. Eyelobe with five large clear lenses. Middorsal carina faint on carapace, wanting posteriorly. Second pedigerous somite narrow, immovably attached to carapace; third to fifth with sprays of plumose setae projecting dorsally in midline. Cara- pace slightly more than twice as long as deep, one and a half times length of free pedigerous somites. Cephalothorax slightly shorter than abdomen. Side- plates of abdominal somites strongly defined ventrally. Antenna | (Fig. 4C) of moderate length, basal segment wide and roundly geniculate. Next two segments subequal in length. Flagellum 1-segmented with two aesthetascs. Accessory flagellum small, 1-segmented. Maxilliped 3 (Fig. 4D) large and stout. Basis flattened, relatively short, almost twice length of rest of limb, serrated on inner edge. Distal prolongation strongly developed, reaching distal tip of carpus. Ischium large, longer than merus. Merus expanded, distally reaching articulation of carpus and propodus. Carpus slightly expanded towards midline. Propodus stout and dactyl slender. Pereiopod 1 (Fig. 4E) elongate, basis short, slightly longer than ischium, merus and carpus together, with blunt distal projection reaching beyond tip of ischium. Ischium short, half length of merus. Carpus, propodus and dacty]l very long, subequal. Pereiopod 2 (Fig. 4F) minute, 6-segmented. Basis only slightly longer than subequal merus and carpus together. Propodus and dactyl stout, subequal in length. 176 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 4. Eocuma winri sp. nov. Adult male, holotype, A. Lateral view. B. Dorsal view of carapace. C. Antenna 1. D. Maxilliped 3. E. Pereiopod 1. F. Pereiopod 2. G. Pereiopod 3. H. Uropod. Ovigerous female. I. Lateral view. J. Dorsal view of carapace. K. Maxilliped 3. L. Uropod. Scale line = 4 mm for A, B, I-J; 2 mm for E; 1 mm for D, G-H, K-L; 0,5 mm for C, F. Pereiopods 3 (Fig. 4G) to 5 similar, basis of pereiopod 3 longest and of pereiopod 5 shortest. Basis and carpus highly setose. Telsonic somite slightly shorter than preceding one, twice as long as wide, not produced between uropods. Peduncle of uropod (Fig. 4H) short and very stout, less than half length of telsonic somite with five plumose setae and numerous serrate spines in several rows on inner edge. Endopod about two SOUTHERN AFRICAN CUMACEA: PART 2 77 and a half times length of peduncle, 1-segmented, with eleven plumose setae and numerous serrate spines in several rows on inner edge and a single short, blunt spine terminally. First segment of exopod about a quarter length of second, unarmed; second armed with six plumose setae on inner edge, four on outer edge and two stout ones terminally. Ovigerous female, length 11,9 mm (slightly damaged), from 80 m off the Natal coast. As male, except as follows: carapace (Fig. 41) slightly more vaulted posteriorly and wider dorsally (Fig. 4J). Eyelobe smaller and bearing three lenses. Middorsal carina evident on carapace and all subsequent somites except the last. Second pedigerous somite wider, third very small, visible laterally as small flattened sideplate only. Abdominal somites much more slender, cylindrical. Flagellum of first antenna 2-segmented. Basis of maxilliped 3 (Fig. 4K) very stout, relatively shorter. Distal projection of pereiopod 1 shorter. Pereiopods 3 to 5 less setose. Peduncle of uropod (Fig. 4L) with seven plumose setae in one row on inner edge. Rami slightly longer, second segment of exopod unarmed except for two terminal spines. Endopod with ten plumose setae in one row on inner edge. Length Adult male 11,6 mm Ovigerous female 11,9 mm Remarks This species closely resembles E. taprobanicum Calman, 1904a from Ceylon, and the two may prove to be synonymous. EF. winri differs from Calman’s figures of E. taprobanicum in the narrower carapace of the female, the less well- developed lateral horns and the shorter basis of perieopod 1. In particular, pereiopod 2 is much smaller, the basis and merus together being shorter than the rest of the limb and the distal spine on the merus is shorter and weaker. The rami of the uropods are slightly shorter and the first segment of the endopod relatively larger. The differences are not considerable and these individuals may be representatives of a single species occurring from Natal to the tropical Indian Ocean and varying slightly from one end of the range to the other. Distribution At present known from the type locality and its vicinity, from 37 to 80 m off the Natal coast. With only six specimens known it is numerically insig- nificant in comparison with the total number of specimens in the collection, but is fairly common in the Natal material, representing almost 5 per cent of the individuals from this region. 178 ANNALS OF THE SOUTH AFRICAN MUSEUM Eocuma aculeatum sp. nov. Fig. 5 Records SAM 327-S 32-E. 550m Pimanea Holotype Manca, unique, in the South African Museum, SAM-A15491, collected by the South African Museum, 22 May 1974. Type locality: 550 m, in the southern Mozambique Channel (27°59’S 32°40’E). Meiring Naude station number SM 86. Description Manca, holotype, length 6,7 mm. Integument roughened by numerous small tubercles and hairs, especially on cephalothorax and first three abdominal somites. Carapace (Fig. 5A) nearly twice as long as deep, oval in lateral outline, with a single pair of well-developed lateral horns about a third of distance from anterior tip. Pseudorostral lobes (Fig. 5B) rounded, meeting for short distance in front of rounded, eyeless eyelobe. Middorsal line very slightly evident on carapace, not at all behind this. Fig. 5. Eocuma aculeatum sp. nov. Manca, holotype. A. Lateral view. B. Dorsal view of carapace. C. Antenna 1. D. Maxilliped 3. E. Pereiopod 1. F. Pereiopod 2. G. Telsonic somite and uropod. Scale line = 2 mm for A-B; 1 mm for D-E, G; 0,25 mm for C, F. SOUTHERN AFRICAN CUMACEA: PART 2 179 Free thoracic somites together two-thirds length of carapace; second pedigerous somite very narrow but not fused with carapace; third narrow, fourth and fifth with some tubercles, each longer than preceding one. Abdomen very long and slender, twice length of cephalothorax; first three somites tuber- culate, a row of tubercles running along the length down each side forming a ventrolateral ridge; last three somites unadorned, slender and translucent. Limbs all covered with very fine detritus. Antenna | (Fig. 5C) of moderate length, first segment stout, twice width of and subequal in length to next two segments. Flagellum l-segmented with two aesthetascs; accessory flagellum small, 1-segmented. Basis of maxilliped 3 (Fig. 5D) slender, distal prolongation long and narrow, reaching articulation of merus and carpus distally. Ischium small, merus very long and stout, distally produced to articulation of carpus and propodus. Carpus small, very slightly widened distally. Pereiopod | (Fig. 5E) of moderate length, basis fairly stout, subequal in length to rest of limb, distal point reaching beyond tip of ischium. Merus and propodus subequal in length, carpus longer. Pereiopod 2 (Fig. 5F) long and slender, 6-segmented. Merus and carpus subequal in length, twice as long as propodus. Dactyl fairly long, subequal in length to carpus and propodus together, armed with a few small spines distally. Telsonic somite (Fig. 5G) more than three times as long as wide, slightly shorter than uropods, very slightly produced between uropods and wider posteriorly. Peduncle of uropod about twice as long as wide, less than half length of rami, unarmed. First segment of exopod less than a third length of second, unarmed; second slender, serrations on inner margin interspersed with a few very fine setae and a single terminal spine. Endopod with three very fine setae on inner margin and one stout serrate seta terminally. Length Manca 6,7 mm Remarks The specimen is the only one known from southern Africa in which the carapace bears a pair of acutely pointed lateral horns and yet lacks lateral carinae. It is unique in the genus in the presence of long hairs and tubercles on the integument. Although it is not normally acceptable to describe a new species on the basis of a single immature specimen, there are several good reasons for doing so in this case. Firstly, very little more material is likely to become available from deep waters off the eastern seaboard. Secondly, the specimen is quite distinct from all other members of the genus in the adornment of the integument, is in a good state of preservation, and it should not prove difficult to match up adults of the same species at a later date. Thirdly, its presence adds considerably to the range of depths recorded for the genus. 180 ANNALS OF THE SOUTH AFRICAN MUSEUM Distribution At present known only from the type locality. Eocuma sp. Fig. 6 Eocuma sarsi: Stebbing, 1910: 414. Records SAM 32°S 28°E 56m 1 subadult male. Pieter Faure station number SAM-A590. Remarks Stebbing (1910) identified this specimen as E. sarsi (Kossmann, 1880), saying that ‘the proportions agree well with Kossmann’s figure’, but giving no figure or description himself. The author has re-examined the specimen, and perhaps due to the ravages of time, the external features no longer correspond well with either Stebbing’s (1913) or Calman’s (1904a) figures. In fact, it is dissimilar enough to suspect that it does not belong to E. sarsi. Since it is the only individual, and in a poor state of preservation, it would be unwise to refer it to a particular species. It is shown in Figure 6, and a very brief description is given below. The carapace (Figs. 6A, C) is extremely irregular, being considerably narrower dorsally and widening out quite abruptly ventrally. The lateral horns are ill-defined, being merely blunt protrusions reaching the level of the pseudo- rostrum on either side. The basis of maxilliped 3 (Fig. 6F) is very much produced distally, and the merus extremely large, being little less than half as long as the basis. The distal tip of the basis of pereiopod | (Fig. 6G) reaches about two- thirds along the length of the merus. The four distal segments of pereiopod 2 (Fig. 6H) are sub-equal in length; pereiopod 3 (Fig. 61) appears to be 6-segmented. The peduncle of the uropod (Fig. 6J) is not much shorter than the telsonic somite. Upselaspis Jones, 1955 Generic diagnosis First pedigerous somite not visible, second visible in females and some males. Second pereiopod 6-segmented. Endopod of uropod 2-segmented, peduncle subequal in length to rami. Type species Upselaspis caparti (Fage, 1951) (as Cyclaspoides caparti). The genus is monotypic. SOUTHERN AFRICAN CUMACEA: PART 2 181 Fig. 6. Eocuma sp. Subadult male. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view of carapace. D. Antenna 1. E. Detail of distal tip of antenna 1. F. Maxilliped 3. G. Pereiopod 1. H. Pereiopod 2. I. Pereiopod 3. J. Telsonic somite and uropod. Scale line = 2 mm for A; 1 mm for C, G-I; 0,5 mm for B, D, F, J; 0,25 mm for E. 182 ANNALS OF THE SOUTH AFRICAN MUSEUM Upselaspis caparti (Fage, 1951) Fig. 7 Cyclaspoides caparti Fage, 1951: 5-8, figs 2-4. Upselaspis caparti Jones, 1955: 284; 1956: 197. Records sub- adult adult ovig. no. of 3S S Q 9 juv. total records WCD 33°S17°E 78 m 1 1 1 (benthic) SAM 23°S 14°E 1m 4 ) 6 3 14 32 3 (planktonic) FISH 20°S12°E 95-O0m 1 1 1 (planktonic) Previous records Near Walvis Bay (22°S 14°E), plankton (Fage 1951; Jones 1955); Sierra Leone Estuary, 10-12 m (Jones 1956). Syntypes Males, females and juveniles, deposited by Fage (1951) in the Museé d’Histoire Naturelle, Paris. Type locality: surface plankton haul, near Walvis Bay, South West Africa (22°S 14°E). Description Ovigerous female, length 3,4 mm, from Sandwich Harbour (23°S 14°E). Integument slightly roughened, white, thin, with minute pits. Carapace (Fig. 7A) rounded, lacking carinae; deeper anteriorly than posteriorly, more than one and a half times as long as deep at deepest point. Antennal notch moderate, anterolateral angle rounded with very fine serrations below. Pseudorostral lobes (Fig. 7B) meeting for short distance in front of rounded, eyeless eyelobe. First pedigerous somite invisible, second half as wide as deep, third to fifth produced laterally to form distinct sideplates. Free pedigerous somites together about two-thirds length of carapace. Abdominal somites cylindrical; abdomen subequal in length to cephalothorax. Antenna | (Fig. 7C) elongate, reaching well beyond pseudorostrum. First and third segments subequal in length, second shorter. Flagellum 2-segmented with two aesthetascs. Accessory flagellum relatively long, 1-segmented. Basis of maxilliped 3 (Fig. 7D) twice length of remaining segments together, strongly angled; distal prolongation short, not reaching articulation of merus and carpus. Merus much expanded externally, distal tip reaching articulation of carpus and propodus. Basis of pereiopod 1 (Fig. 7E) one and a half times length of rest of limb, lower border with fine hairs, hooks and plumose setae. Last three segments subequal in length. Pereiopod 2 (Fig. 7F) elongate, almost as long as pereiopod 1, 6-segmented. SOUTHERN AFRICAN CUMACEA: PART 2 183 Basis subequal in length to remaining segments together. Pereiopods 3 (Fig. 7G) to 5 similar, each shorter than preceding limb. Ischium and merus short, subequal in length. Dactyl very small on pereiopod 3, apparently absent from pereiopod 5. Fig. 7. Upselaspis caparti Ovigerous female. A. Lateral view. B. Dorsal view of carapace. C. Antenna 1. D. Maxilli- ped 3. E. Pereiopod 1. F. Pereiopod 2. G. Pereiopod 3. H. Telsonic somite and uropod. Adult male. I. Lateral view. J. Detail of distal tip of antenna 1. K. Uropod. Scale line = 2 mm for I; 1 mm for A-B; 0,5 mm for C-H, K; 0,25 mm for J. 184 ANNALS OF THE SOUTH AFRICAN MUSEUM Telsonic somite (Fig. 7H) produced between uropods for a short distance, very slightly wider than long. Peduncle of uropod more than one and a half times as long as telsonic somite with four plumose setae and one spine on inner edge. Endopod slightly longer than exopod, first segment longer than second with two plumose setae and seven fine spines on inner edge; second unarmed except for one long stout spine terminally. First segment of exopod less than a quarter length of second with one plumose seta distally on inner edge; second with five plumose setae on inner edge and three slender spines terminally. Adult male, length 3,8 mm, from Sandwich Harbour. As female, except as follows: integument less translucent. Carapace rectangular in lateral outline (Fig. 71), a little less than twice as long as deep. Antennal notch and antero- lateral angle wanting. Second pedigerous somite not visible in some, usually very narrow, pointed dorsally and forming very small rounded sideplates laterally; covered anteriorly by posterior edge of carapace. Abdominal somites with sideplates defined ventrally. Abdomen slightly longer than in female. Flagellum of antenna | (Fig. 7J) stouter, bearing several fine setae as well as two aesthetascs. Basis of maxilliped 3 not angled, prolongation of basis narrower and more pointed distally, reaching articulation of merus and carpus. Basis of pereiopod 1 very stout proximally, bearing ten teeth distally on lower edge. Peduncle of uropod (Fig 7K) with five plumose and ten serrate setae. Exopod as in female. First segment of endopod with five plumose setae and one spine; second as long as first with two plumose setae on inner edge. Length Adult male 3,8-4,1 mm Ovigerous female 3,1-3,6 mm Remarks The specimens correspond in all details with those described by Fage (1951), except that the second pedigerous somite of the male is sometimes visible, and pereiopods 2 to 5 are relatively longer. Distribution Apparently endemic to the coast of west and south-west Africa, usually in estuaries at depths between | and 12 m, and usually found off the bottom, although a single benthic specimen is known from 78 m. The presence of this species in Sandwich Harbour constitutes the only records for any cumacean in estuarine conditions in South West Africa. Its distribution appears to be patchy but it may occur in quite large numbers in individual hauls. Cyclaspoides Bonnier, 1896 Generic diagnosis Carapace laterally compressed, pseudorostral lobes long. First two or three pedigerous somites incorporated in carapace at least dorsally. Second pereiopod SOUTHERN AFRICAN CUMACEA: PART 2 185 6-segmented. Telsonic somite long, peduncle of uropod short. Endopod of uropod I- or 2-segmented. Gut coiled. Type species Cyclaspoides sarsi Bonnier, 1896 Remarks Only one species of Cyclaspoides has previously been reported. The new species is clearly a member of the genus, but since the endopod of the uropod is 2-segmented, the generic diagnosis has been expanded accordingly. Distribution of Cyclaspoides C. sarsi is known from the Bay of Biscay (Bonnier 1896) in 950 m and off the coast of Ireland in 698 m (Calman 1905). It is also widely distributed in deep waters of the tropical and northern Atlantic down to 4934 m (Jones pers. comm.). Calman (1904a) tentatively assigned to this species a single specimen from the atrial cavity of an ascidian from Malaya. Until further material becomes available the identity of his specimen must remain in doubt. Since C. pellucidus is also a deep-water form, it appears that Cyclaspoides is essentially a deep-water genus. Cyclaspoides pellucidus sp. nov. Figs 8-9 Records SAM 34°S 17°E 400m _ 1 subadult 3, 2 ovig. 99 (1 record) SAM 27°S 32°E 820m_ 1 young 8 (1 record) Holotype Ovigerous female, in the South African Museum, SAM-A15490, collected by the S.S. Pieter Faure in about 1900. Type locality: approximately 400 m, off the Cape Peninsula (34°25’S 17°50’E). Pieter Faure station number SAM-— A10602 (PF 17440). Description Ovigerous female, holotype, length 5,2 mm. Integument thin, brittle, almost transparent, with fine reticulations. Coiled gut faintly visible. Exhalant siphon elongate, a third as long as carapace. Carapace (Fig. 8A) oval, pseudorostral lobes elongate; antennal notch (Fig. 8B) elliptical, anterolateral angle small, acute, serrated below for a short distance. Eyelobe (Fig. 8C) bluntly rounded, eyeless. Middorsal carina very slightly evident. First three pedigerous somites fused with carapace dorsally, sideplates of third visible laterally. Free region of thorax very short, about a quarter length 186 ANNALS OF THE SOUTH AFRICAN MUSEUM A Fig. 8. Cyclaspoides pellucidus sp. nov. Ovigerous female, holotype. A. Lateral view. B. Detail of anterior tip of carapace. C. Cara- pace in dorsal view. D. Antenna 1. E. Maxilliped 3. F. Pereiopod 1. G. Pereiopod 2. H. Pereiopod 3. I. Telsonic somite and uropod. Scale line = 2 mm for A; 1 mm for C; 0,5 mm for B, D-I. SOUTHERN AFRICAN CUMACEA: PART 2 187 Fig. 9. Cyclaspoides pellucidus sp. nov. Subadult male, paratype. A. Lateral view. B. Dorsal view. C. Pereiopod 2. Scale line = 1 mm for A-B; 0,5 mm for C. of carapace. Cephalothorax subequal in length to abdomen. Abdomen slender, cylindrical; first five somites with anterior articulatory peg. Antenna | (Fig. 8D) reaching tip of pseudorostrum, basal segment short, geniculate. Flagellum short, 1l-segmented, with two aesthetascs. Accessory flagellum minute, 1-segmented. Maxilliped 3 (Fig. 8E) stout, short, almost lacking setae. Basis nearly twice length of remaining segments together; distal prolongation short, reaching about half-way along merus. Merus somewhat expanded, almost reaching distal tip of carpus. Basis of pereiopod 1 (Fig. 8F) stout, longer than remaining segments together. Ischium and merus subequal in length, as are carpus and propodus. Pereiopod 2 (Fig. 8G) 6-segmented, shorter than pereiopod 3, unarmed except for a single terminal spine. Basis subequal in length to remaining seg- ments together; merus, carpus and propodus subequal in length, slightly shorter than dactyl. Pereiopods 3 (Fig. 8H) to 5 similar, third longest and fifth shortest, armed with a few spines. Telsonic somite (Fig. 81) longer than preceding somite, produced between uropods for about a fifth its length. Peduncle of uropod slightly more than half length of telsonic somite, moderately stout, unarmed. Rami subequal in length. First segment of exopod less than half length of second, unarmed; second armed with two spines on upper edge and two terminally. First segment of endopod three times length of second with six small spines on serrated 188 ANNALS OF THE SOUTH AFRICAN MUSEUM inner edge, and a stout one terminally. Subadult male, paratype, length of cephalothorax 2,7 mm. Last five abdomi- nal somites missing. As female, except as follows: carapace (Fig. 9A) slightly more rectangular, integument thinner, coiled gut clearly visible. Pseudorostral lobes (Fig. 9B) slightly longer. Pedigerous somites 2 and 3 fused with carapace, but suture lines distinct dorsally and laterally. Distal parts of antenna | missing, otherwise as in female. Maxilliped 3 more setose. Basis of pereiopod | straight, slightly broader. Pereiopod 2 (Fig. 9C) stouter, basis shorter, no longer than remaining segments together. Length Ovigerous female 5,2 mm. Remarks This species clearly belongs to Bonnier’s genus, being similar to C. sarsi in general appearance, as well as having only two free pedigerous somites and a coiled gut. It differs from C. sarsi in a number of respects, however: the second and third pedigerous somites are visible laterally in the female, and dorsally as well as laterally in the male, although in both cases they are firmly fused with the carapace (in C. sarsi they are indistinguishable or sometimes separated by a faint suture line). In C. pellucidus the endopod of the uropod is 2-segmented and the entire uropod slightly longer than the telsonic somite: in C. sarsi the endopod is l-segmented, and the uropod distinctly shorter than the telsonic somite. The varying number of segments in the endopod of the uropod is not unusual in genera of this subfamily, but it has necessitated expanding the diagnosis of the genus. Distribution Known only from two records, one at a depth of 400 m off the Cape of Good Hope, and one from 820 m in the southern Mozambique Channel. Mossambicuma gen. nov. Generic diagnosis First pedigerous somite not visible. Ischium of maxilliped 3 larger than merus. Basis of pereiopod 1 without distal projection. Second pereiopod 6-segmented. Telsonic somite shorter than fifth abdominal somite and longer than peduncle of uropod. Rami of uropod longer than peduncle, endopod 1-segmented. Type species M. elongatum sp. noy. (by monotypy). SOUTHERN AFRICAN CUMACEA: PART 2 189 Remarks It is with some hesitation that a new genus is erected for this species. In many ways it resembles some species of Eocoma, but the form of the carapace is quite different and the first pereiopod lacks the characteristic distal projection. Thus it seems better to erect a new genus which may later be submerged than to add species to Eocuma, which is at present well defined. Mossambicuma elongatum gen. et sp. nov. Figs 10-11 Records CON 23°S 32°E 1-2m_ 1 adult J, 1 ovig. 9, 1 young 9, 2 juvs (2 records) Holotype Ovigerous female, in the South African Museum, SAM—A15495, collected by A. C. Connell of the NIWR. Type locality: 1-2 m, near Mongue, Morrum- bene estuary, Mozambique (23°40’S 35°22’E). NIWR station number CON 3. Description Ovigerous female, holotype, length 2,2 mm. Integument uncalcified, flexible. Carapace (Fig. 10A) with scattered shallow pits, one and a half times as long as deep, slightly narrower dorsally due to a shallow depression running mid- laterally along carapace. A second smaller depression runs anteroventrally from posterior middorsal region for a short distance. Antennal notch shallow, anterolateral angle poorly defined, obtuse. Pseudorostral lobes produced beyond rounded, eyeless eyelobe for one-fifth total length of carapace (Fig. 10B). Second pedigerous somite about half as wide as deep, third to fifth about as deep as abdominal somites. Carapace about one and a half times length of free thoracic somites together, cephalothorax equal in length to first five abdominal somites. Abdominal somites cylindrical, elongate. Antenna 1 (Fig. 10C) elongate, protruding well beyond tip of pseudo- rostrum. First two segments subequal in length, together as long as third. Flagellum (Fig 10D) 2-segmented with one aesthetasc; accessory flagellum very small, 1-segmented. Maxilliped 3 (Fig. 10E) short and stout, basis broad and flattened; distal prolongation narrow, reaching beyond articulation of merus and carpus. Ischium as long as next three segments together. Merus wide, distal prolon- gation reaching articulation of carpus and propodus. Basis of pereiopod | (Fig. 10F) short, half length of remaining segments together, not produced distally. Propodus long, almost equal in length to merus and carpus together. Pereiopod 2 (Fig. 10G) 6-segmented, basis shorter than rest of limb. Dactyl twice length of propodus. 190 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 10. Mossambicuma elongatum gen. et sp. nov. Ovigerous female, holotype. A. Lateral view. B. Dorsal view of carapace. C. Antenna 1. D. Detail of distal tip of antenna 1. E. Maxilliped 3. F. Pereiopod 1. G. Pereiopod 2. H. Pereiopod 3. I. Telsonic somite and uropod. Scale line = 1 mm for A; 0,5 mm for B-C, F, I; 0,25 mm for D-E, G—H. Pereiopods 3 (Fig. 10H) to 5 similar, pereiopod 3 longest and 5 shortest. Merus and carpus stout, merus slightly the longer. Telsonic somite (Fig. 101) one and a half times as long as wide, slightly shorter than fifth abdominal somite, little produced between uropods. Peduncle of uropod less than two-thirds length of telsonic somite, less than half length of subequal rami, unarmed. First segment of exopod about a quarter as long as second; second armed with a single small spine dorsally near the tip, and two stouter ones terminally. Endopod 1-segmented, surface covered with small scales, with four very small spines on inner edge and a slightly longer one terminally. SOUTHERN AFRICAN CUMACEA: PART 2 191 Fig. 11. Mossambicuma elongatum gen. et sp. nov. Adult male, paratype. A. Lateral view. B. Detail of distal tip of antenna 1. C. Pereiopod 1. D. Telsonic somite and uropod. Scale line = 0,5 mm for A; 0,25 mm for C-D; 0,1 mm for B. Adult male, paratype, length 2,1 mm. As female, except as follows: cara- pace (Fig. 11A) slightly longer, pseudorostral lobes shorter. Fourth pedigerous somite produced dorsally to form a blunt point. Sideplates poorly defined ventrally. Antenna | (Fig. 11B) with two aesthetascs arising at base of flagellum and one distally. Basis of maxilliped 3 narrower proximally, ischium slightly shorter. Basis of pereiopod 1 (Fig. 11C) longer, merus narrower and propodus not as greatly elongated. Telsonic somite (Fig. 11D) not at all produced. Peduncle of uropod with five spines on inner edge, endopod with sixteen. Endopod and second segment of exopod scaly. Length Adult male 2,1 mm Ovigerous female 2,2 mm Remarks See remarks for genus Mossambicuma. Distribution So far only five specimens known, all from Morrumbene estuary, Mozam- bique, at depths from 1 to 2 m. 192 ANNALS OF THE SOUTH AFRICAN MUSEUM Austrocuma gen. nov. Generic diagnosis Dorsoventrally flattened Bodotriinae with four pedigerous somites visible in the male and five in the female. Eye present. Basis of maxilliped 3 slightly expanded distally. Second to fifth perelopods 6-segmented. Male with three pairs of pleopods, each with an outer process to the inner ramus. Endopod of uropod l-segmented. Type species Austrocuma platyceps sp. nov. (by monotypy) Remarks This genus is the only one in the subfamily in which the male has three pairs of pleopods. The flattened structure of the carapace and the nature of the third maxilliped and first pereiopod are also unusual in the Bodotriinae. Austrocuma platyceps gen. et sp. nov. igaei2 Records CP 34°S 18°E intertidal-2m 3 adult ¢g, 1 subadult J, 10 ovig. 99, 2 mancas (5 records) CPR 33°S 18°E intertidal 3 ovig. 92 (2 records) Holotype Ovigerous female, in the South African Museum, SAM-A15480, collected by UCT, 25 April 1956. Type locality: 1 m, Muizenberg Beach, Cape Peninsula (34°06’S 18°29’E). UCT station number CP 463B. Description Ovigerous female, holotype, length 1,8 mm. Whole animal dorsoventrally flattened (Fig. 12A). Integument smooth, carapace and thorax somewhat calcified, abdomen translucent. Carapace nearly twice as wide as deep with a single pair of lateral carinae, well-defined in the midregion, becoming rounded anteriorly and posteriorly. Anterolateral angle rounded, antennal notch small, triangular. Pseudorostral lobes rounded in lateral view, scalloped in dorsal view (Fig. 12B). Eyelobe wide, slightly pointed anteriorly, bearing reddish pigmented area with two pairs of large lenses on either side and one pair medially. Cephalothorax elliptical in dorsal view. Carapace subequal in length to free thoracic somites, of which first is narrow and second very wide. Cephalo- thorax nearly one and a half times length of narrow subcylindrical abdomen. Telsonic somite short, very slightly produced between uropods. Antenna | (Fig. 12C) stout, elongate. Basal segment subequal in length to next two segments together. Flagellum 2-segmented with one very short aesthetasc. Accessory flagellum minute, l1-segmented. SOUTHERN AFRICAN CUMACEA: PART 2 193 J E Gb \ \\ MN Fig. 12 Austrocuma platyceps gen. et sp. nov. Ovigerous female, holotype. A. Lateral view. B. Dorsal view. C. Antenna 1. D. Maxilliped 3. E. Pereiopod 1. F. Pereiopod 2. G. Pereiopod 3. H. Telsonic somite and uropod. Adult male. I. Lateral view. J. Dorsal view. K. Maxilliped 3. L. Pleopod 2. M. Telsonic somite and uropod. Scale line = 0,5 mm for A, I-J; 0,25 mm for B, D-G, K; 0,15 mm for C, H, L-M. 194 ANNALS OF THE SOUTH AFRICAN MUSEUM Maxilliped 3 (Fig. 12D) stout, basis curved through nearly 90°, produced distally beyond tip of ischium. Ischium, merus, carpus and propodus all flattened but not produced distally. Dactyl slender. Pereiopod 1 (Fig. 12E) very stout, meeting with its partner to form strong protection for the underlying mouthparts. Merus, carpus and propodus sub- equal in length, merus and carpus stout. Pereiopod 2 (Fig. 12F) 6-segmented, basis one and a half times length of rest of limb. Merus and carpus subequal in length, as are propodus and dactyl. Dactyl small, armed with a few small spines. Pereiopods 3 (Fig. 12G) to 5 similar, 6-segmented, shorter than pereiopod 2 and each shorter than preceding one. Ischium and merus subequal in length, as are carpus and propodus. Dactyl apparently wanting. Telsonic somite (Fig. 12H) short, wider than long; entire somite less than half length of peduncle of uropod. Peduncle unarmed, subequal in length to rami. First segment of exopod less than half length of second, unarmed; second serrated on inner edge with one long terminal spine. Endopod 1-segmented, serrated on inner edge, with one long fine spine and two shorter stouter spines terminally. Colour in life speckled above due to many dark brown and fewer greenish- white chromatophores; mainly greenish-white below, fading immediately on immersion in alcohol. Adult male, length 1,4 mm, from Kommetjie, Cape Peninsula. As female, except as follows: cephalothorax narrower, carapace longer than wide (Fig. 12J), nearly twice as long as deep (Fig. 121). Lateral carina confined to posterior half of carapace, which is excavated ventrally to accommodate the large exopods of maxilliped 3 and pereiopod 1. Second pedigerous somite much narrower. Carapace longer than free thoracic somites, cephalothorax longer than abdomen. Antenna | relatively stouter and slightly shorter. Basis of maxilliped 3 (Fig. 12K) less curved, distal segments stouter. Basis of pereiopod 1 straight, propodus and dactyl stouter, limb relatively longer, slightly more slender. Bases of pereiopods 2 to 5 shorter relative to rest of limb. Three pairs of pleopods present (Fig. 12L), rami bearing long plumose setae. Telsonic somite (Fig. 12M) as long as wide, rounded posteriorly. Peduncle slightly longer than rami. Endopod slightly shorter than exopod with four serrate spines on inner edge. Second segment of exopod not serrate. Length Adult male 1,4-1,5 mm Ovigerous female 1,6-1,8 mm Remarks This species is quite distinct from all other members of the subfamily, partly because of its very flattened appearance in both sexes, and more particu- larly because of the presence of only three pairs of pleopods in the male. The SOUTHERN AFRICAN CUMACEA: PART 2 195 reduction in number of pleopods is perhaps less strange than may appear at first sight, since this tendency is found in some of the Vaunthompsoniinae as well as routinely in some of the other families. In these forms the thoracic exopods are usually highly developed, as is the case in the males of Austrocuma platyceps. Distribution At present only a few specimens are known from the shores of the Cape Peninsula, from Muizenberg to Hout Bay, at depths from 0 to | m. Note: Just prior to going to press several more specimens of this species were found intertidally between Ysterfontein and Melkbosch on the south- western Cape coast. Alticuma gen. nov. Generic diagnosis First pedigerous somite visible in both sexes. Second pereiopod 7-segmented. Endopod of uropod 2-segmented. Type species Alticuma carinata (Zimmer, 1921) (as Cyclaspis carinata). Remarks The above combination of characters exhibited by the two species in the collection excludes them from Jphinoe and Cyclaspis, the genera closest to them, since placing them in either would necessitate the expansion of an existing definition. The limits of the genera of the Bodotriinae are discussed on page 163. The differences between the two species included in the genus are mainly in the superficial appearance of the carapace, the body and appendages being rather similar in structure. Thus the genus may in fact prove to be a realistic assemblage of species rather than a merely convenient grouping. Distribution of Alticuma Thus far limited to waters deeper than 183 m off the coast of southern and eastern Africa. Alticuma carinatum (Zimmer, 1921) n. comb. Figs 13-14 Cyclaspis carinata Zimmer, 1921: 126-127, figs 19-21. Records sub- adult adult ovig. no. of 3S ae or Q juv. total records WE Mol eSe li E 208-500 m 5 Wg 2 8 5 WCD 34°S 17°E 320 m 1 1 1 SCD 34°S 20°E-34°S 23°E =183-200 m 2} 3 1 6 3 SST 35°S 22°E 200 m D, 8 G6 5 S) AL By 4 SAM _27°S 32°E-30°S 30°E 550-850 m 1 2 A eS lS 5 196 ANNALS OF THE SOUTH AFRICAN MUSEUM Previous records Type locality, holotype specimen only. Holotype Female, deposited by Zimmer (1921) in the Berlin Zoologisches Museum. Type locality: 693 m, off the east African coast (1°S 41°E). Description Ovigerous female, length 6,7 mm, from 200 m on the Still Bay transect. Integument translucent with fine reticulations and pits visible at high mag- Fig. 13. Alticuma carinatum Ovigerous female (SST). A. Lateral view. D. Dorsal view of carapace. E. Detail of pedigerous somites from the side. F. Antenna 1. G. Maxilliped 3. H. Pereiopod 1. I. Pereiopod 2. J. Pereiopod 3. K. Telsonic somite and uropod. Females. B. Lateral view of SAM specimen. C. Lateral view of LBT specimen. Scale line = 2 mm for A-D; 1 mm for E, H, K; 0,5 mm for F-G, I-J. SOUTHERN AFRICAN CUMACEA: PART 2 197 nifications. Dorsal half of carapace strongly compressed laterally forming a narrow, pointed median carina (Fig. 13A). Pseudorostral lobes elongate, meeting in front of eyelobe for one-sixth total length of carapace. Eyelobe distinct, elliptical, eyeless (Fig. 13D); in lateral view visible above level of pseudorostral lobes and slanting steeply to the elevated posterior region of the carapace. One tooth present midway along middorsal carina. Carapace twice as long as deep. Antennal notch deeply excavate between pseudorostral lobes, antero- lateral angle acutely pointed. (Note: the outline of the carapace in lateral view is very variable. The pseudorostral lobes may be as much as a quarter the total length of the carapace (some SAM specimens, Fig. 13B); the eyelobe may not be elevated above the pseudorostral lobes (Some LBT and SAM specimens, Figs 13B—C); the anterior part of the middorsal carina may bear 0-2 large teeth (most of the SST speci- mens, Fig. 13A), or a long row of many minute ones (SAM and some LBT specimens, Fig. 13B), or ones intermediate in size and number (some LBT specimens, Fig. 13C). Only the carapace is variable, the limbs being similar in all specimens. Too few males are available to determine the degree of variability in them, or the likelihood of there being more than one species represented.) All five pedigerous somites visible laterally, second and third apparently partly fused dorsolaterally (Fig. 13E). Free thoracic somites slightly flanged laterally with a faint middorsal carina. Carapace nearly half as long again as free thoracic somites. Cephalothorax subequal in length to abdomen. Arti- culatory notches present laterally on all abdominal somites except the last. Basal segment of antenna | (Fig. 13F) geniculate, equal in length to next two subequal segments together. Accessory flagellum short, 1-segmented. Flagellum 1-segmented with one aesthetasc longer than the antenna. Maxilliped 3 (Fig. 13G) stout, basis angled, distal prolongation reaching articulation between merus and carpus. Ischium longer than wide; merus sharply expanded externally, prolongation reaching distal tip of carpus. Carpus expanded distally, equal in length to subequal propodus and dactyl. Pereiopod 1 (Fig. 13H) not elongate. Basis equal in length to rest of limb. Carpus and propodus subequal in length, longer than dactyl. Pereiopod 2 (Fig. 131) stout, 7-segmented. Basis longer than remaining segments together. Ischium short but distinct. Pereiopods 3 (Fig. 13J) to 5 similar, less stout than pereiopod 2. Telsonic somite (Fig. 13K) produced between uropods for nearly a third its length, slightly shorter than peduncle of uropod. Peduncle unarmed but serrate on inner margin, two and a half times length of rami. Exopod slightly longer than endopod, first segment unarmed, half length of second; second only with two terminal spines. First segment of endopod twice length of second with one spine distally on inner edge, second with two terminal spines only. First segment of exopod and both of endopod serrated on inner margin. Adult male, \ength 6,7 mm, from 200 m on the Still Bay transect. Differs 198 ANNALS OF THE SOUTH AFRICAN MUSEUM from the female as follows: integument less calcified. Carapace less sharply carinate, less sloping behind eyelobe (Fig. 14A). Pseudorostral lobes shorter (Fig. 14B), anterolateral angle obtuse, blunt. First pedigerous somite obscured laterally by anterior projection of second; second narrow, not fused with third. Ventral sideplates well marked on abdomen. Accessory flagellum of antenna 1 (Fig. 14C) with numerous short aesthetascs. Basis of maxilliped 3 not angled, ischium slightly longer and merus shorter. Basis of pereiopod 1 with eight sharp spines in mid-region. Peduncle of uropod (Fig. 14D) with six fine spines on inner edge, followed by fourteen serrate setae in pairs, with two spines distally. Second segment of exopod with four plumose setae on inner edge; first of endopod with five fine spines, second with three and a single end-spine. Length Adult male 6,7 mm Ovigerous female 5,8-7,4 mm Remarks This species appears to be the same as Zimmer’s Cyclaspis carinata. However, he describes and figures only the whole animal and the telsonic somite and uropod, and the author has not been able to obtain the type for comparison. Zimmer’s figures correspond with the SAM material, except that in the latter the ovigerous female is broader across the posterior part of the cara- pace, the telsonic somite does not appear to be emarginate posteriorly, and there are no setae on the inner border of the peduncle in the females. The most significant difference is that the inner ramus of the uropod is 2-segmented in all specimens examined, whereas Zimmer’s appears to be l-segmented. How- Fig. 14. Alticuma carinatum Adult male. A. Lateral view. B. Dorsal view of carapace. C. Distal tip of antenna 1. D. Uropod. Scale line = 2 mm for A-B; 0,5 mm for D; 0,25 mm for C. SOUTHERN AFRICAN CUMACEA: PART 2 199 ever, the diagram is poor, and the author is disinclined to place too much emphasis on this character. It is not possible to state with certainty that all Specimens under discussion are conspecific, but from the characteristic general appearance of the animals, it is suggested that they probably are. The variability of the available specimens is confined to characters of the carapace, and so at this stage it is assumed that the various morphological variations are intra- specific, rather than that there are as many as four distinct species, all very closely related, and all represented by very few specimens. The situation may well have to be reviewed when further material is available. The fusion of the second and third pedigerous somites is extremely unusual in this family, but once again it is a variable character, not present even in all the ovigerous females. Although it may be of considerable functional significance to the living animal, its sporadic occurrence means that it is of no immediate taxonomic value. Distribution The most common species in deep waters off South and east Africa, from Lambert’s Bay to Kenya, at depths from 183 to 810 m. Alticuma bellum gen. et sp. nov. Figs 15-16 Records SAM 30°S 30°E-26°S 33°E 550-1 300m _ 1 adult g, 47 subadult gd, 9 ovig. 29, 69 gd and 99, 13 juvs, 32 mancas (10 records) Holotype Ovigerous female, in the South African Museum, SAM—A15479, collected by the South African Museum, 22 May 1976. Type locality: 550 m, southern Mozambique Channel (27°59’S 32°40’E). Meiring Naude station number SM 86. Description Ovigerous female, holotype, length 10,0 mm. Integument white, slightly crystalline. Carapace (Fig. 15A) divided transversely by strong ridge running across middorsal region almost to ventrolateral edge; smooth posterior to this, lower and with two slight protuberances on either side anterior to it. Pseudorostral lobes short, not meeting in front of eyelobe. Anterolateral angle acute, antennal notch angular (Fig. 15B). Carapace in dorsal view (Fig. 15C) abruptly wider across posterior part due to transverse ridge. Eyelobe eyeless. Carapace twice as long as deep, one and a half times length of free pedigerous somites together. Cephalothorax equal in length to first five abdominal somites together. Abdominal somites cylindrical. 200 ANNALS OF THE SOUTH AFRICAN MUSEUM Antenna | (Fig. 15D) small, first segment subequal in length to next two. Flagellum l-segmented with two aesthetascs; accessory flagellum minute, 1-segmented. Maxilliped 3 (Fig. ISE) stout, basis strongly angled, about two and a half times length of remaining segments together. Distal prolongation reaching Fig. 15. Alticuma bellum gen. et sp. nov. Ovigerous female, holotype. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view. D. Antenna 1. E. Maxilliped 3. F. Pereiopod 1. G. Pereiopod 2. H. Pereiopod 4. I. Telsonic somite and uropod. Scale line = 4 mm for A; 2 mm for B—-C, F; 1 mm for D, E, G-I. SOUTHERN AFRICAN CUMACEA: PART 2 201 beyond articulation of merus and carpus. Ischium square in outline, merus expanded distally, reaching articulation of carpus and propodus; carpus slightly expanded, propodus and dactyl subequal in length, cylindrical. Pereiopod 1 (Fig. 15F) elongate, basis less than two-thirds length of rest of limb. Ischium small, half length of merus. Carpus subequal in length to merus and ischium together; propodus and dactyl very long: propodus longer than ischium, merus and carpus together, dactyl slightly shorter. Pereiopod 2 (Fig. 15G) 7-segmented. Basis subequal in length to rest of limb, ischium small, merus and carpus subequal, merus stouter. Dactyl twice length of propodus. Pereiopods 3, 4 (Fig. 15H) and 5 similar, basis of 3 longest. Carpus relatively very long with four hooked setae. Propodus and dactyl subequal. Telsonic somite (Fig. 151) two-thirds lengths of peduncle of uropod, produced between uropods for about one-third its length. Peduncle of uropod less than twice length of exopod, slender, with eight small spines on inner edge. Exopod slightly longer than endopod, first segment less than one-third length of second, unarmed; second armed with two small spines on inner edge and four terminally. First segment of endopod one and a half times length of second, with six spines on inner edge, interspersed with scale-like serrations; outer edge serrated; second segment with one fine spine on inner edge and a short one terminally (probably broken). Adult male, \ength 10,3 mm, from near the type locality. As female, except as follows: carapace (Fig. 16A) less than twice as long as deep, transverse ridge very much fainter, anterolateral angle (Fig. 16B) smaller and less acute; anterior and ventral edges of antennal notch serrated. First pedigerous somite visible dorsally only. Abdominal sideplates distinctly defined ventrally. Anterior end of single specimen damaged. Free pedigerous somites flanged laterally (Fig. 16C). Basal segment of antenna | (Fig. 16D) larger, setose; numerous aesthetascs surrounding flagellum. Basis of maxilliped 3 (Fig. 16E) less angled, merus narrowly expanded. Propodus and dactyl of pereiopod | relatively shorter. Merus, carpus and dactyl of pereiopod 2 slightly longer and stouter. Bases of pereiopods 3 and 4 (Fig. 16F) shorter, carpus relatively large. Both uropods damaged. Subadult male, paratype, length 10,5 mm. Carapace (Fig. 16G) strongly sculptured, transverse ridge very pronounced, posterior and ventral regions with scattered rounded projections. Distinct ridges running posteriorly from antennal notch and posteroventrally from eyelobe suture nearly to ventral edge. Peduncle of Uropod (Fig. 16H) with twelve spines on inner edge. Second segment of exopod serrated distally on inner edge. First segment of endopod serrated on both edges and second on inner edge. Manca, paratype, length 4,6 mm (Fig. 161). Carapace smoother than in adults, transverse ridge strongly pronounced, edge defined by a row of small rounded tubercles. 202 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 16. Alticuma bellum gen. et sp. nov. Adult male. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view. D. Antenna |. E. Maxilliped 3. F. Pereiopod 4. Subadult male, paratype. G. Lateral view of carapace. H. Uropod. Manca, paratype. I. Lateral view of carapace. Scale line = 4mm for A; 2 mm for B-C, G; 1 mm for D-F, H-I. Length Adult male 10,3 mm Ovigerous female 10,0 mm Remarks Although in outward appearance this species is very different from A. carinatum, the presence of a free first pedigerous somite, 7-segmented second pereiopod and 2-segmented endopod of the uropod would place them in the same genus. Certainly the appendages are very similar, but the lengths of the pseudorostral lobes are very different and the sculpturing of the carapace is quite dissimilar. The species may easily be distinguished from all other South African members of the subfamily by the transverse ridge on the carapace. It is not very different in external appearance from some of the members of Hale’s ‘exculpta’ group of Cyclaspis (Hale 1944a), but is clearly separated from all of them by the nature of pereiopod 2 and the endopod of the uropod. SOUTHERN AFRICAN CUMACEA: PART 2 203 The uropods of a number of specimens are rather different from those described above. The peduncle is shorter by about a third than the telsonic somite and the rami are subequal in length to the peduncle. The proportions of the rami are the same as those described above. It would appear that all of these specimens are immature, and that this is merely a juvenile variation, as others from the same sample have uropods almost identical with those of adults. The sculpturing of the carapace is also less pronounced in these speci- mens. Otherwise they seem to be identical to those described above. Distribution One of the more common deep-water species from Natal to the southern Mozambique Channel at depths from 550 to 1 300 m. Iphinoe Bate, 1856 Generic diagnosis Five pedigerous somites visible, the first short. Second pereiopod 6-segmented. Endopod of uropod 2-segmented, rami no longer than peduncle. Type species I. trispinosa Bate, 1856—Europe, to 150 m. Distribution of Iphinoe While the greatest depth recorded for I. producta is 280 m, for I. serrata 1175 m (Jones, pers. comm.) and for J. trispinosa 150 m, the others appear to be confined to depths less than 100 m. Of the 33 species so far described, only 2 are found exclusively north of 40°N, the ranges of 3 extend north of 40°N and south of 40°S and the other 28 are found between these latitudes. Thus Iphinoe is essentially a genus of warm, shallow waters. 24 species are found in European and African waters; 3 of these also extend to India, where there are also 2 endemic species; 6 species are found in other regions (Indochina and Australia). Although the known ranges of many species will undoubtedly be increased by further collecting, the rate of endemism appears to be high. 25 species are known only from rather restricted areas, while 3 occur in two or more oceans. 7 of the 9 South African species are endemic, | also occurs in west Africa and | in both west Africa and India. Since the South African species of Iphinoe appear to be linked only to those from west Africa and India, the following key applies to species occurring in these areas only. KEY TO THE AFRICAN AND INDIAN SPECIES OF JPHINOE il CAarApace at least inv© ainal 2 Gerace wimnes AS llomie AS CEE. oo 565 bborcsonbonsuoeogous D; = (CAAOAGEe JESS WAIN TWO amcl A CWAIKISE tines AS OME AS GIES. 55 56500005000040500000008 B) 2, PEO DOGS ZF ginal 3 Sulosciwell im WEAN. ssoncccsccccbonconcceoeonencr producta sp. nov. REKclopodesaboumhialiglenctimolmperclOpOdesey anita. Ge okie caite acre erie ede 3 204 ANNALS OF THE SOUTH AFRICAN MUSEUM 3 Nn | O Carapace about two and a quarter times as long as deep; basis of pereiopod 2 longer than rest: OF lily sel aake oun ces nee Eee Ree africana Zimmer, 1908—South Africa Carapace about three times as long as deep; basis of pereiopod 2 shorter than rest of limb.4 Adults less than 10 mm in length; basis of pereiopod 1 about one and a half times length of rest of limb; basis of pereiopod 2 hardly longer than wide......................... brevipes Hansen, 1895—west Africa Adults more than 12 mm in length; basis of pereiopod 1 about one and three-quarters length of rest of limb; basis of pereiopod 2 about twice as long as wide................ stebbingi Jones, 1956—South Africa Carapace no more than one and two-thirds times as long as deep, or if nearly twice as long as deep in male then basis of pereiopod | no more than three times as long as wide. . .6 Carapace twice as long as deep; basis of pereiopod 1 about six times as long as wide... .10 Pseudorostrum upturned, lower edge curled inwards, truncate anteriorly.............. 7 Pseudorostrum straight, not curled inwards, not obviously truncate.................. 8 Antennal notch very deeply excavated, anterolateral angle strongly produced and serrate in both sexes; a pair of dorsal ridges running back from eyelobe almost to posterior edge OlMCARAPACE Sek 28. oe erent eee ene cae pokoui LeLoeuff & Intes, 1972—west Africa Antennal notch absent in male, moderate in female; anterolateral angle normal, slightly serrate ventrally; faint dorsal ridges on front half of carapace only................... crassipes Hansen, 1895—South African and Indian form Middorsal line of carapace serrate. . tenella Sars, 1878 —India, west Africa, Mediterranean = Miuddorsal line of carapace not serrate... 95.6056 on lee ee nee cn eee 9 10 14 15 Prolongation of basis of maxilliped 3 not reaching articulation of merus and carpus; OMSOIS TOE QOMINGEE 4 oo dob ondsovmroboodooboo‘ES capensis (Zimmer, 1921)—South Africa Prolongation of basis of maxilliped 3 reaching beyond articulation of merus and carpus; menus very slightly expandedme nse ase ena truncata Hale, 1953—South Africa (estuarine) Prolongation of basis of maxilliped 3 reaching articulation of carpus and propodus; merus strongly expanded....... crassipes Hansen, 1895—west African and Mediterranean form Carapace witha pain of dorsolateralicaninaesae saeco leer eee 11 Carapace lacking dorsolateral carimae.. 54 2295.2 cos oe ee to Lon eee iy Carinae confined to dorsal half of carapace. . .plicata LeLoeuff & Intes, 1972—west Africa Carinae running diagonally from anterolateral corner almost to mid-dorsal line......... robusta Hansen, 1895—west Africa 0=3 minute serrations middorsallly 3.75 0. Seu oo oe eee eee eee 13 Numerous! serrations ontat least halivon middorsalicaninds inn eens 14 Prolongation of basis of maxilliped 3 comprising one-third its total length; carapace slightly less than twice as long as deep. . . senegalensis Jones, 1956—South and west Africa Prolongation of basis of maxilliped 3 comprising one-quarter its total length; carapace slightly more than twice as long as deep............... fagei Jones, 1955—South Africa Basis of pereiopod 1 subequal in length to rest of limb. ..dayi Jones, 1960—South Africa Basis of pereiopod 1 no longer than next four segments together.................... 15 Merus of maxilliped 3 expanded, basis little longer than remaining segments together..... sanguinea Kemp, 1916—India (lentic) Merus of maxilliped 3 not expanded, basis one and a half times length of remaining seg- ments togethers ase eee ee pigmenta Kurian, 1961 —India (lentic) Iphinoe stebbingi Jones, 1956 Figs 17-18 Iphinoe brevipes (non Hansen, 1895): Stebbing, 1910: 410. Jones, 1955: 288. Iphinoe stebbingi Jones, 1956: 203-205, figs 10-12; 1960: 175. SOUTHERN AFRICAN CUMACEA: PART 2 205 Records sub- adult adult Ovig. no. of 3S € € & Q jw tora wecoras WCD 33-34°S 18°E 65-84 m 2 Di 6 1 2 FAL & FBY 34°S 18°E 7m 22 143) ily SS Zils Wy Bye 7 SST BAS WP 18 50-80 m 8 8 Ta NG Bil 74 8 SCD 34°S 21°E-33°S 27°E 36-100 m 7 1S BS I) 30, BO TN a NIWR 30°S 30°E-29°S 31°E 30-62 m 1 i 2 4 4 SAM 34°S 18°E-34°S 22°E 55-87 m 10 ihe TE 9 30 8 Previous records Cape Point to St Francis Bay, 44-62 m (34°S 18°E-33°S 25°E) (Stebbing 1910 (= Jones 1956)); False Bay to Cape Agulhas (34°S 18°E-34°S 19°E), 20-82 m (Jones 1960). Syntypes Adults of both sexes deposited by Jones in the British Museum (Natural History): specimens previously identified by Stebbing as J. brevipes. Type locality: not specified; material from St Francis Bay (33°S 25°E), off Cape Point Lighthouse (34°S 18°E) and off Sebastian Bluff (34°S 22°E). Description Ovigerous female, length 15,9 mm, from False Bay. Slender, elongate. Body cylindrical, carapace slightly compressed laterally. Integument shiny with faint reticulations at high magnifications. Carapace (Fig. 17A) slightly less than three times as long as deep (slightly more in non-ovigerous females), with a faint middorsal carina, especially on posterior half. Antennal notch excavate, anterolateral angle acute, tooth present (Fig. 17B). Pseudorostral lobes meeting for a short distance in front of elongate, eyeless eyelobe (Fig. 17C). First pedigerous somite visible, about a third as long as second; second longer than third. Thorax slightly longer than carapace, cephalothorax longer by two somites than abdomen. Antenna | (Fig. 17D) of moderate length, first and third segments subequal in length, second slightly shorter. Flagellum 1-segmented with two aesthetascs. Accessory flagellum minute, 1-segmented. Basis of maxilliped 3 (Fig. 17E) three times length of remaining segments together; distal prolongation reaching half way along merus (Fig. 17F), merus slightly expanded. Basis of pereiopod 1 (Fig. 17G) one and three-quarters times length of rest of limb, slender, with several spines on outer distal edge. Ischium and merus subequal in length, as are next three segments. Pereiopod 2 (Fig. 17H) 6-segmented, short, stout, equal in length to basis of pereiopod 3. Basis twice as long as broad, merus stout. Pereiopods 3 (Fig. 171) and 4 stout, 7-segmented. Pereiopod 5 (Fig. 17J) with ischium, merus and carpus very much enlarged, ANNALS OF THE SOUTH AFRICAN MUSEUM Iphinoe stebbingi |Eue INI Ovigerous female. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view of carapace. D. Antenna 1. E. Maxilliped 3. F. Detail of distal tip of maxilliped 3. G. Pereiopod 1. H. Pereiopod 2. I. Pereiopod 3. J. Pereiopod 5. K. Telsonic somite and uropod. 2mm for E,G; 1 mm for B, D, I, K; 0,5 mm for F, H, J. 9 Scale line = 4 mm for A, C; SOUTHERN AFRICAN CUMACEA: PART 2 207 Fig. 18. Iphinoe stebbingi Adult male. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view of carapace. D. Pereiopod 2. E. Uropod. Scale line = 3 mm for A, C; 1 mm for B; 0,5 mm for D-E. merus and carpus subequal in length and width to basis, each of the first four segments furnished with fans of stout setae. Propodus and dactyl small, cylindrical. Telsonic somite (Fig. 17K) one and a half times as long as broad, equal in length to peduncle of uropod. Peduncle with two spines proximally followed by thirteen plumose setae in one row on inner edge. Rami subequal in length. First segment of exopod shorter than second, with two plumose setae on inner edge; second with six plumose setae dorsally and sixteen on inner edge and four short terminal spines. First segment of endopod half length of second with six serrate setae on inner edge, second with fourteen on inner edge and nine plumose 208 ANNALS OF THE SOUTH AFRICAN MUSEUM setae on tip and inner edge distally. Adult male, length 14,9 mm, from False Bay. As female, except as follows: integument thinner and less polished in appearance. Carapace almost exactly three times as long as deep (Fig. 18A). Anterolateral angle wanting, antennal notch only slightly excavated (Fig. 18B). Posteroventral corners of carapace produced backwards to cover part of first pedigerous somite, which is narrowly visible. Eye consisting of five lenses (Fig. 18C), no pigment visible in preserved specimens at least. Sideplates of pereion produced backwards, of fourth also produced forwards to overlap third; abdominal sideplates defined ventrally. Ventral sternites poorly armed, first with two spines produced anteriorly between first perelopods; second with raised protuberance bearing anteriorly- directed spine; third unarmed; fourth as second; fifth with slight, rounded protuberance. Carpus of maxilliped 3 slightly less expanded. Pereiopod 2 (Fig. 18D) less stout, relatively longer. Peduncle of uropod (Fig. 18E) stouter, heavily armed with several rows of spines, serrate and plumose setae (about seventy in all). Spines on inner edge of endopod much longer, not serrate. Length Adult male 12-18 mm. Ovigerous female 15-22 mm. Remarks The individuals of this species are morphologically uniform, only varying to some extent in size throughout the range, being on the whole a little larger in the south. They compare in all features with Jones’s descriptions and figures. For a discussion of the brevipes—stebbingi-africana group of species, see p. 213. Distribution Endemic to South Africa from the Cape Peninsula to Durban at depths from 17 to 100 m. Absence of records from the west coast almost certainly indicates a real distribution limit at the Cape Peninsula, as J. stebbingi has not been found on the west coast despite intensive sampling in the area. The majority of individuals was found between False Bay and Port Elizabeth, only isolated specimens occurring on the Natal coast. There is less evidence that Durban is the northern limit, however, for little material has been collected from shallow waters in northern Natal, and sampling in Mozambique and further north has been scanty or non-existent. This appears to be the second most abundant species on the southern African coasts (after I. africana), constituting more than 26 per cent of the total number of individuals in the collection. SOUTHERN AFRICAN CUMACEA: PART 2 209 Iphinoe africana Zimmer, 1908 Figs 19-20 Iphinoe africana Zimmer, 1908: 163-164, pl. 2; 1942: 190-191. Fage, 1951: 4-5. Jones, 1955: 288; 1956: 202. Iphinoe brevipes (non Hansen, 1895): Stebbing, 1910: 411; 1913: 45. Records sub- adult adult ovig. no. of 3 6 6 2 2 Juv. total records SWD_ 22°S 14°E TD tan 4 Vf 13 26 50 i\ WCD 33°S 17°E-34°S 18°E = 62-130 m 2 3 1s NAS 10 3 SB 33°S 17°E 3-29 m 199 221 61 195 206 661 1543 5S Previous records ‘Great Fish Bay’ (16°S 11°E) (Zimmer 1908); Walvis Bay (23°S 14°E) (Fage 1951); northern South West Africa to Liideritz (i9°S 12°E-25°S 14°), plankton (Jones 1955); Kunene River Mouth to Walvis Bay (17°S 11°E- 23°S 14°E), 6-100 m (Jones 1956). Syntypes Ovigerous females, deposited by Zimmer (1908) in the Berlin Zoologisches Museum. Type locality: no depth given, ‘Great Fish Bay’, near Kunene River Mouth (16°S 11°E). Description Ovigerous female, length 12,3 mm, from Saldanha Bay. Animal elongate, almost cylindrical (Fig. 19A). Integument shiny, slightly translucent, with minute reticulations visible at high magnifications. Carapace about two and one third times as long as deep with distinct middorsal carina bearing eleven teeth (number varies between four and sixteen, usually ten to twelve). Pseudo- rostral lobes vertically blunted in lateral view (Fig. 19B), short, not much produced anterior to eyelobe. Antennal notch moderately excavate, anterolateral angle acute with several small teeth below along ventral margin. Eyelobe (Fig. 19C) rounded, some reddish pigment visible well below surface, even after long preservation in alcohol; no lenses. First pedigerous somite visible dorsally and laterally, second almost as wide as deep, third and fourth produced posteriorly. Pedigerous somites together longer than carapace, cephalothorax longer than abdomen by one segment. Abdominal somites cylindrical, fifth longest. Antenna | (Fig. 19D) fairly long, first and third segments subequal in length, second a little shorter. Flagellum 1-segmented with one aesthetasc; accessory flagellum 1-segmented. Basis of maxilliped 3 (Fig. 19E) more than twice length of remaining seg- ments together; distal prolongation not greatly expanded, reaching junction 210 ANNALS OF THE SOUTH AFRICAN MUSEUM of ischium and merus. Merus somewhat expanded externally. Last three seg- ments almost cylindrical, subequal in length. Pereiopod | (Fig. 19F) elongate, basis little longer than rest of limb, serrate on distal third of inner edge. mM * x 4 \ v Ass Fig. 19. Iphinoe africana Ovigerous female. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view of carapace. D. Antenna 1. E. Maxilliped 3. F. Pereiopod 1. G. Pereiopod 2. H. Pereiopod 3. I. Telsonic somite and uropod. Scale line = 2,5 mm for A, C; 1,5 mm for B, E-F, 1; 0,5 mm for D, G-H. SOUTHERN AFRICAN CUMACEA: PART 2 Dill Pereiopod 2 (Fig. 19G) characteristically short and broad, equal in length to basis of perelopod 3, 6-segmented. Basis twice as long as broad, edged with many plumose setae. Merus broader than long with one very long spine at the expanded distal tip, reaching apex of limb. Carpus and propodus subequal in length, together equal in length to dactyl. Pereiopods 3 (Fig. 19H) to 5 similar, basis of peretopod 3 longest. Carpus of pereiopod 5 relatively longer than that of 3 or 4. Telsonic somite as wide as long at widest point, with two anal setae; in lateral view with small middorsal tooth a third from posterior end, anterior to a slight depression. Peduncle of uropod (Fig. 191) slightly longer than tel- sonic somite, subequal in length to rami, with 12 fine sharp spines on inner edge. First segment of exopod two-thirds length of second with two plumose setae on inner border; second segment with twenty-two plumose setae around entire border. First segment of endopod half length of second with five spines on inner and four on outer border; second segment with fourteen sharp spines on inner and five on outer border, plus three terminally. Adult male, length 9,2 mm, from Saldanha Bay. As female, except as follows: Mid-dorsal carina not serrate (Fig. 20A), antennal notch very shallow, anterolateral angle obsolete, but some serrations present along anteroventral margin for a short distance. Carapace relatively wider anteriorly (Fig. 20B). Second and third pedigerous somites narrower, sideplates of fourth overlapping third and fifth ventrolaterally. Abdominal sideplates defined ventrally. Carapace subequal in length to rest of thorax, cephalothorax and abdomen subequal in length. Armature of thoracic sternites simple: second forming a raised trans- verse ridge with five spines at equal intervals across the width; third and fourth forming slightly raised projections each with a forward-pointing midventral hook, fifth a low rounded projection. Aesthetascs of antenna | (Fig. 20C) annulated only proximally, accessory flagellum 2-segmented with two short aesthetascs. Pereiopods 2 to 5 less stout and more heavily armed. Pereiopod 2 longer relative to pereiopod 3. Peduncle of uropod with 35 spines of varying length in several rows on inner edge. (Fig. 20D). Second segment of exopod unarmed on outer edge. Armature of endopod much stouter, and including plumose setae. Length Adult male 8,0-11,5 mm Ovigerous female 9,0-15,2 mm Remarks Zimmer (1908) described I. africana from ‘several female individuals’. Apart from slight differences in the degree of expansion of the merus of maxilliped 3 and the carpus of pereiopod 3, the females in the present collection are identical with Zimmer’s description and figures. Except for the second pereiopod and uropod figured by Fage (1951), the male has not previously PAD ANNALS OF THE SOUTH AFRICAN MUSEUM \ N Nt Fig. 20. Iphinoe africana Adult male. A. Lateral view. B. Detail of eyelobe. C. Detail of distal tip of antenna 1. D. Uropod. Scale line = 2 mm for A; 1 mm for B; 0,5 mm for D; 0,3 mm for C. been figured or described. Note: the number of teeth on the middorsal carina is related to age and sex. Juveniles of indeterminate sex have between two and seven teeth, the number increasing in females to between seven and ten in young adults and between ten and sixteen in ovigerous females. In males the reverse is true, two or three teeth occurring in young males with asetous pleopods, while in fully adult males there are no teeth at all. Distribution I. africana is endemic to the cold south-western coast of Africa from the Kunene River Mouth to Cape Point. Sufficient collecting has been done on either side of this region to make it probable that the species is indeed confined SOUTHERN AFRICAN CUMACEA: PART 2 213 to this rather narrow range. It is apparently restricted to areas with a mean annual temperature of less than 20°C. The depth range is 3 to 130 m. It is the most abundant species in these waters, constituting almost 35 per cent of the individuals in the collection. The brevipes-africana—stebbingi group I. brevipes was described by Hansen in 1895 from the Gulf of Guinea in west Africa and J. africana by Zimmer (1908) from the Great Fish Bay in southern Angola. Stebbing (1910, 1913) was of the opinion that they were too similar to warrant the maintenance of two species, and referred all specimens (including some from South Africa) to J. brevipes. Zimmer (1916, 1942) main- tained not only that these two species were distinct from one another, but that a third species would probably have to be erected for the South African material. This was in fact done by Jones (1956), who gave the name I. stebbingi to the new species. Although distinct from one another, the three species fall into a sub- group within the genus, since they share a number of characters not found in the other members of the genus, in particular the shortness of pereiopod 2 and the elongate, almost cylindrical carapace. Their distinctness from each other is evident ecologically as well as morphologically, since the distribution of the three species is disjunct. They may be distinguished from each other as follows: I. brevipes pseudorostral lobes pointed anteriorly in lateral view antennal notch small, con- fined to ventral part of pseudorostrum anterolateral tooth small and blunt with ventral serrations basis of pereiopod 1 one and a half times length of rest of limb basis of pereiopod 2 as long as broad telsonic somite rounded posteriorly with two small teeth in midline maximal length of adult female 10 mm, of adult male about 8,6 mm serrations on middorsal carina in both sexes southern limit 5°S I. africana pseudorostral lobes bluntly truncate anteriorly antennal notch of moderate size, confined to ventral part of pseudorostrum anterolateral tooth small, pointed, with serrations above and below basis of pereiopod 1 equal in length to rest of limb basis of pereiopod 2 twice as long as broad telsonic somite bluntly truncate posteriorly with two very small teeth in midline length of adult female 9-15 mm, of adult male 8,0-11,5 mm serrations on middorsal carina in juveniles and adult females only northern limit 17°S, eastern limit 18°E I. stebbingi pseudorostral lobes some- what truncate anteriorly antennal notch large, not confined to ventral part of pseudorostrum anterolateral tooth long and pointed, reaching beyond anterior tip of pseudo- rostrum, without serrations basis of pereiopod 1 twice length of rest of limb basis of pereiopod 2 twice as long as broad telsonic somite rounded posteriorly, without teeth minimal length of adult female 15 mm, of adult male 12 mm middorsal serrate carina never western limit 18°E 214 ANNALS OF THE SOUTH AFRICAN MUSEUM Iphinoe producta sp. nov. Fig. 21 Records LBT 32°S 17°E 200-280 m _ 1 subadult g, 3 ovig. 92 Ijuv. (3records) Holotype Ovigerous female, in the South African Museum, SAM-A15494, collected during the UCT benthic survey, 24 September 1971. Type locality: 200 m, off Lambert’s Bay (32°04’S 17°12’E). UCT station number LBT 67D. Description Ovigerous female, holotype, \length 9,3 mm. Very slender and elongate. Integument translucent, slightly calcified, very finely reticulate. Anterior three- quarters of middorsal carina of carapace bearing well-developed forward- pointing denticles (Fig. 21A). Pseudorostral lobes (Fig. 21B) about a seventh of total length of carapace. Eyelobe narrow with three very small lenses (Fig. 21C). Anterolateral angle acute, antennal notch distinct but small, confined to ventral half only. Carapace two and a half times as long as deep, slightly longer than free thoracic somites together. Cephalothorax subequal in length to abdomen. First pedigerous somite visible dorsally and laterally, second slightly wider than third. Abdominal somites cylindrical, lacking defined sideplates ventrally. Middorsal carina present up to and including third pleon somite. Basal segments of antenna | (Fig. 21D) subequal in length. Accessory flagellum short, 2-segmented. Flagellum 2-segmented with one long aesthetasc. Basis of maxilliped 3 (Fig. 21E) more than two and a half times length of remaining segments together, distal prolongation reaching beyond junction of merus and carpus (Fig. 21F). Merus short and slightly expanded externally. Pereiopod | (Fig. 21G) very slender and elongate, basis slightly shorter than rest of limb with a few stout spines distally along outer edge. Carpus, propodus and dactyl all very slender, more or less subequal in length. Pereiopod 2 (Fig. 21H) fairly stout, 6-segmented, slightly shorter than posterior pereiopods. Basis equal in length to carpus, propodus and dactyl together. Merus and carpus stout, subequal in length. Pereiopods 3 (Fig. 211) to 5 similar, basis of pereiopod 3 longest, merus and carpus of pereiopod 5 longest. Telsonic somite slightly produced between uropods (Fig. 21J). Peduncle of uropod a little longer than telsonic somite with eleven blunt spines on inner edge. Exopod slightly longer than endopod, first segment unarmed, a third length of second; second with eleven plumose setae on inner edge and six spines terminally, three very long. First segment of endopod less than half length of second, with five spines on inner edge; second with eleven small spines on inner edge and four long ones terminally. SOUTHERN AFRICAN CUMACEA: PART 2 DAS) A single damaged subadult male was taken at the same station as the holotype female. It appears to be similar in most details, but is not sufficiently whole to allow an adequate description. Its length is approximately 7,5 mm. Fig. 21. Iphinoe producta sp. nov. Ovigerous female, holotype. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view of carapace. D. Antenna 1. E. Maxilliped 3. F. Detail of distal tip of maxilliped 3. G. Pereiopod 1. H. Pereiopod 2. I. Pereiopod 3. J. Telsonic somite and uropod. Scale line = 2 mm for A, C; 1 mm for B, E, G, J; 0,5 mm for D, F, H-I. 216 ANNALS OF THE SOUTH AFRICAN MUSEUM Length Ovigerous female 8,6—-9,5 mm. Remarks I. producta is undoubtedly a member of the genus Jphinoe, most closely resembling J. ischnura Zimmer, 1952, from Indo-China. Apart from other minor differences, the two species may be distinguished by the more elongate carapace with more numerous serrations, the longer bases of pereiopods 2 to 5 and the longer first segment of the endopod of the uropod in J. producta. I. ischnura reaches 4,5 mm in length and J. producta 9,6 mm. It may be distinguished from J. dayi Jones, 1960, by the more robust form and the presence of two aesthetascs on the flagellum of the first antenna in the latter species, and from I. tenella Sars, 1878, I. elisae Bacescu, 1950, and J. serrata (Norman, 1867) by the shorter carapace in these three species. Distribution Five specimens known, all from the south-western coast of South Africa at depths between 200 and 280 m. Iphinoe dayi Jones, 1960 Figs 22-23 Iphinoe dayi Jones, 1960: 175-177, fig. 2. Records sub- adult adult Oovig. no. of 3 6 6 & 2 juve totalitecords WCD 33°S 18°E 65 m 1 1 1 FAL & FBY 34°S 18°E 23-87 m 28 LO 162224 ee SST 34°S 21°E 80 m y 1 7 1 2 Saal 4 SCD 34°S 25°E-33°S 27°E 26-84 m 3° 23 6 4 SAM ? ? 1 1 1 Previous records False Bay (34°S 18°E), 20-58 m (Jones 1960). Holotype Adult male, designated by Jones (1960), in the British Museum (Natural History). Type locality: 20 m, False Bay (34°S 18°E). Description Ovigerous female, length 9,6 mm, from False Bay. Integument finely reticulate, neither shiny nor translucent. Carapace somewhat compressed laterally, middorsal carina bearing eight teeth (varies between eight and ten) on anterior half (Fig. 22A). Carapace twice as long as deep, pseudorostral lobes SOUTHERN AFRICAN CUMACEA: PART 2 Dil Fig. 22. Iphinoe dayi Ovigerous female. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view of carapace. D. Antenna 1. E. Maxilliped 3. F. Pereiopod 1. G. Pereiopod 2. H. Pereiopod 3. I. Telsonic somite and uropod. Scale line = 2 mm for A, C; 1 mm for B, F-I; 0,5 mm for D-E. 218 ANNALS OF THE SOUTH AFRICAN MUSEUM well developed, produced beyond eyelobe for about one-seventh of total length of carapace. Anterolateral angle (Fig. 22B) rounded, bearing several small teeth. Antennal notch deeply excavate, set back from pseudorostral lobes. Eyelobe somewhat elongate, eyeless (Fig. 22C). First pedigerous somite visible dorsally and laterally, second and third of equal width, third produced backwards to overlap fourth. Carapace slightly longer than free thoracic somites together. Cephalothorax longer than abdomen by one somite. Abdominal somites cylindrical. Antenna | (Fig. 22D) fairly short, basal segment half as wide as long, edged with denticles. Flagellum short, 2-segmented, with two aesthetascs and one fine seta. Accessory flagellum short, 1-segmented. Basis of maxilliped 3 (Fig. 22E) curved, two and a half times length of remaining segments together; distal prolongation elongate, almost reaching distal tip of carpus. Merus short, little expanded, wider and denticulate distally. Carpus inserted on inner half of merus only. Basis of pereiopod | (Fig. 22F) equal in length to rest of limb. Ischium half length of merus. Last three segments subequal in length. Pereiopod 2 (Fig. 22G) 6-segmented, very slightly shorter than pereiopod 3. Basis shorter than rest of limb, dactyl furnished with a number of short blunt spines. Pereiopods 3 (Fig. 22H) to 5 similar, pereiopod 3 longest. Telsonic somite somewhat produced between uropods, less than one and a half times as long as wide, about two-thirds length of peduncle of uropod. Peduncle (Fig. 221) with fifteen short, blunt spines on inner edge. Exopod slightly longer than endopod, two-thirds length of peduncle. First segment a third length of second, unarmed; second with five plumose setae on inner edge and five terminal spines. Segments of endopod subequal in length, each with six spines on inner edge, second also with two spines terminally. Adult male, length 8,9 mm, from False Bay. As female, except as follows: teeth of middorsal carina smaller and between five and seven in number (Fig. 23A). Anterolateral angle obsolete, antennal notch shallow (Fig. 23B) with a few small serrations below. Eye present (Fig. 23C) in the form of two small lenses and a little pigment (although in some the eye is well developed). Side- plates of fourth pedigerous somite overlapping third anteriorly and fifth posteriorly. Abdominal sideplates defined ventrally. Thoracic sternites simple, that of first pedigerous somite forming a single mid-ventral spine and of third and fifth forming small rounded projections (no armature on second or fourth). Merus of maxilliped 3 (Fig. 23D) a little more expanded, carpus and propodus narrower. Peduncle of uropod (Fig. 23E) stouter, with about forty short blunt spines and serrate setae in several rows on inner edge. Second segment of exopod with seven plumose setae on inner edge and three long serrate setae terminally. First segment of endopod with twelve spines of various types, unevenly spaced; second with an even row of slender spines on inner edge and two stout serrate setae terminally. SOUTHERN AFRICAN CUMACEA: PART 2 219 Fig. 23. Iphinoe dayi Adult male. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view of cara- pace. D. Distal tip of maxilliped 3. E. Uropod. Scale line = 4 mm for A; 2 mm for B—C; 0,5 mm for D-E. Length Adult male 8,6-10,6 mm Ovigerous female 8,3-11,2 mm Remarks Only the male of this species has previously been described (Jones 1960). The present specimens have been compared with paratypes identified by Jones, and there are no differences of any degree, apart from the varying degree of 220 ANNALS OF THE SOUTH AFRICAN MUSEUM development of the eye in adult males. A much-mutilated specimen of J. dayi, now in the possession of the South African Museum, was tentatively labelled by Stebbing (unpublished) as J. tenella, and the two species are not dissimilar. But J. dayi may be distinguished from all other species having a serrated mid- dorsal carina in the male as follows: I. tenella, I. ischnura, I. elisae and I. producta have only one aesthetasc on the flagellum of the first antenna and J. serrata is serrate for the whole length of the mid-dorsal carina. J. dayi may further be distinguished from J. producta, the only other species from southern Africa in which the male is serrate, by the greater length of the carapace in J. producta and the generally far more slender body and limbs. Distribution Apparently endemic to South Africa from the Cape Peninsula to East London at depths from 23 to 87 m. The relative rarity of this species means that the limits of its range can be determined with less finality than can those of I. africana and I. stebbingi. Nevertheless, it is fairly common in those areas where it is known to occur, and accounts for more than 3 per cent of the total number of individuals in the collection. Iphinoe fagei Jones, 1955 Figs 24-25 Iphinoe fagei Jones, 1955: 285-287, figs 3-4; Jones, 1956: 199. Records sub- adult adult ovig. no. of 3 é& & Q Q juv. total records SWD_ 26°S 15°E 26 m 8 8 1 WCD 32°S 17°E-32°S 18°E 11-172m 15 Pe 2 als S24 aos) 6 Previous records South West Africa (south of Walvis Bay) (22°S 14°E-25°S 14°E), plankton (Jones 1955); South West Africa (23°S 14°E), 22-76 m (Jones 1956). Holotype Not designated. Type locality: plankton, South of Walvis Bay (22°S 14°E). Description Ovigerous female, length 6,9 mm, from St Helena Bay (32°S 18°E). Integu- ment translucent, finely reticulate, appearing slightly crystalline. Carapace oval with two (varying between none and three) small teeth on middorsal carina, little over a third of the distance from anterior tip (Fig. 24A). Middorsal carina present only on anterior two-thirds of carapace. Pseudorostral lobes meeting for a short distance in front of eyelobe (Fig. 24B). Eyelobe (Fig. 24C) rounded SOUTHERN AFRICAN CUMACEA: PART 2 DAI Fig. 24. Iphinoe fagei Ovigerous female. A. Lateral view. B. Anterior tip of carapace. C. Dorsal view of carapace. D. Antenna 1. E. Maxilliped 3. F. Detail of distal tip of maxilliped 3. G. Pereiopod 1. H. Pereiopod 2. I. Pereiopod 3. J. Telsonic somite and uropod. Scale line = 2 mm for A, C; 1 mm for B, E; 0,5 mm for D, F-J. with three (or more) small, indistinct lenses, slightly pigmented. Anterolateral angle acute with small tooth at apex, antennal notch excavate, angular. Carapace twice as long as deep, longer than free thoracic somites together. First pedigerous somite exposed for a short distance, second broad. Cephalo- DO), ANNALS OF THE SOUTH AFRICAN MUSEUM thorax little longer than abdomen; abdominal somites cylindrical. Antenna 1 (Fig. 24D) of moderate length, first segment longest and second shortest. Flagellum 2-segmented with one aesthetasc; accessory flagellum small, 1-segmented. Basis of maxilliped 3 (Fig. 24E) more than two and a half times length of remaining segments together; distal prolongation narrow, nearly reaching distal tip of carpus. Ischium (Fig. 24F) slightly longer than wide, merus pro- duced externally, tip reaching half-way along carpus. Pereiopod | (Fig. 24G) elongate, basis curved, subequal in length to rest of limb. Last three segments fairly stout, subequal in length. Pereiopod 2 (Fig. 24H) stout, 6-segmented. Basis slightly shorter than rest of limb. Dactyl fairly long, strongly armed at tip. Pereiopods 3 (Fig. 241) to 5 similar, basis of pereiopod 3 longest. Telsonic somite slightly produced between uropods (Fig. 24J), more than half as long as penducle. Peduncle nearly twice length of rami with twenty-one fine spines on inner edge. First segment of exopod shorter than second, unarmed; second with eight plumose setae on inner edge and three terminal spines. First segment of endopod shorter than second with four spines on inner edge; second with nine short spines on inner edge and two long ones terminally. Adult male, length 7,5 mm, from St Helena Bay. As female except as follows: integument thinner, carapace (Fig. 25A) without middorsal carina; antennal notch much shallower (Fig. 25B), anterolateral angle obtuse, poorly defined. Eye present (Fig. 25C), consisting of three to five large lenses (distinct if white pigment present below, otherwise difficult to see). Sideplates present on all abdominal somites, fourth thoracic sideplate overlapping third and fifth. First pedigerous somite visible only dorsally and dorsolaterally. Sternite of first pedigerous somite armed with three plumose setae pointing posteriorly on either side; second with large rounded projection bearing forward-directed tooth; third with slight transverse ridge; fourth unarmed; fifth with small rounded projection. Flagellum of antenna | (Fig. 25D) surrounded by several short aesthetascs. Basis of pereiopod | armed with ten spines proximally on lower edge. Pro- longation of basis of maxilliped 3 somewhat longer (Fig. 25E). Merus of pereiopod 2 (Fig. 25F) longer. Peduncle of uropod (Fig. 25G) more than one and a half times length of rami, with about forty-two spines on inner edge. Second segment of exopod with about ten plumose setae on inner edge and four spines terminally. First segment of endopod with twelve spines on inner edge, second with thirteen, plus two larger ones subterminally and three plumose setae terminally. Length Adult male 6,7-8,3 mm Ovigerous female 6,7—9,8 mm SOUTHERN AFRICAN CUMACEA: PART 2 223 Remarks I. fagei was first described by Jones (1955) from plankton collected by the R.R.S. William Scoresby off the coast near Walvis Bay. Further benthic samples were collected by the Galathea, also off Walvis Bay, and identified by Jones (1956). The author has examined some ovigerous females from the latter collection and finds that they agree in morphological detail with those of the present collection. However, the Galathea specimens are transparent and slightly smaller, whereas those of the author are quite translucent and brittle due to a very much thicker integument. The adult males in the present collection agree with Jones’s description and figures. Most females may be distinguished from other females in the genus by the presence of one to three small teeth behind the eyelobe, but the teeth are absent WY AL "WU D 4) Vy HF, Ly Ni 4 i) yt i ’ ¢ : ORO SS . y : \ \ RS. N WSN} WSS eS SSS SOAS OSS SST oe <= aes ene DZ LE COE IE LIYUTY Hf Fig. 25. Iphinoe fagei Adult male. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view of carapace. D. Antenna 1. E. Distal tip of maxilliped 3. F. Pereiopod 2. G. Uropod. Scale line = 2 mm for A, C; 1 mm for B; 0,5 mm for D-G. 224 ANNALS OF THE SOUTH AFRICAN MUSEUM in some specimens. The latter may be distinguished by the smooth carapace which is almost exactly twice as long as deep, and the similar lengths of pereio- pods 2 and 3. The species most closely resembling J. fagei is I. senegalensis Jones, 1956, from the shore in west Africa. The two may be distinguished primarily by the longer basis of maxilliped 3 and the greater body size of I. fagei. Distribution Apparently endemic to the south-western coast of southern Africa from Walvis Bay to St Helena Bay, at depths from 11 to 172 m, and constituting a little over 2 per cent of the individuals in the collection. Its distribution appears to be patchy, a few hauls yielding large numbers of individuals. Iphinoe senegalensis Jones, 1956 I. senegalensis Jones, 1956: 200-201, figs 7-8; Jones 1960: 175. A single adult male from an earlier UCT collection (not in the author’s possession) was identified by Jones (1960) as J. senegalensis. The position (32°S 18°E) at a depth of | m suggests that the specimen came from Langebaan Lagoon. As Jones pointed out, it is not easy to identify with confidence a single adult male of this genus, and confirmation of the occurrence of J. senegalensis in South Africa will have to await the collection of more material. The species, which previously has been found only on the beach at Goreé in Senegal, is very similar to J. fagei. Differences between the two are mentioned in the dis- cussion of I. fagei above. Iphinoe crassipes Hansen, 1895 Figs 26-27 Iphinoe crassipes Hansen, 1895: 53-54, pl. 4 (figs 44f). Stebbing, 1910: 412-413, pl. 45; 1913: 43-44, figs 21-22. Fage 1928: 331. Zimmer, 1942: 191-192. Kurian, 1951: 84-86; 1954: 276. Jones, 1956: 202, fig. 9. Bacescu, 1961: 501, fig. 4. LeLoeuff & Intes, 1972: 43. Iphinoe macrobrachium Calman 1904a: 173, pl. 4 (figs 72-75). Records sub- adult adult Oovig. no. of 3 $$ & 2 2 juve totaltreconds SB 33°S 18°E 13-29 m 1 A 2) 6) seals 4 FAL & FBY 34°S 18°E 16-39 m 4 le al 33 IS) 35 7 SCD 34°S 21°E-33°S 25°E 44-79 m 8 3 1 3 1 3 Al 7 SAM A 688 33°S 26°E 92 m 1 1 1 NIWR 30°S 30°E-28°S 32°E _23-103m _ 12 7 SO AS Sie 0 20 Previous records Gulf of Guinea, plankton (Hansen 1895); Ceylon, 8-14 m (Calman 1904a); South Africa (East London) (32°S 28°E), 75 m (Stebbing 1910); tropical west SOUTHERN AFRICAN CUMACEA: PART 2 DD5 Africa (Fage 1928; Jones 1956; LeLoeuff & Intes 1972); India, 6-32 m (Kurian 1951, 1954); Red Sea (Bacescu 1961). Holotype Immature male, length 3,2 mm, unique. Type locality: Gulf of Guinea. Description Ovigerous female, length 7,2 mm, from the south coast near Knysna. Integument rather thin and delicate, little calcified, somewhat transparent. Reticulations and pits visible at high magnifications producing a finely crystalline appearance. Carapace little more than one and a half times as long as deep with several shallow longitudinal furrows (Fig. 26A). Middorsal carina forming double row of very small serrations on middle part of carapace only. Eyeless eyelobe elevated above pseudorostral lobes in lateral view (Fig. 26B). Pseudorostral lobes short, rounded, slightly upturned and truncate anteriorly, curled inwards in dorsal view (Fig. 26C). Anterolateral angle acute with several small serrations below. Antennal notch rounded, of moderate size. Five free pedigerous somites, together as long as carapace; first narrow, second wider ventrally. Cephalothorax equal in length to first five abdominal somites. Abdominal somites almost cylindrical with sideplates poorly defined posteroventrally on each somite. Antenna 1 of moderate length (Fig. 26D), second segment about half length of first or third. Flagellum 2-segmented with two slender spines. Acces- sory flagellum 1-segmented, without setae. Basis of maxilliped 3 (Fig. 26E) very wide and short, about one and a half times length of remaining segments together, serrate on inner edge. Distal prolongation particularly long, narrowed distally, reaching half-way along propodus. Merus also much expanded, distal edges parallel for some distance, reaching more than half-way along propodus. Basis of pereiopod 1.(Fig. 26F) particularly short, equal in length to next three segments together, slightly curved. Carpus wide, slightly longer than subequal propodus and dactyl. Pereiopod 2 (Fig. 26G) relatively short and stout, basis very short, subequal in length to merus, carpus and propodus together. Dactyl stout, strongly armed, subequal in length to basis. Pereiopods 3 (Fig. 26H) to 5 similar, merus and carpus stout, carpus strongly armed distally. Telsonic somite (Fig. 261) well produced between uropods, with two fine anal setae. Peduncle of uropod little longer than telsonic somite, armed with seven evenly-spaced stout spines on inner edge. Exopod of uropod two-thirds length of endopod, only slightly longer than first segment of endopod. First segment of exopod half length of second, unarmed; second segment armed with two very small spines on inner edge and six long stout ones terminally. First 226 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 26. Iphinoe crassipes Ovigerous female. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view of carapace. D. Antenna 1. E. Maxilliped 3. F. Pereiopod 1. G. Pereiopod 2. H. Pereiopod 3. I. Telsonic somite and uropod. Scale line = 2 mm for A; 1 mm for B-C, F; 0,5 mm for D-E, G-I. SOUTHERN AFRICAN CUMACEA: PART 2 Di segment of endopod nearly twice length of second, armed with four spines on inner edge and one distally on outer edge. Second segment narrower than first with four strong distal spines. Adult male, length 7,8 mm, from UCT’s SCD programme near Knysna. As female, except as follows: anterolateral angle and antennal notch wanting (Figs 27A & B). Serrations of middorsal carina of carapace much less distinct. Eye (Fig. 27C) well developed, consisting of a single large central lens surrounded by eight smaller ones, with pigment below. Antenna 1 (Fig. 27D) stouter, flagellum bearing about nine short aesthetascs. Prolongation of basis of maxilliped 3 (Fig. 27E) shorter and broader, of merus wider, distally reaching articulation of propodus and dactyl. Peduncle Fig. 27. Iphinoe crassipes Adult male. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view of carapace. D. Antenna 1. E. Distal tip of maxilliped 3. F. Uropod. Scale line = 2 mm for A; 1 mm for B—C; 0,5 mm for E-F; 0,3 mm for D. 228 ANNALS OF THE SOUTH AFRICAN MUSEUM of uropod (Fig. 27F) armed with fourteen spines on inner edge and five small plumose setae distally. First segment of exopod shorter relative to second with six very long plumose setae on inner edge. First segment of endopod with eleven stout spines on inner edge, second with seven. Length Adult male 5,/—8,1 mm Ovigerous female 5,6-7,8 mm Remarks The relationship between J. crassipes Hansen, 1895, and I. macrobrachium Calman, 1904a, has never been satisfactorily determined. J. crassipes was described on the basis of a single immature male, 3,2 mm in length, from west Africa. In 1904 Calman described specimens from Ceylon, for which he erected a new species, J. macrobrachium. (In his paper he gives the length as | mm, but in fact both are about 4,5 mm in length.) He considered his species to be distinct from J. crassipes, differing mainly in ‘the outline of the carapace’, the length of the basis of pereiopod | and the unequal lengths of the rami of the uropods in his specimens. Stebbing (1910) described a single adult male from South Africa which he called I. crassipes (although it exhibited some of the characters used by Calman to differentiate [. macrobrachium from Hansen’s species), considering the differences between Calman’s and Hansen’s specimens to be due solely to sexual dimorphism between obviously immature individuals. The author does not have Stebbing’s specimen, but it is presumably similar to those in her possession, which are very uniform in structure throughout the range. His figures do not correspond well with any specimens, particularly in the outline of the carapace, the basis of pereiopod 1 and the uropod. Since then Fage (1928), Jones (1956) and LeLoeuff & Intes (1972) have all recorded specimens referred to as JI. crassipes from west Africa, Jones figuring the previously undescribed female. Kurian (1951, 1954) has recorded some speci- mens from India and Bacescu (1961) has erected a new subspecies, I. c. haifae, for material from the Red Sea. The question still remains as to whether Stebbing was correct in assuming I. crassipes and I. macrobrachium to be synonymous. The author has examined Calman’s two syntypes and several specimens from P. LeLoeuff’s west African material. Despite the striking differences between the figures given by different authors, particularly those of the carapace, the external appearance of the animals is rather uniform throughout the range. In all specimens examined the carapace is characterized by shallow longitudinal furrows on the anterior part. Individuals differ mainly in that in some the eyelobe is elevated above the level of the pseudorostrum, and in others the two are flush with each other; in some the pseudorostrum is clearly upturned and truncate anteriorly while in others it is more or less anteriorly directed and rounded. In all cases the pseudorostal lobes are curled inwards and downwards in dorsal view, with a number of downward-directed spines. The other major variations occur in the first 229 SOUTHERN AFRICAN CUMACEA: PART 2 pereiopods and uropods. The basis of pereiopod | varies between a third and nearly half the total length of the limb; the exopod of the uropod may be two- thirds the length of the endopod or the rami may be subequal in length; the first segment of the endopod may be twice the length of the second or the two segments may be of equal length. Since these variations are not found uniformly throughout the geographical range, the author has come to the conclusion—after some deliberation—that the differences exhibited between individuals from different areas are not con- sistent enough to warrant the existence of two species or even subspecies. Thus Stebbing was correct in his conclusion although, contrary to his suggestion, few of the differences are sexual. The author also suspects that, from his figures and very brief description, Bacescu’s J. c. haifae will be found to fit within the range of variability of I. crassipes without subspecific differentiation, being very like the West African forms in most respects. The major variable characters shown by individuals from different regions are tabled below. Ceylon and India (UI. macrobrachium sensu Calman) eyelobe strongly ele- vated in 2 (no ¢ available) pseudorostrum upturned in 2 distal process of basis of maxilliped 3 reaching distal tip of carpus basis of pereiopod 1 three-sevenths total length of limb endopod of uropod one and a_ third length of exopod first segment of endo- pod slightly longer than second peduncle of uropod shorter than endopod South Africa U. crassipes sensu Stebbing) eyelobe strongly ele- vated in 9, slightly in 3 pseudorostrum upturned in 9, very slightly in 3 distal process of basis of maxilliped 3 reaching at least distal tip of carpus basis of pereiopod 1 three-fifths total length of limb endopod of uropod one and a_ half length of exopod first segment of endopod twice length of second peduncle and endo- pod subequal in length Red Sea (subspecies I. c. haifae sensu Bacescu) ? eyelobe elevated in ¢ and 92 pseudorostrum rounded, ? not up- turned distal process of basis of maxilliped 3 reaching _half-way along carpus basis of pereiopod 1 half total length of limb West Africa UI. crassipes sensu Hansen) eyelobe elevated in 6, less in 2 pseudorostrum upturned in dg, rounded in 2 distal process of maxilliped 3 reach- ing one-third length of carpus basis of pereiopod 1 less than half total length of limb exopod and endopod of uropod subequal in length segments of endopod subequal in length penducle of uropod longer than endopod It can be seen that in most respects the west African and Red Sea forms are rather similar, as are the South African and Ceylonese forms, but the degree of overlap of distinguishing characters is such that taxonomic differentiation is inappropriate. I. crassipes is obviously closely related to J. pokoui LeLoeuff & Intes, 1972, from West Africa, the only other species bearing long stout first pereiopods and an upturned pseudorostrum. It may be distinguished from J. crassipes by 230 ANNALS OF THE SOUTH AFRICAN MUSEUM the very wide antennal notch with a narrow serrate elongate anterolateral angle and more slender uropods with subequal rami. Both species are rather aberrant for the genus, the pereiopods and maxilliped 3 having some similarity with those of Eocuma. The nature of the carapace and thoracic somites, how- ever, ensures that they remain in the genus Jphinoe. Distribution India; Ceylon; South Africa: Saldanha Bay (13-29 m) to Natal (23-103 m); Gulf of Guinea; Red Sea. The species is absent from the colder waters of South West Africa, but otherwise probably occurs round the entire Atlantic and Indian Ocean coasts of Africa and India from about 8 to little over 100 m in depth. It constitutes a little more than 3 per cent of the individuals in the collection, but represents more than 70 per cent of the individuals from Natal waters. Iphinoe truncata Hale, 1953 Fig. 28 Iphinoe truncata Hale, 1953: 48-50, figs 3-4. Records sub- adult adult ovig. no. of 6 3S 2 g&8F juv. total records CON Morrumbene (23°S 35°E) D) 3 9 14 5 PEM Richards Bay (28°S 32°B) 1 1 1 RU _ St Lucia (28°S 32°B) estuarine D 5 5 1 Yrs 1 UCT Keurboom’s River (34°S 23°E) ( plankton 10 4415) 9 543 D UCT Knysna (34°S 23°E) D D 4 1 UCT Great Brak River (34°S 22°E) 1 1 1 Previous records Estuarine plankton from: The Haven (32°S 28°E), Port St Johns (31°S 29°E), Umkomaas (30°S 30°E) (Hale 1953); Knysna (34°S 23°E) (Jones 1960). Holotype Not designated. Type localities: ovigerous female, Umkomaas River Mouth; adult male, Port St Johns River Mouth; both in sand, estuarine. Description Ovigerous female, length 2,8 mm, from Keurboom’s River Mouth. Integu- ment translucent, finely spotted with black chromatophores. Carapace one and two-thirds as long as deep (Fig. 28A). Anterolateral angle very small, acute; antennal notch tiny, too small to accommodate first antenna. Distinct mid- dorsal carina present on carapace (Fig. 28B), widening slightly about half-way back along the carapace, and absent on posterior third. Pseudorostral lobes short, barely meeting in front of eyelobe. Eyelobe rounded, eye consisting of large patch of black pigment; no lenses visible. SOUTHERN AFRICAN CUMACEA: PART 2 235i Fig. 28. Iphinoe truncata Ovigerous female. A. Lateral view. B. Dorsal view of carapace. C. Antenna 1. D. Maxilliped 3. E. Pereiopod 1. F. Pereiopod 2. G. Pereiopod 3. H. Telsonic somite and uropod. Adult male. I. Lateral view. J. Dorsal view of carapace. K. Detail of distal tip of antenna 1. L. Maxilliped 3. M. Pereiopod 1. N. Telsonic somite and uropod. Scale line = 1 mm for A-B, J; 0,5 mm for C-J, L-N; 0,1 mm for K. First pedigerous somite visible, second very wide. Carapace almost as long as free thoracic somites together, cephalothorax as long as abdomen and peduncle of uropod together. Marsupium very large. Exopods of maxilliped 3 and pereiopod 1 large and reflexed upwards. D3) ANNALS OF THE SOUTH AFRICAN MUSEUM Antenna | (Fig. 28C) fairly long and slender, first segment of basal region short, subequal in length to third. Accessory flagellum very small, 1-segmented. Flagellum 2-segmented, bearing two aesthetascs and a number of fine setae. Maxilliped 3 (Fig. 28D) short and stout, basis a third as wide as long; prolongation narrow, reaching junction of merus and carpus. Merus and carpus both somewhat expanded. Basis of pereiopod | (Fig. 28E) stout, very slightly longer than rest of limb. Remaining segments also stout, particularly ischium and merus. Pereiopod 2 (Fig. 28F) 6-segmented, slightly shorter than pereiopod 3, last four segments subequal in length. Pereiopods 3 (Fig. 28G) to 5 similar, slender. Basis of pereiopod 3 subequal in length to rest of limb, bases of others relatively shorter. Telsonic somite (Fig. 28H) almost square in dorsal view, notched in mid- line. Peduncle of uropod slightly longer than telsonic somite with eight small serrated spines on inner edge. Rami subequal in length. First segment of exopod about half length of second, unarmed; second armed with six very slender spines on inner edge and three stouter terminal spines, one very long. Segments of endopod subequal in length, first armed with four spines on inner edge, second with three on inner edge and two terminally. Adult male, length 1,7 mm, from Keurboom’s River Mouth. As in female, except as follows: much smaller in size, fewer chromatophores present. Carapace (Fig. 281) almost twice as long as deep; anterolateral angle wanting; no antennal notch. First pedigerous somite visible only dorsally, second much narrower. Anterior part of carapace (Fig. 28J) relatively wider. Flagellum of antenna | with five aesthetascs, three around base (Fig. 28K). Basis of maxilliped 3 (Fig. 28L) somewhat narrower, less angled; prolongation narrower. Basis of pereiopod | (Fig. 28M) slightly narrower, ischium and merus less stout. Peduncle of uropod (Fig. 28N) armed with twelve spines, second segment of exopod with three spines, second segment of endopod serrated on inner edge with two terminal spines, one constricted half-way along and serrated distally. Length Adult male 1,7—2,5 mm Ovigerous female 2,5—3,1 mm Remarks This species has previously been recorded by Hale (1953) and Jones (1960). Hale described it on the basis of adult females only, although he did also desig- nate an adult male holotype without description. The present specimens are clearly members of Hale’s species, not differing from those figured by him in any significant way. This species can be distinguished from others in the genus by the absence of a serrated dorsal carina in conjunction with the rami of the uropods being subequal in length to the peduncle and the very short, stout SOUTHERN AFRICAN CUMACEA: PART 2 233 bases of maxilliped 3 and pereiopod 1. The adult males are all considerably smaller than the ovigerous females, and the size of these females varies rather more than is common in the genus. Distribution The only bodotriid known in estuaries from Great Brak River to Morrum- bene, where it is found in fairly small numbers. Iphinoe capensis (Zimmer, 1921) Fig. 29 Bodotria capensis Zimmer, 1921: 123-124, figs 12-14. Iphinoe brevidactyla Hale, 1953: 145-148, figs 1-2. Records sub- adult adult ovig. no. of 3g 3 OQ BegQ juny, total records SWD 26°S 15°E 26 m 1 D 3 1 LBT 32°S 18°E 10-13 m 9 3 1 13 8 LB 33°S 18°9E O2m 5 D 1 3 11 6 @P* B47S 18sE) . 022m 1 1 1 FAL & FBY 34°S18°E 0-23m 71 2 1 4 4 82 Ans NIWR 29°S 31°E 30 m 1 1 1 *Kommetjie **Two records from light-traps Previous records ‘Cape Town’ (Zimmer 1921); Langebaan Lagoon (33°S 18°E), 1-3 m (Hale 1953). Syntypes Two adult males, deposited by Zimmer (1921) as Bodotria capensis in the Berlin Zoologisches Museum. Type locality: ‘Cape Town’. Description Ovigerous female, length 3,1 mm, from Kalk Bay Harbour, False Bay. Integument translucent, shiny. Carapace (Fig. 29A) more than one and a half times as long as deep, anterolateral angle obtuse, small; antennal notch very shallow (Fig. 29B). Pseudorostral lobes short, rounded. Eyelobe rounded, eye large and distinct with maroon pigment below; three lenses visible (Fig. 29C). First pedigerous somite very short, second very wide. Carapace subequal in length to free thoracic somites, cephalothorax longer than abdomen by two somites. Abdominal somites cylindrical. Antenna | (Fig. 29D) short; first segment longest and second shortest. Accessory flagellum very small; flagellum short, 1-segmented, with a single short aesthetasc. 234 ANNALS OF THE SOUTH AFRICAN MUSEUM Maxilliped 3 (Fig. 29E) stout, basis angled; prolongation short and blunt, hardly reaching level of articulation of ischium and merus. Merus, carpus and propodus all flattened and of equal width. Dactyl short and cylindrical. Fig. 29. Iphinoe capensis Ovigerous female. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view of carapace. D. Detail of distal tip of antenna 1. E. Maxilliped 3. F. Pereiopod 1. G. Pereiopod 2. H. Pereiopod 3. I. Telsonic somite and uropod. Adult male. J. Lateral view. K. Dorsal view of carapace. L. Antenna 1. M. Telsonic somite and uropod. Scale line = 1 mm for A, C, J-K; 0,5 mm for B, E-I, L-M; 0,25 mm for D. SOUTHERN AFRICAN CUMACEA: PART 2 235 Pereiopod 1 (Fig. 29F) short and stout, basis equal in length to remaining segments together. Distal segments stout, cylindrical. Pereiopod 2 (Fig. 29G) stout, 6-segmented, poorly armed. Dactyl very slightly longer than propodus. Pereiopods 3 (Fig. 29H) to 5 similar, well armed with long spines on merus, carpus and propodus. Propodus and dactyl very short, relatively stout. Telsonic somite (Fig. 291) little produced between uropods, about half length of peduncle. Peduncle about one and a half times length of rami, unarmed. Exopod slightly shorter than endopod, first segment unarmed; second toothed on inner edge with two stout serrated terminal spines. Both segments of endopod toothed on inner edge, first with two stout serrate setae distally, second with two terminally. Colour in life beige with numerous white chromatophores dorsally, especially on carapace, and melanophores scattered over the body and limbs ventrally, also forming a wide transverse band across second pedigerous somite. White pigment fades rapidly, but melanophores may still be evident after some months or years in alcohol. Adult male, length 3,8 mm, from Kalk Bay Harbour, False Bay. As female, except as follows: antennal notch (Fig. 29J) less evident. Eye bigger with six large lenses (Fig. 29K). Sideplates of last three pedigerous somites slightly produced laterally, abdominal sideplates produced ventrally. Aesthetasc of antenna | (Fig. 29L) extremely short. Basis of maxilliped 3 slightly longer and less curved. Basis of pereiopod | slightly longer and stouter. Peduncle of uropod (Fig. 29M) longer and more slender, bearing sixteen long slender and ten short, stout, serrate setae on inner edge. Second segment of exopod bearing eight very fine plumose setae on inner edge. First segment of endopod with thirteen short, stout serrate spines on inner edge and two larger ones distally; second segment toothed on inner edge with two large, strong serrate spines terminally. Length Adult male 3,1-4,3 mm Ovigerous female 3,1-3,4 mm Remarks Zimmer (1921) described Bodotria capensis from Cape Town on the basis of two adult males, which he ascribed to Bodotria on the basis that only four pedigerous somites were visible. He also stated that the posterior part of the carapace was bounded by a ring or collar. In his diagram this looks very like the first pedigerous somite. Apart from a figure of the whole animal, only the third maxilliped and uropod were figured. As far as can be seen from the descrip- tion and figures, the specimens in the present collection are identical with his. Hale (1953) described I. brevidactyla from male and female individuals from Langebaan Lagoon. His descriptions and figures tally with those of the 236 ANNALS OF THE SOUTH AFRICAN MUSEUM author in all respects except as follows: the adult female is shown with an acutely pointed pseudorostrum (but in some of the present specimens the siphons are calcified and appear as a forward projection of the pseudorostrum); more lenses are visible in the eye of the female; in the female also, the second segment of the uropodal exopod bear four fine setae, whereas the present ones are toothed; in the male the peduncle of the uropod has fewer setae on the inner border than in the present males. It is noticeable that individuals vary even within a single sample in all these characters, and the differences are minor. so that the present specimens are clearly members of Hale’s species. In fully ovigerous females the first pedigerous somite is quite evident dorsally and dorsolaterally but in others, including most young females and some adult males, the somite is totally invisible. Thus, within one species the main character distinguishing between Jphinoe and Bodotria becomes obscure. However, even when the first pedigerous somite is not evident, it can be seen on flexing the animal. Thus it seems appropriate to concur with Hale and place the species in Jphinoe. It is not then necessary to expand the definition of Bodotria, which would cause considerable confusion since the boundaries between the genera are not clear-cut. I. capensis is easily distinguished from the other species of Jphinoe by the very distinctive distal segments of maxilliped 3 and pereiopods 3—5 as well as by the serrations of the second segment of the uropodal endopod. Distribution Widely distributed from Liideritz to Durban at depths from 0 to 50 m, but occurring in rather small numbers. The species constitutes about 2 per cent of the individuals in the collection. Some specimens were collected by light trap in Kalk Bay Harbour and others were taken very near to Lambert’s Bay Harbour. These are both fishing harbours, indicating that the species is resistant to at least some degree of organic pollution. Cyclaspis Sars, 1865 Generic diagnosis Carapace variable in shape. Eye present or absent. Four pedigerous somites visible in male, sometimes five in ovigerous females. Second pereiopod 7-segmented. Endopod of uropod 1-segmented. Type species C. longicaudata Sars, 1865. Remarks A general discussion of the genera of the Bodotriinae is to be found on page 163 in the Introduction. The South African members of Cyc/aspis are poor SOUTHERN AFRICAN CUMACEA: PART 2 237 in number both of samples and individuals. In comparison with the warmer waters of the Indo-Pacific region with several tens of species, only three occur in South African waters, and only one other species has been recorded off the rest of Africa. It seems fairly certain that the species occurring here are quite distinct from those from other regions, but the amount of variability in the genus is so great that it is difficult to state with certainty that the species are distinct entities and not genetic variations of species from other regions. Since only three species are known from southern Africa, no key is given. The reader is referred to Hale (1944a) for.a key to all species known up to that time. Distribution of Cyclaspis The genus is a very large and widespread one, representatives being found in all oceans. Of approximately 80 species nearly 60 per cent are found exclu- sively in Australasian waters, 20 per cent in the Indo-Chinese region and 10 per cent occur around the coasts of the Americas; 3 species occur in South Africa, 1 in east Africa, 3 in European and 2 in Antarctic waters. Fully 57 per cent of the species occur between 20 and 45°S (mainly due to the very great pre- ponderance in Australasian waters) and only 5 per cent north or south of 45°. This indicates that the genus is essentially one of south temperate latitudes. The majority of species has a depth range from about 10 to 50 m, and almost all are confined to depths less than 100 m. However, a group of morpho- logically distinct species occurs at much greater depths. These are C. subgrandis Jones 1969 (3 290 m off Kenya), C. tasmanica Jones, 1969 (610 m in the Tasman Sea), C. longicaudata Sars, 1865 (120-3 834 m off Europe), C. gigas Zimmer, 1907 (193-640 m in the Antarctic) and C. spectabilis Zimmer, 1908 (140-1 300 m off South Africa). C. sibogae Calman, 1905, is also a deep-water species, occurring at a depth of 411 m in the Philippine Sea. Cyclaspis australora sp. nov. Figs 30-31 Records SST 34°S 21°E 15m _ 1 subadult J, 4 ovig. 92 (1 record) SCDF 335 25-E 7m 4subadult $3, 1 9, 3 juvs (1 record) NIWR 29°S 31°E-27°S 32°E 20-27 m 1 adult J, 1 juv. @ records) Holotype Ovigerous female, in the South African Museum, SAM-—A15488, collected during the UCT benthic survey, 21 June 1972. Type locality: 21 m, off Still Bay (34°23’S 21°26’E). UCT station number SST 66B. Description Ovigerous female, holotype, length 5,6 mm. Integument clean, white, shiny, slightly brittle and translucent with minute reticulations and scattered pits 238 ANNALS OF THE SOUTH AFRICAN MUSEUM visible at high magnifications. Carapace (Fig. 30A) oval, one and a half times as long as deep. Anterolateral angle (Fig. 30B) acute, antennal notch fairly small, rounded. Pseudorostral lobes (Fig. 30C) very short, not meeting in front of eyelobe. Eyelobe (Fig. 30D) rounded, eye consisting of eight small lenses SS Fig. 30. Cyclaspis australora sp. nov. Ovigerous female, holotype. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view of carapace. D. Detail of eyelobe. E. Antenna 1. F. Detail of distal tip of antenna 1. G. Maxilliped 3. H. Pereiopod 1. I. Pereiopod 2. J. Pereiopod 3. K. Telsonic somite and uropod. Scale line = 2 mm for A, C; 1 mm for B, D, H, K; 0,5 mm for E, G, I-J; 0,1 mm for F SOUTHERN AFRICAN CUMACEA: PART 2 239 above a red pigmented area. Middorsal region marked by a suture-line, carina hardly present. First pedigerous somite visible dorsally and dorsolaterally second narrower dorsally than ventrally. Dorsolateral edges of third to fifth pedigerous somites raised. Carapace twice length of free thoracic somites. Cephalothorax and abdomen subequal in length. Abdominal somites cylindrical with small lateral articulatory pegs on second to fifth. First segment of antenna | (Fig. 30E) slightly longer than each of next two. Flagellum 2-segmented with three setae, one rigid and two irregular and flexible. Accessory flagellum (Fig. 30F) minute, 1-segmented. Basis of maxilliped 3 (Fig. 30G) stout, curved, twice length of rest of limb. Distal prolongation triangular, reaching half-way along merus. Outer distal portion of merus expanded, reaching junction of carpus and propodus. Basis of pereiopod 1 (Fig. 30H) curved, equal in length to rest of limb. Remaining segments stout, propodus longest. Pereiopod 2 (Fig. 301) 7-segmented. Basis slightly shorter than rest of limb. Ischium short, merus and carpus subequal in length, dactyl twice as long as propodus with eleven sharp serrate spines. Pereiopods 3 (Fig. 30J) to 5 similar, stout. Merus and carpus subequal in length and as wide as basis. Telsonic somite (Fig. 30K) square in dorsal view, not produced between uropods, half length of peduncle. Peduncle half as long again as rami with seven very short spines on inner edge. First segment of exopod a quarter length of second, neither armed. Endopod very slightly shorter than exopod with six short spines distally on inner edge. Adult male, length 6,2 mm, from Natal. As female, except as follows: anterolateral angle and antennal notch rounded (Fig. 31A). Carapace narrower in dorsal view (Fig. 31B). Eye (Fig. 31C) with nine large, clear lenses. Carapace not carinate. Sideplates of abdominal somites ventrally defined posterior to insertion of pleopods only. Second segment of antenna 1 shorter and stouter, flagellum 1-segmented. Basis of maxilliped 3 less curved. Basis of pereiopod 1 (Fig. 31D) stout. pro- duced to a strong point distally; subsequent segments more slender. Basis, merus and carpus of pereiopod 2 a little shorter. Bases of pereiopods 3 to 5 shorter. Telsonic somite (Fig. 31E) slightly longer. Peduncle of uropod with twelve short and seventeen long serrate spines on inner edge as well as three dorsally on proximal half. Endopod with twenty-three short spines on inner edge. Second segment of exopod with seven very small spines on inner edge. Length Adult male 6,2 mm Ovigerous female 5,4-5,6 mm 240 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 31. Cyclaspis australora sp. nov. Adult male. A. Lateral view. B. Dorsal view of carapace. C. Detail of eyelobe. D. Pereiopod 1. E. Telsonic somite and uropod. Scale line = 2 mm for A-B, D; 1 mm for C, E. Remarks C. australora is one of the three species of Cyc/aspis occurring in southern African waters. It may be distinguished from C. spectabilis by its very much longer and more slender uropods and from C. scissa by the lack of a transverse groove across the carapace. It apparently falls within Hale’s (1944a) levis group, most closely resembling a number of unsculptured Indo-Pacific species. Within this group only C. australora and C. herdmanni Calman, 1904a have the basis of pereiopod 1 subequal in length to the rest of the limb. Comparison with the syntypes of C. herdmanni shows a number of significant differences between the two species. C. australora is about twice the size of C. herdmanni; the anterior face of the carapace is perpendicular in C. australora and normally slanted in C. herdmanni. The distal prolongation of the basis of pereiopod 1 is shorter, while the whole limb is longer in C. australora; the uropods are equal in length to the last three somites together and the peduncle is nearly one and a half times the length of the rami in C. australora, while in C. herdmanni the uropods are little longer than the last two somites together and the peduncle is subequal in length to the rami. Distribution Apparently endemic to the warmer waters of the south-eastern coast of Africa from Still Bay to northern Natal at depths from 9 to 27 m; not a common species. SOUTHERN AFRICAN CUMACEA: PART 2 241 Cyclaspis scissa sp. nov. Jae, BZ Records NIWR 30°S 30°E-29°S 31°E 15-52 m_ 1 subadult 3, 1 immature 3, 2 29 (3 records) Holotype Young female, in the South African Museum, SAM-—A15489, collected by the NIWR, 19 November 1973. Type locality: 15 m, off Durban (29°53’S 31°04’E). NIWR station number BL D1(G). Description Young female, holotype, length 5,8 mm. Integument white, velvety, with fine reticulations interspersed with scattered pits visible at high magnifications. Carapace (Fig. 32A) oval in outline with a sharply-delineated groove running transversely from middorsal region to ventral edge of carapace, ending a little behind the anterolateral angle and a small indentation dorsolaterally on either side slightly anterior to the major groove. Anterolateral angle small and acute, antennal notch small, semicircular. Pseudorostral lobes (Fig. 32B) short, ending level with eyelobe. Eyelobe (Fig. 32C) rounded, bearing three transparent lenses with reddish pigment below. Middorsal carina not strongly evident, most marked anteriorly. First pedigerous somite not visible, second narrow, third and fifth with sideplates defined dorsolaterally. Free thoracic somites less than half length of carapace, cephalothorax slightly shorter than abdomen. Abdominal somites almost cylindrical with sideplates poorly defined ventrally. Articulatory pegs present on abdominal somites 1-5. Antenna 1 (Fig. 32D) elongate, first segment geniculate, longer than second and third together. Flagellum 2-segmented bearing a single unmodified seta. Accessory flagellum minute, l-segmented. Basis of maxilliped 3 (Fig. 32E) nearly twice length of rest of limb, inner part strongly calcified where exposed ventrally. Distal elongation reaching articulation between merus and carpus. Outer edge of merus widely expanded, distal tip reaching articulation of carpus and propodus. Pereiopod | (Fig. 32F) elongate, reaching tip of pseudorostrum with carpus. Basis subequal in length to rest of limb. Merus slightly shorter than subequal propodus and dactyl. Pereiopod 2 missing. Pereiopods 3 (Fig. 32G) to 5 similar, stout. Ischium small, merus and carpus both relatively wide, strong, subequal in length. Telsonic somite (Fig. 32H) protruding between uropods for about a third its length, one and a half times as long as broad. Peduncle of uropod less than one and a half times length of telsonic somite, unarmed except for three small 242 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 32. Cyclaspis scissa sp. nov. Young female, holotype. A. Lateral view. B. Dorsal view of carapace. C. Detail of eyelobe. D. Antenna 1. E. Maxilliped 3. F. Pereiopod 1. G. Pereiopod 3. H. Telsonic somite and uropod. Young male. I. Lateral view. J. Dorsal view of carapace. K. Antenna 1. L. Detail of distal tip of maxilliped 3. M. Pereiopod 2. N. Telsonic somite and uropod. Scale line = 2 mm for A-B, I-J; 1 mm for F, H, N; 0,5 mm for C-E, G, K-M. SOUTHERN AFRICAN CUMACEA: PART 2 243 spines distally on inner edge. Endopod 1-segmented with eight small spines on inner edge and one terminally. First segment of exopod about a quarter length of second, unarmed; second with three very small spines on inner edge and one terminally. Young male, length 5,8 mm, from Natal. (Note: a single subadult male is too badly damaged for descriptive purposes.) As female, except as follows: carapace (Fig. 321) more vaulted posteriorly, transverse groove present only on dorsal half of carapace. Anterolateral angle rounded. Eyelobe (Fig. 32J) protruding between pseudorostral lobes, pigmented area larger. Antenna | (Fig. 32K) shorter and stouter, with two aesthetascs. Greater part of basis of maxilliped 3 (Fig. 32L) exposed ventrally, carpus and propodus both slightly expanded internally. Pereiopods 2 (Fig. 32M) stout, relatively short. Ischium wider than long, merus large, carpus and propodus short, sub- equal in length. Dactyl with a few terminal spines. Abdominal somite protruding less between uropods. Peduncle (Fig. 32N) shorter and stouter, unarmed. Second segment of exopod clearly serrated on inner and outer margins, first segment relatively longer. Endopod unarmed. Length Subadult male 6,3 mm Young female 5,8 mm Remarks C. scissa may be distinguished from all other southern African bodotriids by the strong transverse groove on the carapace of both sexes. It most closely resembles C. uniplicata Calman, 1904a, C. longipes Calman, 1904a and C. nubila Zimmer, 1936. In C. longipes there is no transverse groove and the basis of pereiopod 1 is considerably shorter than the rest of the limb; in C. nubila the basis of pereiopod | is considerably longer than the rest of the limb, and there is no transverse groove. C. uniplicata does possess a groove but anterior to it is a very distinctive middorsal tooth, the basis of pereiopod | is about a third the total length of the limb, which is slender. Distribution Four specimens known from the south coast of Natal at depths from 15 to 52 m. Cyclaspis spectabilis Zimmer, 1908 Fig. 33 Cyclaspis spectabilis Zimmer, 1908: 161-162, pl. 1; 1921: 124. Records SAM 34°S 18°E 460-560 m 1 subadult 3 SAM 27°S 32°E-30°S 30°E 680-1 300m 2 subadult gg, 1 dg, 11 juvs, 6 mancas (5 records). 244 ANNALS OF THE SOUTH AFRICAN MUSEUM Previous records Holotype only. Holotype Ovigerous female, deposited by Zimmer (1908) in the Berlin Zoologisches Museum. Type locality: 565 m, Agulhas Bank (35°S 18°E). Description Subadult male, length 7,8 mm, from material collected by the S.S. Pieter Faure off the Cape Peninsula. Handsome, integument appearing polished. Carapace (Fig. 33A) almost spherical, nearly as deep as long or wide. Antero- lateral angle rectangular, very slightly serrate below (Fig. 33B). Antennal notch rectangular, shallow. Eyelobe (Fig. 33C) pointed anteriorly, eyeless. Pseudo- rostral lobes very short, just meeting in front of eyelobe. Second pedigerous somite firmly fused with carapace, last three narrow. Carapace more than twice as long as free thoracic somites together, cephalothorax shorter than abdomen. Abdominal somites cylindrical, each slightly constricted anteriorly, with lateral articulatory pegs present on somites 1-4 anteriorly. Antenna | (Fig. 33D) short, basal segments subequal in length. Flagellum (Fig. 33E) 2-segmented with two long aesthetascs. Accessory flagellum minute, 1-segmented. Basis of maxilliped 3 (Fig. 33F) stout, about three times length of rest of limb; distal prolongation widely triangular, reaching more than half-way along carpus. Merus and carpus both expanded externally, distal tip of merus reaching half-way along carpus. Pereiopod 1 (Fig. 33G) reaching beyond anterior tip of carapace. Basis wide and stout, shorter than remaining segments together. Pereiopod 2 (Fig. 33H) 7-segmented. Basis equal in length to next four segments together. Lower edge of dactyl slightly serrate with three terminal spines. Pereiopods 3 (Fig. 331) to 5 similar, basis and carpus of pereiopod 3 longest. Pleopods (Fig. 33J) not setose. Telsonic somite (Fig. 33K) subequal in length to preceding somite, more than twice length of peduncle of uropods, produced between them for more than a quarter its length. Peduncle hardly twice as long as broad, unarmed. Rami subequal in length, about twice length of peduncle. Endopod |-segmented, unarmed. Exopod 2-segmented, first segment about a fifth length of second, unarmed; second serrated on inner edge but lacking spines or setae. Length Subadult male 7,8—-8,8 mm SOUTHERN AFRICAN CUMACEA: PART 2 245 J Fig. 33. Cyclaspis spectabilis Subadult male. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view of carapace. D. Antenna 1. E. Detail of distal tip of antenna 1. F. Maxilliped 3. G. Perciopod 1. H. Pereiopod 2. I. Pereiopod 3. J. Pleopod. K. Telsonic somite and uropod. Scale line = 2mm for A, C; 1 mm for B, G; 0,5 mm for D, F, H-K; 0,2 mm for E. 246 ANNALS OF THE SOUTH AFRICAN MUSEUM Remarks Zimmer (1908) described C. spectabilis from ‘several . . . females’ collected by the Deutsches Tiefsee-Expedition from 565 m off the Cape Peninsula, and a single, dried specimen from about 140 m on the Agulhas Bank. His specimens were all females while all those in the present collection are subadult males, juveniles and mancas. Thus it is difficult to state with certainty that the specimens in the author’s possession are indeed the same species as Zimmer’s, although the differences between them and his figures are consistent with sexual variations within a species. The subadult males differ from those figured by Zimmer as follows: the males have no transverse suture across the carapace; the basal segment of antenna | is shorter; the prolongations of the basis and merus of maxilliped 3 are slightly more expanded; the carpus and propodus of pereiopod 1 are slightly shorter and the uropods slightly longer. In other respects they are very similar. The combination of an almost spherical carapace and very short uropods is unique in the South African bodotriid fauna. C. spectabilis is very similar to a number of deep-water species: C. tasmanica, subgrandis, gigas and longi- caudata. It may be distinguished from the latter two species by their longer uropodal peduncles and from the former two by their more laterally compressed carapaces. Distribution Fairly common in deep waters from the Cape Peninsula to the southern Mozambique Channel at depths from 140 to 1 300 m. Bodotria Goodsir, 1843 Generic diagnosis Carapace often with lateral carinae. First pedigerous somite not visible, second moderately wide in male and very wide in female. Second pereiopod 6-segmented. Peduncle of uropod much longer than rami; endopod I- or 2-segmented. Type species Bodotria arenosa Goodsir, 1843 Remarks The genus Bodotria at present consists of about forty species, most of which are clearly very closely related to each other. The most characteristic feature of the majority is the presence of at least one pair of lateral carinae on the carapace, frequently extending along the pedigerous somites and sometimes the abdomen as well. The integument is usually strongly calcified and brittle, the animals appearing heavy and compact. Due to the fact that in most species the details of the sculpturing of the carapace are variable both between and SOUTHERN AFRICAN CUMACEA: PART 2 247 within the sexes, it is not always easy to define the limits of each species. As a result, unless large numbers of individuals are available, there is danger of variable species being split into several species, at least until the extent of variation is known. Thus the whole genus is in serious need of revision and until such time as this is done the distinctness of many species must remain in question. This probably applies equally to the new species described here. Distribution of Bodotria All forty-odd species of Bodotria so far described are from the Eastern hemisphere. 8 species are known from Indo-China or Australia and 15 from Japan (although it is likely that some of the latter will prove to be synonymous). Of the 5 species known from the Red Sea and the Mediterranean, 3 also occur in the colder waters off Norway and the British Isles; 2 of these (B. pulchella and B. scorpioides) are very widespread, being found off the west African coast as well. The western and southern coasts of Africa are very rich in species, 20 having been described so far (including those in the paper by LeLoeuff & Intes (in press)). 10 species are apparently endemic to southern Africa, 7 to west Africa, and | occurs on the geographic boundary of 20°S. All species but one are confined to depths less than 120 m, B. tenuis sp. nov. occurring between 78 and 550 m in the southern Mozambique Channel. This is also the northernmost record on the east coast of Africa. KEY TO THE AFRICAN AND EUROPEAN SPECIES OF BODOTRIA Note: the differences between many of the species of Bodotria, although constant in large numbers of individuals, are none the less so minor that a number of polymorphic species may well have been described under more than one name. Since only South African material has been available for study, no attempt has been made to group species from other areas. Each species has been keyed according to the most apparently satisfactory descriptions and figures in the literature. 1 Endopod of uropod 1-segmented in female at least (and in all males except B. /aevigata)...2 EMG OVOdOMmuUukO pode -seementeduin: bot SeXeS eee eam cea ae oe 5 2 Carapace with no trace of lateral ridges or carinae in either sex (endopod of uropod Z-Sesimemte dalmeimnalle)eeneeneinen en laevigata LeLoeuff & Intes, in press—west Africa meter acanimacwpokeschtOnepaltOmcarapaccradull Castanea eer eenieeiciere ciao cor 3 3 Lateral carina present on anterior of carapace only...... glabra Jones, 1955—west Africa SMeAenalucakiMmameacMinespOStehOmoOkd chOlmcakapaceme rare aaa ete ee ke rea 4 4 Basis of pereiopod 1 subequal in length to rest of limb; lateral carina widely produced making width of carapace subequal to length............. alata Bacescu, 1975—Red Sea — Basis of pereiopod 1 much longer than rest of limb; lateral carina not widely produced... arenosa Goodsir, 1843 —Norway, Mediterranean Se Carapace withoutlatenale Carinae user yan ss Siewa cl cles oe cde Ae eects H ehoiee eles. cro eels 6 = (CaRajoacs with IENGRIkKCAaIHIey 5 6 Kel loced 650 SOO OOO On Os oo oo Ee ee oer 7 6 Integument rugose; anterolateral angle not produced anterior to tip of pseudorostral lobes; basis of pereiopod 1 less than one and a half times length of rest of limb......... magna Zimmer, 1921—southern Africa — Integument smooth; antero-lateral angle produced anteriorly as far as tip of pseudo- rostral lobes; basis of pereiopod 1 nearly twice length of rest of limb....... nitida sp. NOV. 248 ANNALS OF THE SOUTH AFRICAN MUSEUM 7 12 13 14 115) A single lateral carina omearapace j...235. 8s 3% socked uae dae. de oe eee ee eee 8 Carapace with two lateral carinae, or lower edge of midlateral depression forming an angular ridge-as second ‘canna... . 5... f a2 atone eke © ce Oe et one) eee 17 Carina extending hardly more than half length of carapace. .).5..52- 4. eee 9 Carina extending along full length of carapace, or absent only at extreme anterior tip. ..10 Second pedigerous somite in female elevated to a point middorsally, fourth and fifth MOtelevatedsinecholic lise xe tenet ee elevata Jones, 1960—South Africa Second pedigerous somite not elevated in either sex, fourth and fifth elevated to points middorsally on both sexes, more so in female... .vertebrata semicarinata sp. et subsp nov. Endopod of uropod Jess than half lengthiof peduncles... 4... ssn eee ooo eee 11 Endopod' of uropodiat least half lengthioi pedtncles. 25.4422 ae eee 12 Third pedigerous-somite abruptly lower than second in lateral view; endopod of uropod about a third length of peduncle.......... cribaria LeLoeuft & Intes, 1972—west Africa Pedigerous somites sloping gently in lateral view, third not abruptly lower than second; endopod of uropod slightly less than half length of peduncle, =. -= >. 4.44. eae scorpioides (Montagu, 1804)—west Africa, Europe Some or all of pedigerous somites strongly elevated middorsally................-+-- 13 Pedigerous somites not elevated middorsally 42 54.4 4-5500 4.202 eee 14 Integument squamous; second and third pedigerous somites strongly elevated middorsally, forming an enormous point in female or rectangular plate in male; abdominal somites NOt; cleVatede Gece ete se ee et eee gibba (Sars, 1879)— Mediterranean Integument smooth; second and third pedigerous somites not elevated middorsally; fourth and fifth pedigerous and first abdominal somites at least elevated to points mid- dorsally xe cso ee pe A eae vertebrata vertebrata sp. et subsp. nov. Prolongation of basis of maxilliped 3 pointed distally, reaching level of carpus........ 15) Prolongation of basis of maxilliped 3 rounded distally, not reaching level of carpus..... 16 Lateral carina dorsal to midlateral line; carapace nearly twice as long as deep.......... intermedia LeLoeuft & Intes, in press—west Africa Lateral carina midlateral; carapace less than one and a half times as long as deep....... montagui Stebbing, 1912—South Africa Carpus of pereiopod 1 very stout, less than three times as long as broad, with spines on lower edse:"carapace ofsfemale) widen thamlonse a] soe ee eee ee eeeee falsinus sp. nov. Carpus of pereiopod | more than three times as long as broad, lower edge without spines; carapace of female andemalel oncenthannwide nessa iene serica Sp. NOV. Lower carina very distinct, much longer than upper, curving strongly to meet it at postero- dorsaliconenoticarapaccees = ene enenee pulchella (Sars, 1879)—Europe, west Africa Lower carina no longer than upper, forming lower edge of midlateral depression, inter- rupted half-way along if curved, or meeting anterior to posterodorsal corner........- 18 Second and fourth pedigerous somites produced to form narrow, plate-like middorsal keel; gap between distal prolongation of basis of maxilliped 3 and ischium...... clara sp. Nov. Second and fourth pedigerous somites not platelike dorsally, no gap between prolongation of basis anduschium'of maxilliped $45. 45 een oe eee neo eee 19 Rami of uropods more than-half length of peduncle..............95. 50. - eee eee 20 Rami of uropods half length of peduncleron less. ....454.26 54. eee eee an Basis of maxilliped 3 twice length of remaining segments together; major carina dorso- Feater al scsth sc 0c, pila a ae en ec sar eee ata lata Jones, 1956—west Africa Basis of maxilliped 3 nearly two and a half times length of remaining segments together; major carina apparently midlateral............... australis Stebbing, 1912 —South Africa Carinae meeting in front of posterior border of carapace to form a deep depression...... bineti LeLoeuff & Intes, in press—west Africa Carinae mot meeting wns 3 2a ae a eee ee ee eee 22 Basis of pereiopod | one and a half times length of rest of limb................++405- armoricana LeLoeuff & Intes, in press—west Africa Basis of pereiopod) (equal in lengthito test of limbs. 5. 44sec eee oe eee 23 Carapace less than twice as long as deep; propodus of pereiopod 1 longer than carpus OF dacty lewis. « 3 ane day at mee nee ey Pee ee africana Zimmer, 1921 —West Africa Carapace more than twice as long as deep; propodus of pereiopod 1 subequal in length to dactyl-shorter than canpuse.) ost hele oO ee oe tenuis Sp. NOV. SOUTHERN AFRICAN CUMACEA: PART 2 249 Bodotria clara sp. nov. Figs 34-35 Records FAL 34°S 18°E 2-4m _ 1 adult 3, 1 ovig. 2 2 records) SED 33°S 25°E (oiled, Holotype Adult male, in the South African Museum, SAM-A15481, collected during the UCT benthic survey, 5 February 1962. Type locality: 7 m, off Port Elizabeth (33°52’S 25°38’E). UCT station number SCD 383L. Description Adult male, holotype, length 2,6 mm. Integument thick, white, brittle, with many large pits. Carapace (Fig. 34A) one and a half times as long as deep, dorsal outline irregular in lateral view. Sharp dorsolateral carina running from eyelobe to posterior edge of carapace, abruptly elevated in three steps, joined rear posterior border by second carina running ventrally and then anteriorly for a short distance, continued forward as slight fold and distinct again on anterior third of carapace, reaching point of anterolateral angle. Anterolateral angle acute, bluntly rounded. Antennal notch narrow and deeply indented. Carapace in dorsal view (Fig. 34B) slightly longer than wide, pseudorostral lobes meeting for short distance in front of eyelobe. Eyelobe rounded, eye visible as two lenses laterally above some pigment. No middorsal carina on carapace. Second pedigerous somite produced to form very distinct plate-like mid- dorsal carina and a pair of lateral ridges, the latter forming strongly marked sideplates, excavated anteriorly by continuations in pits of integument. Third pedigerous somite elevated slightly, dorsally only; fourth strongly elevated middorsally forming thin median plate and produced into a pair of carinae laterally; last elevated middorsally only and somewhat excavate posteriorly. Free thoracic somites together little more than half length of carapace. Cephalo- thorax and abdomen subequal in length. Abdominal somites large, as deep as long, sideplates strongly defined ventrally; middorsal carina evident on first and second; third to fifth with lateral articulatory pegs. Antenna | (Fig. 34C) short, first segment twice length of next two together. Flagellum (Fig. 34D) 1l-segmented with one aesthetasc and three fine setae. Accessory flagellum 1-segmented with one aesthetasc. Basis of maxilliped 3 (Fig. 34E) strongly angled distal to midpoint, more than twice length of remaining segments together; distal prolongation narrow, reaching half-way along merus. Merus slightly expanded, distal tip reaching half-way along carpus. Pereiopod | (Fig. 34F) very stout, basis curved, more than twice length of rest of limb with four small spines half-way along on median edge. Ischium very 250 ANNALS OF THE SOUTH AFRICAN MUSEUM short, merus and carpus subequal in length, as are propodus and dactyl. Pereiopod 2 (Fig. 34G) fairly stout, 6-segmented. Basis shorter than rest of limb. Merus slightly shorter than carpus, propodus half length of dactyl. Pereiopods 3 (Fig. 34H) to 5 similar, basis of pereiopod 3 longest. Merus and carpus very stout, merus curved. Fig. 34. Bodotria clara sp. nov. Adult male, holotype. A. Lateral view. B. Dorsal view of carapace. C. Antenna 1. D. Detail of distal tip of antenna 1. E. Maxilliped 3. F. Pereiopod 1. G. Pereiopod 2. H. Pereiopod 3. I. Telsonic somite and uropod. Scale line = 1 mm for A; 0,5 mm for B, C, E-I; 0,2 mm for D. SOUTHERN AFRICAN CUMACEA: PART 2 Dy) Telsonic somite (Fig. 341) longer than wide, slightly produced between uropods. Peduncle of uropod nearly half as long again as telsonic somite with five long plumose setae and five serrated spines on inner edge. Exopod slightly longer than endopod, less than half length of peduncle. First segment half length of second, unarmed; second with five plumose setae on inner edge, one long plumose seta and one long slender spine terminally. First segment of endopod nearly four times length of second with five serrate setae on inner edge and one plumose seta distally on outer edge; second with three fine spines terminally. Ovigerous female, length, 1,9 mm, from False Bay. As male, except as follows: integument softer, poorly calcified, with smaller, less evident pits. Lower lateral carina of carapace (Fig. 35A) not joining upper, longer and more marked, anterior part not reaching anterolateral angle. Anterolateral angle sharper. Carapace wider ventrally than dorsally (Fig. 35B). Eye indistinct. Second pedigerous somite much wider, sideplates of fourth more distinct, fifth not elevated. Abdominal somite smaller. First segment of antenna | (Fig. 35C) shorter, both flagella (Fig. 35D) without aesthetascs. Basis of maxilliped 3 (Fig. 35E) wider, more strongly Fig. 35. Bodotria clara sp. nov. Ovigerous female. A. Lateral view. B. Dorsal view of carapace. C. Antenna 1. D. Detail of distal tip of antenna 1. E. Maxilliped 3. F. Pereiopod 3. G. Telsonic somite and uropod. Scale line = 1 mm for A-B; 0,5 mm for C, E-G; 0,2 mm for D. O52 ANNALS OF THE SOUTH AFRICAN MUSEUM angled, prolongation incised proximally. Merus slightly longer, carpus slightly expanded. Basis of pereiopod | shorter, less curved, lacking spines. Basis of pereiopod 2 shorter. Merus and carpus of pereiopods 3 (Fig. 35F) to 5 less stout. Telsonic somite (Fig. 35G) shorter. Peduncle of uropod somewhat less than twice length of exopod, unarmed. First segment of endopod with only one spine distally on inner edge, second with two terminally. Length Adult male 2,6-2,7 mm Ovigerous female 1,9 mm Remarks The species is unique in the genus in the combination of a strongly pitted integument, the dumb-bell-shaped fourth pedigerous somite and the lateral carina rising in steps towards the posterior end of the carapace. It is reminiscent of B. gibba (Sars, 1879), from which it is easily distinguished by the lack of a plate-like median carina on the fourth pedigerous somite, the much longer prolongation of the basis of maxilliped 3 and the extreme elevation of the second pedigerous somite in B. gibba. Distribution So far only three specimens known, all from False Bay or Algoa Bay (Port Elizabeth), between 2 and 7 m. Bodotria magna Zimmer, 1921 Fig. 36 Bodotria magna Zimmer, 1921: 121-123, figs 8-11. Records sub- adult adult Ovig. no. of 3 S 5 Q Q juv. total records FAL & FBY 34°S 18°E 48-87 m 15 13 1 A) 18 16 88 14 SST 34°S 21°E 50-80 m nt gy D) 17 IS) 34111 3 SCD SAS) DIE 67 m 1 1 1 Previous records ‘Great Fish Bay’ (16°S 11°E), no depth given. Holotype Ovigerous female, deposited by Zimmer (1921) in the Berlin Zoologisches Museum. Type locality: “Great Fish Bay’ (16°S 11°E). Description Ovigerous female, length 6,0 mm, from False Bay. Integument very rugose, ridges running longitudinally for the most part, most evident on cephalothorax. SOUTHERN AFRICAN CUMACEA: PART 2 255 Carapace (Fig. 36A) less than one and a half times as long as deep, dorsal outline very gently arched. Antennal notch deeply excavate, anterolateral angle acute, pointed. Eyelobe (Fig. 36B) rounded, eye wanting. Pseudorostral lobes short, not meeting in front of eyelobe. Carapace lacking dorsal and lateral carinae. Second pedigerous somite slightly elevated above level of carapace, fairly wide. Third to fifth pedigerous somites hardly elevated, with well-developed sideplates. Free thoracic and first two abdominal somites with distinct mid- dorsal carina. Carapace subequal in length to free thoracic somites together, cephalothorax and abdomen subequal in length. Abdominal somites less rugose, sideplates defined ventrally. Antenna | (Fig. 36C) short, first segment longer than next two together. Flagellum 2-segmented with two aesthetascs, accessory flagellum small, 1-segmented. Basis of maxilliped 3 (Fig. 36D) twice length of remaining segments together, strongly calcified ventrally where exposed (stippled in figure). Distal prolongation broad and of moderate length, reaching articulation of merus and carpus. Merus slightly expanded, carpus cylindrical. Pereiopod | (Fig. 36E) stout, basis slightly longer than remaining segments together, curved. Ischium short, merus and carpus stout, merus twice and carpus three times length of ischium. Pereiopod 2 (Fig. 36F) stout, 6-segmented. Basis longer than rest of limb. Merus and carpus stout, subequal in length. Propodus and dactyl] slender. Pereiopods 3 (Fig. 36G) to 5 similar, ischium, merus and carpus all stout, dactyl small. Telsonic somite (Fig. 36H) rectangular in dorsal view, not produced between uropods, little more than half length of peduncle of uropod. Peduncle unarmed, more than twice length of rami. Rami short, exopod slightly shorter than endo- pod. First segment of exopod a quarter length of second, unarmed; second with four plumose setae on inner edge and two terminally. Endopod 2-segmented, second segment less than half length of first; first with two small spines on inner edge, second with two, and one very stout terminal spine. Adult male, length 6,3 mm, from False Bay. As female, except as follows: rugosities distinct, but forming more regular honeycomb pattern. Antennal notch shallower (Fig. 361), anterolateral angle obtuse, rounded. Carapace more elongate, without carinae, but sculpturing absent dorsolaterally, resulting in the formation of a rounded edge. Second pedigerous somite narrower, not elevated; third produced to form a point middorsally. Abdominal sideplates deeper, sideplates more marked ventrally. Antenna | (Fig. 36J) bearing four aesthetascs around flagellum as well as two at tip. Basis of maxilliped 3 less angled, distal prolongation a little longer, merus less expanded and carpus more. Basis of pereiopod 1 less angled, narrower, with about ten sharp spines on inner ventral edge. Carpus not stout. Pereiopod 2 (Fig. 36K) more strongly armed, basis somewhat shorter, dactyl 254 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 36. Bodotria magna. Ovigerous female. A. Lateral view. B. Dorsal view of carapace. C. Antenna 1. D. Maxilliped 3. E. Pereipod 1. F. Pereiopod 2. G. Pereiopod 3. H. Telsonic somite and uropod. Adult male. I. Lateral view. J. Antenna 1. K. Pereiopod 2. L. Uropod. Juvenile. M. Lateral view. Scale line = 2 mm for A-B, I; 1 mm for E, H, M; 0,5 mm for C—D, F-G, J-L. SOUTHERN AFRICAN CUMACEA: PART 2 DSS shorter and stouter. Inner edge of peduncle of uropod (Fig. 36L) armed with six plumose setae proximally followed by thirty-six sharp serrate setae in two rows. Second segment of exopod with six plumose setae on inner edge and two spines terminally, one with a brush of fine filaments distally. First segment of endopod with eight and second with one blunt seta on inner edge, second terminating in one spine. Juvenile (Fig. 36M), length 3,1 mm. Integument less calcified, sculpturing restricted to thorax and ventral and posterior regions of carapace, in the form of individual raised ridges—more extensive in some than in others. Second pedigerous somite strongly elevated to a point, last two thoracic and first three abdominal somites also pointed dorsally. The second pedigerous somite is also elevated to a point in subadult males. Length Adult male 4,3-6,3 mm Ovigerous female 4,4-6,0 mm Remarks Although Zimmer’s (1921) description and figures are rather incomplete, this species is quite obviously the same as his B. magna, especially since in his description Zimmer mentions the very characteristic sculpturing of the integu- ment. The species is unique in the genus in this respect. The species nearest to B. magna are B. nitida sp. nov. and B. laevigata LeLoeuff & Intes, in press. In both of these the integument is smooth. The endopod of the uropod in males of B. /aevigata is 1-segmented and the basis of the first pereiopod in B. nitida is longer than that of B. magna. Distribution A fairly common species, constituting over 4 per cent of the individuals in the collection. It is apparently endemic to the south-western and southern coasts of southern Africa from the mouth of the Kunene River to Still Bay, at depths from 48 to 87 m. Bodotria nitida sp. nov. Figs 37-38 Records sub- adult adult ovig. no. of 3 3 3 2 2 juv. total records WCD 33°S 17°E 11-26 m 12 3 3 16 14 48 5 LBT B2-S See) 83—15am 3 6 3 7 10 y oil lS) FAL & FBY 34°S 18°E 2-11m 5 7 1 84 8 73) NB} 12 SST 34°S 21°E 15 m 1 1 1 SAM 33°S 18°E iy 2 ) 1 5 if SCD 33°S 25°E TL xon\ 5 y 4 1 3 15 1 *from stomach of Rhabdosargus globiceps 256 ANNALS OF THE SOUTH AFRICAN MUSEUM Holotype Ovigerous female, in the South African Museum, SAM-—A15483, collected during the UCT benthic survey, 24 April 1962. Type locality: 11 m, outside Saldanha Bay (33°07'S 17°58’E). UCT station number WCD 129M. Description Ovigerous female, holotype, length 6,7 mm. Large, rotund. Integument yellowish-white in alcohol, lustrous at low magnifications, reticulate with faint pits at high magnifications. Carapace (Fig. 37A) smooth, no trace of carinae; nearly twice as long as deep. Antennal notch (Fig. 37B) deeply excavate, anterolateral angle acutely pointed and protruding beyond tip of pseudorostral lobes. Eyelobe (Fig. 37C) rounded, eyeless, protruding beyond anterior tip of pseudorostral lobes. Articulatory peg present between carapace and second pedigerous somite dorsolaterally. This somite wide, smooth, very slightly elevated above level of carapace; third to fifth slightly elevated dorsally, with well-defined sideplates. Middorsal carina present on thorax. Carapace slightly longer than free thoracic somites. Cephalothorax and abdomen subequal in length. Abdominal somites with sideplates defined ventrally, each overlapping succeeding one. Antenna | (Fig. 37D) of moderate length, first segment subequal in length to next two together. Flagellum short, 2-segmented, with two aesthetascs. Accessory flagellum minute, 1-segmented. Maxilliped 3 (Fig. 37E) elongate, basis more than twice length of remaining segments together; distal prolongation short, reaching articulation of ischium and merus. Merus slightly expanded, carpus hardly at all. Basis of pereiopod | (Fig. 37F) curved, wider at base, nearly twice length of rest of limb. Ischium short, merus and carpus subequal in length, dactyl short. Pereiopod 2 (Fig. 37G) stout, 6-segmented. Basis longer than rest of limb, merus stouter than carpus, dactyl poorly armed, twice length of propodus. Pereiopods 3 (Fig. 37H) to 5 similar, merus and carpus stout, propodus and dactyl narrow. Telsonic somite (Fig. 371) rectangular in dorsal view, less than one and a half times as long as broad, very slightly produced between uropods. Peduncle of uropod one and a half times length of telsonic somite, twice length of rami, unarmed. First segment of exopod about a third length of second, unarmed; second with eight plumose setae on inner edge and two short terminal spines. Endopod 2-segmented, first segment four times length of second with three small spines on inner edge; second unarmed except for two small terminal spines. Adult male, length 6,4 mm, from False Bay. As female, except as follows: carapace (Fig. 38A) more rectangular in lateral view, anterolateral angle obtuse and rounded, antennal notch shallow. Eyelobe (Fig. 38B) shorter, carapace slightly narrower in dorsal view. Abdominal somites stouter. Antenna | (Fig. 38C) shorter, accessory flagellum surrounded by four aesthetascs. Distal prolongation of basis of maxilliped 3 (Fig. 38D) slightly SOUTHERN AFRICAN CUMACEA: PART 2 DSi more produced. Telsonic somite (Fig. 38E) protruding even less between uro- pods, a little longer than wide. Peduncle relatively longer, armed proximally with six plumose setae and distally with 25-30 serrate setae on inner edge. Second segment of exopod with six plumose setae and first of endopod with eleven fine sharp spines on inner edges. Fig. 37. Bodotria nitida sp. nov. Ovigerous female, holotype. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view of carapace. D. Antenna 1. E. Maxilliped 3. F. Pereiopod 1. G. Pereiopod 2. H. Pereiopod 3. I. Telsonic somite and uropod. Scale line = 2 mm for A, C: 1 mm for B, F, H-I; 0,5 mm for D-E, G. 258 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 38. Bodotria nitida sp. nov. Adult male. A. Lateral view. B. Dorsal view of carapace. C. Antenna 1. D. Distal tip of maxilliped 3. E. Telsonic somite and uropod. Juvenile, paratype. F. Lateral view. Scale line = 2 mm for A-B, F; 1 mm for E; 0,5 mm for C-D. Juvenile, paratype (Fig. 38F), length 4,2 mm. Differs from the adults in that the third to fifth pedigerous somites are more elevated dorsally, both in the midline and laterally. The carapace is relatively smaller and smoothly arched dorsally and the peduncle of the uropod is shorter and stouter. Length Adult male 5,8-7,2 mm Ovigerous female 4,5-7,7 mm Remarks The only other species of Bodotria totally lacking lateral carinae on the carapace are B. choprai Kurian, 1951, from India, B. magna Zimmer, 1921, from South West Africa and B. laevigata LeLoeuff & Intes (in press), from west Africa. Females of the new species may be distinguished from all of these by the very characteristic protrusion of the anterolateral angle beyond the point of the pseudorostral lobes. The male of B. choprai has the anterior part of the SOUTHERN AFRICAN CUMACEA: PART 2 259 carapace deepened and of B. laevigata has a l-segmented endopod of the uropod. Distribution Apparently endemic to the Cape, from Saldanha Bay to Port Elizabeth, at depths from 2 to 26 m. Constituting over 5 per cent of the individuals in the collection, this a fairly common species, particularly in False Bay and further north. Bodotria elevata Jones, 1960 Figs 39-40 Bodotria elevata Jones, 1960: 173-175, fig. 1. Records sub- adult adult Ovig. no. of i) 6 6 DQ @ jer, tora mecorrals LBT 33°S 17°E 20 m eal 1 3 D) FAL & FBY 34°S 18°E 9-69 m 2 1 4 1 8 7 SST 35°S 22°E 15-50 m 3 2, 1 6 4 SCD 34°S 20°E-33°S 27°E = 7-87 m 6 3 Seeslsr. slo 33 11* *type locality and some paratypes included. Previous records Off Hermanus, south-western Cape (34°S 19°E), 22-37 m, and Lambert’s Bay (32°S 18°E), 17 m (Jones 1960). Holotype Not specified. Deposited by Jones (1960) in the British Museum (Natural History). Type locality: 22 m, off Hermanus, south-western Cape (34°S 19°E). Description Ovigerous female, length 4,1 mm, from the south coast near Mossel Bay. Integument hard, brittle, chalky-white with regularly scattered deep pits. Carapace (Fig. 39A) more than one and a half times as long as deep, middorsal carina only slightly evident. Dorsolateral carina present but not strongly marked. Antennal notch (Fig. 39B) fairly deep, anterolateral angle acute. Articulatory notch present dorsolaterally on posterior border of carapace. Eyelobe (Fig. 39C) eyeless, rounded. Second pedigerous somite highly elevated, twice as deep as long. Third to fifth pedigerous somites all with sideplates well defined ventrally and dorsally. All free thoracic somites with middorsal carina continuing faintly along abdomen. Carapace slightly longer than free thoracic somites together, cephalo- thorax longer than abdomen. Abdominal somites rounded, first slightly elevated forming a posteriorly directed point. 260 ANNALS OF THE SOUTH AFRICAN MUSEUM Antenna | (Fig. 39D) short, first segment longer than next two together. Flagellum 2-segmented with two aesthetascs. Accessory flagellum minute, 1-segmented. Basis of maxilliped 3 (Fig. 39E) very long, more than two and a half times length of remaining segments together. Distal prolongation short, reaching half-way along unexpanded part of merus. Merus bluntly expanded, reaching two-thirds length of carpus. Pereiopod | (Fig. 39F) fairly long, basis very stout, broadest at midpoint. Merus and carpus subequal in length, propodus and dactyl short, slender. Pereiopod 2 (Fig. 39G) stout, 6-segmented. Basis longer than rest of limb. Merus and carpus subequal in length, propodus half length of dactyl. Dactyl armed with a few setae tipped with a brush of filaments. Fig. 39. Bodotria elevata Ovigerous female. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view of carapace. D. Antenna 1. E. Maxilliped 3. F. Pereiopod 1. G. Pereiopod 2. H. Pereiopod 3. I. Telsonic somite and uropod. Scale line = 2 mm for A-B; 1 mm for C, F; 0,5 mm for D-E, G-I. SOUTHERN AFRICAN CUMACEA: PART 2 261 Pereiopods 3 (Fig. 39H) to 5 similar; ischium short and stout; merus and carpus stout, subequal in length. Propodus and dactyl short, slender. Telsonic somite (Fig. 391) slightly longer than wide, rectangular in dorsal outline. Peduncle of uropod one and half times length of telsonic somite, two and a half times length of rami, unarmed. Exopod slightly longer than endopod, first segment about a quarter length of second, neither armed except for four terminal spines on second. First segment of endopod twice length of second, unarmed. Second armed with two slender terminal spines. Adult male, length 4,7 mm, from the south coast near Mossel Bay. As female except as follows: carapace (Fig. 40A) more rectangular in lateral outline, antennal notch and anterolateral angle obscure. Second pedigerous somite only very slightly elevated dorsally. Sideplates of abdominal somites defined ventrally. Antenna | (Fig. 40B) with three aesthetascs arising between flagella. Pro- longation of basis and merus of maxilliped 3 (Fig. 40C) slightly longer. Basis of pereiopod | narrower with two spines half-way along length, carpus narrowed proximally. Pereiopod 2 narrower and shorter. Propodus of pereiopods 3 to 5 longer. Uropod (Fig. 40D) strongly armed: peduncle with eight plumose setae proximally and twenty-eight serrate spines distally in two rows on inner edge. Second segment with five plumose setae on inner edge, one spine and two plumose setae terminally. First segment of endopod with eight spines on inner edge, second with two on inner edge and one terminally. Juvenile (Fig. 40E), length 2,4 mm, from type locality. Second pedigerous somite very strongly elevated to a sharp point dorsally, first abdominal somite elevated to two points dorsolaterally. Juveniles may be confused with juveniles Fig. 40. Bodotria elevata Adult male. A. Lateral view. B. Distal tip of antenna 1. C. Distal tip of maxilliped 3. D. Uropod. Juvenile. E. Lateral view. Scale line = 2 mm for A, E; 0,5 mm for C-D; 0,2 mm for B. Dor ANNALS OF THE SOUTH AFRICAN MUSEUM of B. magna which also have the elevated second pedigerous somite, but there is always some degree of sculpturing in the latter, whereas in B. elevata the integument is always smooth. Also, in B. magna the first abdominal somite is elevated to a single point middorsally. Length Adult male 4,7-5,0 mm Ovigerous female 3,8-4,3 mm Remarks The author has examined Jones’s paratypes which, apart from being poorly calcified, are very much the same as those in the present collection. There are a number of differences between individuals from the west coast and those (frequently more calcified) from the south coast. Those from False Bay are intermediate in most respects. In both male and female it is extremely difficult to detect the second lateral carina in Jones’s specimens and in the newer material these seem to be entirely absent; the lateral carinae do not extend on to the second pedigerous somite and there appear to be no lenses in the eye. Neither the peduncle nor the rami of the female uropod is serrated, but the armature of the male uropod is identical with that described by Jones. These variable characters are in accordance with the thesis that the species of Bodotria are more variable than those of most genera in the Bodotriidae. B. elevata is the only species in the genus in which the second pedigerous somite alone is elevated in the adult female. The male is less readily identifiable but the distal segments of maxilliped 3 are characteristic. Distribution Apparently endemic to the west and south coasts of the Cape, occurring in small numbers from Lambert’s Bay to East London at depths from 7 to 87 m. Bodotria montagui Stebbing, 1912. Fig. 41 Bodotria montagui Stebbing, 1912: 141-142, pl. 51(A). Records sub- adult adult ovig. no. of 3 db 3 g @ juv. total records SWD 26°S 15°E 26 m y) yy 3 7 1 LBT 32°S 18°E 20-33 m 2 1 1 4 4 SB 33°S 17°E 31m 1 1 1 FAL & FBY 34°S 18°E 29-87 m 1 1 y) 3 1 8 6 SST 34°S 21°F 15) inal 1 I 1 SCD 33°S 27°E 88 m 1 1 1 Previous records Off East London (32°S 28°E), 75 m. SOUTHERN AFRICAN CUMACEA: PART 2 263 Holotype Female deposited by Stebbing (1912) in the British Museum (Natural History). Type locality: 75 m, off East London. Description Ovigerous female, length 4,8 mm, from the Lambert’s Bay transect. Integu- ment solid, brownish in colour, with fine pitting and reticulation at higher magnifications. Carapace (Fig. 41A) less than one and a half times as long as deep, about one and a third times as long as wide, slightly vaulted dorsally. Single lateral carina running along entire length of carapace except at extreme anterior tip, almost straight. Anterolateral angle acute, rounded; antennal notch rounded, moderately excavate. Pseudorostral lobes meeting for short distance in front of rounded, eyeless eyelobe (Fig. 41B). Sides of carapace slightly oval in dorsal view. Middorsal carina present but not strongly defined on carapace and free pedigerous somites. Second pedigerous somite with strong lateral carina continuing from that on carapace, moderately wide, not elevated dorsally; third to fifth with sideplates rounded, keeled dorsolaterally and slightly produced middorsally. Carapace slightly longer than free thoracic somites together, cephalothorax slightly longer than abdomen. Abdominal somites rounded ventrolaterally. First segment of antenna | (Fig. 41C) wide, subequal in length to remaining segments together. Next two segments subequal in length. Flagellum 2-segmented with two aesthetascs. Accessory flagellum minute, 1-segmented. Basis of maxilliped 3 (Fig. 41D) more than twice length of remaining segments together, distal prolongation fairly long and relatively narrow, reaching beyond articulation of merus and carpus. Ischium longer than wide, longer than merus. Merus slightly expanded distally, carpus widened. Propodus and dactyl cylindrical. Basis of pereiopod | (Fig. 41E) stout proximally, longer than remaining segments together. Ischium very small, carpus longer than ischium and merus or propodus and dactyl together. Propodus slightly expanded distally. Pereiopod 2 (Fig. 41F) fairly short, basis subequal in length to rest of limb. Merus longest of distal segments, dactyl small and poorly armed. Pereiopods 3 (Fig. 41G) to 5 similar, pereiopod 5 very short and slender. Basis of pereiopod 3 subequal in length to rest of limb, merus and carpus stout, carpus slightly longer. Telsonic somite (Fig. 41H) little longer than wide, produced slightly between uropods. Peduncle of uropod one and a half times length of telsonic somite, nearly twice length of rami, unarmed. First segment of exopod less than half length of second, unarmed. Second segment serrated on inner edge with three spines terminally. First segment of endopod more than three times length of second with inner edge serrated proximally and with four spines distally. Second segment armed only with two terminal spines. Adult male, length 5,8 mm, from the Lambert’s Bay transect. As female 264 ANNALS OF THE SOUTH AFRICAN MUSEUM except as follows: integument lighter. Carapace (Fig. 411) slightly longer, second pedigerous somite narrower. Abdominal somites larger, sideplates defined ventrally. Basal segment of antenna 1 longer, accessory flagellum surrounded by five aesthetascs. Distal portion of maxilliped 3 (Fig. 41J) much more slender, particularly prolongation of basis. Basis of pereiopod 1 with seven spines near Fig. 41. Bodotria montagui Ovigerous female. A. Lateral view. B. Dorsal view of carapace. C. Antenna 1. D. Maxilliped 3. E. Pereiopod 1. F. Pereiopod 2. G. Pereiopod 3. H. Telsonic somite and uropod. Adult male. I. Lateral view. J. Distal tip of maxilliped 3. K. Uropod. Scale line = 2 mm for A-B, I; 1 mm for E-F, J-K; 0,5 mm for C-D, G-H. SOUTHERN AFRICAN CUMACEA: PART 2 265 mid-region, propodus not expanded. Merus of pereiopod 2 slightly longer. Carpus of pereiopods 3 to 5 shorter and propodus stouter. Peduncle of uropod (Fig. 41K) armed with about thirty-one fine, sharp spines on two rows on inner edge. Second segment of exopod with six plumose setae on inner edge. First segment of endopod with three short and eight long fine spines on inner edge. Length Adult male 5,8 mm Ovigerous female 3,8-5,0 mm Remarks Despite the rather indefinite nature of Stebbing’s diagrams of B. montagui, it seems fairly certain that the present specimens can be referred to his species. The main differences between the two are: Stebbing figures the propodus of maxilliped 3 of his unique female as being expanded; the propodus of pereiopod 1 is longer and not expanded; the telsonic somite is shorter; the second segment of the exopod of the uropod has a few plumose setae on the inner edge. On the other hand, the very great similarity in the rest of the uropod and maxilliped 3 suggests that these are indeed referable to the same species. The rather extra- ordinary placing of the lateral carina very far ventrally on the carapace and second pedigerous somite in Stebbing’s drawing should perhaps be discounted, since if this is an accurate representation, then the individual would indeed be unique in the genus. The species is closest to B. intermedia LeLoeuff & Intes (in press), from which it may be distinguished by the longer and more vaulted carapace, the shorter ischium and merus of maxilliped 3, the more dorsally situated lateral carina on the carapace and second pedigerous somite and the slightly longer basis of pereiopod | in B. intermedia. Distribution Apparently endemic to the coasts of South and South West Africa, being found occasionally between Liideritz and East London at depths from 15 to 88 m, more commonly in the west. Bodotria tenuis sp. nov. Fig. 42 Records sub- adult adult Oovig. no. of 3 E € © @ juv. total records SST 34°S 21°E-35°S 22°E 80-200 m Di 3 11 4 20 4 SCD 34°S 20°E-33°S 27°E 78-200 m 28 Dy, 1 ily 1 49 7 SAM 29°S 32°E 550 m 1 2D 3 1 266 ANNALS OF THE SOUTH AFRICAN MUSEUM Holotype Ovigerous female, in the South African Museum, SAM-—A15485, collected during the UCT benthic survey, 20 June 1972. Type locality: 200 m, off Still Bay (35°22’S 22°31’E). UCT station number SST 17K. Description Ovigerous female, holotype, length 5,2 mm. Integument dull white, reticu- lations not very evident even at high magnifications. Body slender, elongate. Carapace (Fig. 42A) more than twice as long as deep with strong dorsolateral carina, almost rectangular in cross-section, sides more or less vertical. Major carina running full length of carapace, second minor carina (almost absent in some) midlateral, forming lower edge of lateral depression, running from below anterolateral angle to posterior third of carapace, slightly curved. Anterolateral angle acute, rounded. Antennal notch fairly deeply excavate and small. Eyelobe (Fig. 42B) eyeless, rounded, pseudorostral lobes meeting for a very short distance in front of it. Carapace about a quarter again as long as wide. Middorsal carina present (but not strongly marked) on carapace and thorax only. A single pair of very characteristic black pigmented areas present at level of upper carina very slightly anterior to midpoint of carapace. Carapace slightly longer than free thoracic somites together, abdomen longer than cephalothorax by one somite. Second pedigerous somite wide, not elevated, dorsolateral carina strongly defined. Third to fifth pedigerous somites low, all with well-defined sideplates formed by posterior continuation of lateral carina. First two abdominal somites also with lateral carina, rest cylindrical. Antenna | (Fig. 42C) fairly small, first segment subequal in length to remaining segments together, fairly slender. Flagellum 2-segmented with two aesthetascs. Accessory flagellum minute, l-segmented. Basis of maxilliped 3 (Fig. 42D) more than twice length of remaining segments together, fairly stout; distal prolongation relatively pointed, slender, reaching articulation of merus and carpus. Ischium longer than wide. Merus slightly expanded, reaching half length of carpus. Carpus slightly expanded, propodus and dactyl cylindrical. Basis of pereiopod | (Fig. 42E) stouter proximally, slightly longer than rest of limb. Ischium very short, carpus slightly expanded, longer than ischium and merus together and shorter than subequal propodus and dactyl together. Pereiopod 2 (Fig. 42F) stout, basis slightly longer than rest of limb. Merus and carpus stout, subequal in length; propodus more than half length of dactyl. Pereiopods 3 (Fig. 42G) to 5 similar, slender. Each segment distal to basis narrower than the preceding one. Telsonic somite slightly produced between uropods. Peduncle of uropod (Fig. 42H) stout, twice length of endopod, unarmed but finely serrate on inner edge. Exopod slightly longer than endopod, first segment third length of second, unarmed; second segment armed with six slender plumose setae on inner edge and three fine spines terminally. First segment of endopod more than three SOUTHERN AFRICAN CUMACEA: PART 2 267 Fig. 42. Bodotria tenuis sp. nov. Ovigerous female, holotype. A. Lateral view. B. Dorsal view of carapace. C. Antenna 1 D. Maxilliped 3. E. Pereiopod 1. F. Pereiopod 2. G. Pereiopod 3. H. Uropod. Adult male, paratype. I. Lateral view. J. Antenna 1. K. Uropod. Scale line = 2 mm for A-B, I; 1 mm for E; 0,5 mm for C—D, F-H, J-K. times length of second, scalloped proximally on inner edge and with three small spines distally; second segment with a single terminal spine. Adult male, paratype, \ength 6,5 mm. As female except as follows: dorso- lateral carina of carapace (Fig. 421) slightly more laterally situated, ventro- lateral carina better defined, upper edge scalloped. Two pairs of pigmented areas on carapace. Anterolateral angle obtuse, antennal notch very shallow. 268 ANNALS OF THE SOUTH AFRICAN MUSEUM No lateral carinae on abdomen; sideplates defined ventrally. Basis of antenna | (Fig. 42J) stouter and shorter with two aesthetascs. Basis of maxilliped 3 narrower, distal prolongation reaching half-way along carpus, carpus slightly longer. Basis of pereiopod 1 straight. Merus, carpus and propodus of pereiopods 3 to 5 longer and narrower. Peduncle of uropod (Fig. 42K) stouter, nearly two and a half times length of endopod, armed with twenty serrate setae in two rows on inner edge. Second segment of exopod with seven plumose setae on inner edge and a single terminal spine. First segment of endopod wide, strongly serrate on inner edge, serrations almost forming a row of small setae; second segment with three very small spines on inner edge. Length Adult male 6,5-6,7 mm Ovigerous female 4,8-6,0 mm Remarks The most remarkable feature of this species is its relatively great depth range. No members of Bodotria have previously been found at depths greater than 120 m, yet the deepest at which B. tenuis is known to occur is 550 m The specimens from the deep stations in the southern Mozambique Channel differ slightly from those further south in that there are no pigmented spots on the carapace and the lower carina is very faint. The second pedigerous somite is not carinate at all. The species closest to B. tenuis may be distinguished from it as follows: B. intermedia Le Loeuff & Intes (in press) has no lower lateral carina on the carapace and the basis of pereiopod | is longer. The carapace in both B. africana Zimmer, 1921 and B. armoricana Le Loeuff & Intes (in press) is less than twice as long as deep; in B. armoricana the bases of maxilliped 3 and pereiopod | are longer and in B. africana they are shorter. Distribution From the Cape Peninsula to the southern Mozambique Channel at depths from 78 to 550 m; one of the more common species at these depths. Bodotria falsinus sp. nov. Fig. 43 Records sub- adult adult ovig. no. of 3S 3 Q other total records FAL & FBY 34°S18°E 40-69 2 1 2) D 7 5 SST 35°S 22°E 50-80 4 13 10 i 34 4 SCD 33°S 25°R 32 1 1 1 SOUTHERN AFRICAN CUMACEA: PART 2 269 Holotype Ovigerous female, in the South African Museum, SAM-—A15482, collected during the UCT benthic survey, 20 June 1972. Type locality: 80 m, off Still Bay (34°40’S 21°39’E). UCT station number SST 29X. Description Ovigerous female, holotype, length 3,1 mm. Integument clear white, slightly reticulate, with a number of small pits on carapace. Carapace (Fig. 43A) very flat, almost twice as long as deep, slightly wider than long. Middorsal carina poorly defined; lateral carina dorsal to midlateral line, very evident, making carapace wider dorsally than ventrally in cross-section. A second much more evanescent carina present below major one, formed by lower edges of a series of small pits in a row, evident anteriorly only, beginning immediately behind eyelobe. Anterolateral angle (Fig. 43B) acute, antennal notch deep and narrow. Pseudorostral lobes short, not meeting in front of narrow, eyeless eyelobe (Fig. 43C). Second pedigerous somite almost as wide as long, with midlateral carina continuing from carapace; ventrolaterally with rounded, yellowish protuberance. Third and fourth pedigerous somites with sideplates defined, not elevated dorsally; fifth slightly elevated in middorsal line. Middorsal carina present on thorax, absent from abdomen. Carapace longer than free thoracic somites together, abdomen equal in length to carapace and first two free thoracic somities together. Antenna | (Fig. 43D) small, first segment rectangular in outline, nearly twice length of remaining segments together. Flagellum 1-segmented with two aesthetascs. Accessory flagellum minute, 1-segmented. Maxilliped 3 (Fig. 43E) stout, basis curved, about twice length of rest of limb. Distal prolongation short, rounded, reaching half-way along merus. Ischium longer than wide, merus slightly expanded, carpus very wide distally. Basis of pereiopod 1 (Fig. 43F) slightly longer than rest of limb, curved. Ischium short, wider than long; merus stout, carpus very stout, hardly more than twice as long as broad, with eight spines on lower edge, longer than subequal propodus and dactyl together. Pereiopod 2 (Fig. 43G) stout, basis more than one and a half times length of remaining segments together. Suture-line of basis and ischium faintly visible on one side, marked by a single plumose seta. Merus longer and stouter than carpus; dactyl and propodus both short and stout, dactyl about twice length of propodus. Pereiopods 3 (Fig. 43H) to 5 similar, carpus of each longest of distal seg- ments. Pereiopod 5 by far the shortest. Telsonic somite (Fig. 431) longer than wide, produced between uropods for a short distance. Peduncle of uropod little longer than telsonic somite, one and a half times length of endopod, with six small, fine spines each on inner and outer edges. Exopod slightly shorter than endopod, first segment a third 270 ANNALS OF THE SOUTH AFRICAN MUSEUM length of second, unarmed; second armed with six plumose setae on inner edge and two spines terminally. First segment of endopod four times length of second, armed only with two small spines distally on outer edge; second armed with two spines terminally. Fig. 43. Bodotria falsinus sp. nov. Ovigerous female, holotype. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view of carapace. D. Antenna 1. E. Maxilliped 3. F. Pereiopod 1. G. Pereiopod 2. H. Pereiopod 3. I. Telsonic somite and uropod. Adult male, paratype (large). J. Lateral view. K. Dorsal view of carapace. L. Antenna 1. M. Uropod. Adult male, paratype (small). N. Lateral view of carapace. Scale line = 2 mm for A, C, J-K; 1 mm for I, M—N; 0,5 mm for B, D-H, L. SOUTHERN AFRICAN CUMACEA: PART 2 271 Note: although the appendages of all female specimens referred to this species are very similar, the gross appearance of animals, even from the same sample, varies quite considerably. In ovigerous females the sides of the carapace may be pitted or quite smooth; the lower carina may be fairly evident, evanescent (Fig. 43A) or absent; the sharpness of the lateral carina is variable; the second pedigerous somite may or may not be carinate laterally and may or may not possess a rounded protuberance. The size of individuals varies between 2,4 and 4,1 mm. These variable characters are not always found together in the same individuals, suggesting that they are genetic variations. Adult male, paratype, length 4,2 mm. As female, except as follows: carapace (Fig. 43J) almost rectangular in lateral outline, fewer pits present. Antennal notch not as deep. Eyelobe (Fig. 43K) wider, carapace longer than wide. Abdominal somites larger, sideplates defined ventrally. First segment of antenna | (Fig. 43L) larger, accessory flagellum surrounded by six aesthetascs. Basis of maxilliped 3 less curved; ischium and prolongation of basis longer. Median edge of basis of pereiopod | with four spines. Merus and carpus of pereiopods 3 to 5 stouter. Telsonic somite slightly more produced between uropods. Peduncle of uropod (Fig. 43M) with seven plumose setae and ten small sharp spines on inner edge. Second segment of exopod terminating in four spines. First segment of endopod serrated, serrations alternating with eight small spines; second segment terminating in three spines. Note: as in the female, there are a number of variable characters in the males. The secondary carina of the carapace may be well (Fig. 43N) or poorly (Fig. 43J) defined; the lateral carina may be present or absent from the second pedigerous somite; the integument may be strongly or slightly pitted. The size varies considerably. Length Adult male 2,9-4,6 mm Ovigerous female 2,4-4,1 mm Remarks B. falsinus is similar to B. vertebrata sp. nov., B. lata Jones, 1955 and B. australis Stebbing, 1912. It may be distinguished from B. vertebrata by the absence of elevated points on the pedigerous and abdominal somites and by its much wider carapace, and from B. /ata and B. australis also by its wider carapace, deeper antennal notch and enlarged carpus of pereiopod 1. Distribution Apparently endemic to the south coast of South Africa from False Bay to Port Elizabeth at depths from 32 to 80 m; not a commonly encountered species. Diz ANNALS OF THE SOUTH AFRICAN MUSEUM Bodotria vertebrata vertebrata sp. et subsp. nov. Fig. 44 Records sub- adult adult ovig. no. of 3 3 2 $&¢ total records FAL & FBY 34°S 18°E 11-31 m 13 1 11 6 31 15 SST 34°S 21°E 15m 1 3 4 1 SCD 34°S 23°E 11-42m 2 1 yD 2 7 3 Holotype Ovigerous female, in the South African Museum, SAM-A15487, collected during the UCT benthic survey, 13 July 1967. Type locality: 23 m, False Bay (34°08’S 18°30’E). UCT station number FBY 90V. Description Ovigerous female, holotype, length 3,1 mm. Integument white, crystalline; no reticulations visible even at high magnifications. Carapace (Fig. 44A) smooth, less than twice as long as deep. Single distinct lateral carina running from posterior edge to level of anterolateral angle, with a few small, rounded pits in a single row beneath. Carapace rounded in cross-section, no middorsal carina, lateral carina not very sharp. Anterolateral angle acute, rounded; antennal notch narrow, moderately excavate. Pseudorostral lobes meeting for a short distance in front of rounded, eyeless eyelobe (Fig. 44B). Carapace less than one and a half times as long as wide. Second pedigerous somite very wide, lacking lateral carina; third produced dorsally and laterally to form three rounded spines; fourth and fifth somites elevated to points middorsally only, sideplates defined dorsolaterally. First two abdominal somites produced to a wider elevated band dorsally. Carapace and free thoracic somites subequal in length, cephalothorax longer than abdomen by two segments. Antenna | (Fig. 44C) very small, basis longer than remaining segments together. Flagellum with two aesthetascs, accessory flagellum minute. Basis of maxilliped 3 (Fig. 44D) strongly curved, less than one and a half times length of remaining segments together; distal prolongation short, rounded, reaching proximal third of merus. Ischium relatively long and narrow, merus wide but very slightly expanded distally; carpus wider than long. Basis of pereiopod 1 (Fig. 44E) very stout, slightly longer than remaining segments together, slightly curved. Ischium very short, much wider than long. Merus stout, carpus very stout, longer than subequal propodus and dactyl together. Pereiopod 2 short and stout (Fig. 44F). Basis longer than rest of limb; junction of ischium and basis evident, marked by a single plumose seta. Merus SOUTHERN AFRICAN CUMACEA: PART 2 273 Fig. 44. Bodotria vertebrata vertebrata sp. et subsp. nov. Ovigerous female, holotype. A. Lateral view. B. Dorsal view of carapace. C. Antenna 1. D. Maxilliped 3. E. Pereiopod 1. F. Pereiopod 2. G. Pereiopod 3. H. Telsonic somite and uropod. Adult male, paratype. I. Lateral view. J. Distal tip of maxilliped 3. K. Uropod. Scale line = 2 mm for I; 1 mm for A-B, D-G; 0,5 mm for C, H, J-K. 274 ANNALS OF THE SOUTH AFRICAN MUSEUM and carpus wide and stout, propodus short, more than half length of narrow dactyl. Pereiopods 3 (Fig. 44G) to 5 similar, slender; pereiopod 5 much shorter than 4. Telsonic somite (Fig. 44H) more than one and a half times as long as wide, rounded posteriorly, slightly produced between uropods. Peduncle of uropod one and a third times length of telsonic somite, more than one and a half times length of rami, unarmed, but serrated on inner edge. First segment of endopod two and a half times length of second, armed only with two spines distally; second slightly serrated on inner edge with a single spine terminally. First segment of exopod more than three times length of second; second with five plumose setae on inner edge and two terminally. Adult male, paratype, length 3,5 mm. As female, except as follows: carapace (Fig. 441) more rectangular in lateral outline, carina situated less dorsally, a few more pits below. Antennal notch shallower. Pedigerous somites narrower, less elevated dorsally, abdominal somites not at all. Abdominal somites bigger, sideplates defined ventrally. Antenna 1 shorter, accessory flagellum surrounded by five aesthetascs. Ischium, merus and carpus of maxilliped 3 (Fig. 44J) longer. carpus wider. Basis of pereiopod | longer. Pereiopods 3 to 5 shorter. Peduncle of uropod (Fig. 44K) slightly more than twice length of rami, armed with nine long serrate spines proximally and six very short ones distally on inner edge. First segment of endopod with seven serrate setae on inner edge, second with two short and three very short spines. Length Adult male 3,1-3,8 mm Ovigerous female 2,9-4,1 mm Remarks The presence of elevated spines middorsally on thoracic and abdominal somites 1s unique in the genus. There are two distinct forms of the species: those from the south coast and False Bay have the lateral carina continuous along almost the whole length of the carapace and a number of small rounded pits below: those from the west coast have the carina extending only along half of the carapace without pits below. The forms are so similar that it seems unnecessary to split them into two species, but the differences are consistent enough to require subspecific differentiation. The differences between B. vertebrata vertebrata and B. verte- brata semicarinata are described in the discussion of the latter below. Distribution Found occasionally from False Bay to Knysna at depths from 11 to 42 m; less common than B. y. semicarinata. SOUTHERN AFRICAN CUMACEA: PART 2 DAS Bodotria vertebrata semicarinata sp. et subsp. nov. Fig. 45 Records sub- adult adult ovig. no. of 3 5 Q Q juv. total records WED) 3327S 17-E-34°S 18°B 11-32m 8 6 16 5 ill 36 fi WBE 32-S 18 E 20-33 m 1 1 D, y) SB 33S IAL 7-29 m 6 10 13 11 40 11 SAM 33°S 18°E ? D 3 5 (| *from stomach of Rhabdosargus globiceps. Holotype Ovigerous female, in the South African Museum, SAM-A15486, collected during the UCT benthic survey, 25 April 1962. Type locality: 26 m, off Saldanha Bay (33°07'S 17°57’'E). UCT station number WCD 134X. Description This subspecies is identical in most respects with B. v. vertebrata, differing from it as follows: Ovigerous female, holotype, length 3,4 mm. Integument browner, velvety, finely reticulate. Carapace (Fig 45A) deeper, about one and a half times as long as deep, hardly longer than wide. Lateral carina extending no more than half length of carapace, lacking pits below. Pseudorostral lobes deeper and shorter, not meeting in front of eyelobe. Second pedigerous somite deeper, with mid- dorsal carina; third less elevated dorsally. Abdomen shorter, subequal in length to cephalothorax. Basis of antenna | slightly wider. Bases of maxilliped 3 and pereiopod | longer, of pereiopod | (Fig. 45B) nearly twice length of rest of limb. Pereiopod 2 (Fig. 45C) longer and more slender. Peduncle of uropod (Fig. 45D) twice length of rami, not serrate. Adult male, paratype, length 3,8 mm. The same differences occur in the external anatomy of the males (Fig. 45E), apart from which the peduncle of the uropod (Fig. 45F) is about one and three-quarters times the length of the rami with sixteen long, slender spines evenly spaced along inner edge. The first segment of the exopod is slightly longer relative to the second. Length Adult male 3,6-4,1 mm Ovigerous female 2,9-4,3 mm 276 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 45. Bodotria vertebrata semicarinata sp. et subsp. nov. Ovigerous female, holotype. A. Lateral view. B. Pereiopod 1. C. Pereiopod 2. D. Telsonic somite and uropod. Adult male, paratype. E. Lateral view. F. Uropod. Scale line = 2 mm for E; 1 mm for A; 0,5 mm for B-D, F. Remarks In the genus Bodotria in most cases it is difficult to determine the limits of species because there are such great individual differences that there is a danger of creating more species than is perhaps realistic. For this reason the number of species proliferates greatly in almost every area where members of the genus are to be found. B. vertebrata is one of the few species in which the differences between individuals are consistent in two contiguous geographical areas, and for this reason subspecific separation is possible. The two forms are clearly very similar to each other and do not warrant specific differentiation. Distribution Apparently endemic to the west coast of South Africa from Lambert’s Bay to Table Bay at depths from 11 to 33 m. Somewhat’more commonly found than B. y. vertebrata, this subspecies accounts for nearly 2 per cent of the individuals in the collection. SOUTHERN AFRICAN CUMACEA: PART 2 Hig Bodotria serica sp. nov. Fig. 46 Records sub 3} adult adult ovig. & no. of 3 re) 2 @ juv. total records WCD 34°S 17°E-33°S 18°E 65-79 m 1 1 D, 2D FAL & FBY 34°S 18°E 17-87 m 8 Ae Gn ie 7 47 24 SST 35°S 22°E 80 m 15 1B w22 3 6 59 yy Holotype Ovigerous female, in the South African Museum, SAM-—A15484, collected during the UCT benthic survey, 21 June 1972. Type locality: 80 m, off Still Bay (34°40’S 21°39’E). UCT station number SST 29W. Description Ovigerous female, holotype, length 3,7 mm. Integument silky white, trans- lucent, not strongly calcified, without reticulations. Carapace (Fig. 46A) nearly twice as long as deep, about one and a quarter times as long as wide. Dorsal carina faintly visible on posterior half of carapace and thoracic somites, almost invisible on abdomen. Single pair of rounded lateral carinae present about a third distance from dorsal edge of carapace with slight longitudinal depression below (not present in all specimens). Anterolateral angle acutely pointed, antennal notch evident, fairly deep, rounded. Pseudorostral lobes short, not meeting in front of eyeless eyelobe (Fig. 46B). Second pedigerous somite very wide, carinate dorsally but not laterally; third deep, moderately wide, not carinate laterally; fourth and fifth very slightly elevated dorsally with sideplates defined dorsally and ventrally. Carapace slightly longer than free thoracic somites together, cephalothorax and abdomen subequal in length. Abdominal somites cylindrical. Antenna | (Fig. 46C) small, first segment subequal in length to remaining segments together. Flagellum 1l-segmented with two aesthetascs; accessory flagellum 1-segmented. Maxilliped 3 (Fig. 46D) fairly long, basis more than twice length of rest of limb. Distal prolongation rounded, hardly reaching articulation of merus and carpus. Merus widely expanded, carpus wider distally than proximally. Basis of pereiopod | (Fig. 46E) slightly longer than rest of limb; carpus longest of remaining segments, propodus and dactyl subequal in length. Pereiopod 2 (Fig. 46F) stout, basis about one and a half times length of rest of limb; merus and carpus subequal in length, propodus more than half length of dactyl. Pereiopods 3 (Fig. 46G) to 5 long, bases longer than rest of limbs. Telsonic somite (Fig. 46H) one and a third times as long as wide, hardly produced between uropods, two-thirds length of peduncle of uropod. Peduncle 278 ANNALS OF THE SOUTH AFRICAN MUSEUM unarmed, very slightly less than twice length of rami. First segment of exopod half length of second, second with seven plumose setae on inner edge and a single spine terminally. First segment of endopod more than twice length of second, serrate proximally and with seven sharp spines distally on inner edge; Fig. 46. Bodotria serica sp. nov. Ovigerous female, holotype. A. Lateral view. B. Dorsal view of carapace. C. Antenna 1. D. Maxilliped 3. E. Pereiopod 1. F. Pereiopod 2. G. Pereiopod 3. H. Telsonic somite and uropod. Adult male, paratype. I. Lateral view. J. Distal tip of maxilliped 3. K. Telsonic somite and uropod. Scale line = 2 mm for B,I; 1 mm for A, E, H, K; 0,5 mm for C_D, F-G, J. SOUTHERN AFRICAN CUMACEA: PART 2 279 second with three terminal spines and serrate on inner edge. Adult male, paratype, length 3,4 mm. As female, except as follows: carapace (Fig. 461) with several small depressions below lateral carina; anterolateral angle obtuse, antennal notch shallower. Carapace narrower dorsally, with no middorsal carina. Sideplates of abdominal somites defined ventrally. Fifth pedigerous somite produced ventrally to form a rounded protuberance. Four small aesthetascs between flagella of antenna 1. Merus of maxilliped 3 (Fig. 46J) smaller and carpus larger. Pereiopod 2 smaller than in female. Telsonic somite (Fig. 46K) produced less between uropods. Peduncle of uropod narrower with eighteen setae in two rows on inner edge. Second segment of exopod with three plumose setae, first of endopod with eleven small spines on inner edge. Length Adult male 3,3-4,8 mm Ovigerous female 3,1-4,6 mm Remarks This species is most closely allied with B. tenuis sp. nov. and B. australis Stebbing, 1912, differing from B. australis mainly in the lack of a lateral carina on the second and third pedigerous somite in the female or second and fourth in the male, and the longer peduncle of the uropod; and from B. tenuis in the lack of the carinae, the rounded prolongation of the basis of maxilliped 3 and the absence of a second lower ridge below the main carina. In recorded depth range as well as in morphological detail, this species seems to be intermediate between the two named above. Distribution Apparently endemic to the south-western coast of South Africa from Saldanha Bay to Still Bay, at depths from 17 to 87 m, this is a fairly common species, especially in the Still Bay region. It accounts for about 2 per cent of the individuals in the collection. Bodotria australis Stebbing, 1912 Bodotria australis Stebbing, 1912: 142, pl. 51(B). Remarks Stebbing described this species from a single female 3,25 mm in length. Although corresponding in many features with several of the new species described here, Stebbing’s figures show several characters which, if accurately portrayed, are sufficiently distinctive to separate this species from all the others known from South Africa. In particular the lateral carina is present on pedige- rous somites 2 to 4 and abdominal somites 2 to 5. A lower lateral carina is present on the carapace, which is more than one and a half times as long as 280 ANNALS OF THE SOUTH AFRICAN MUSEUM broad. The distal prolongation of maxilliped 3 is rounded and the carpus of perelopod | is not particularly stout. This combination of characters separates Stebbing’s individual from B. montagui Stebbing, 1912, B. falsinus sp. nov., B. vertebrata sp. nov., and B. serica sp. nov. B. australis is most similar to B. tenuis sp. nov., which differs in the absence of a lateral carina on abdominal somites 3 to 5 and a dorsally situated lateral carina which makes the carapace flat on top. Distribution Off East London (32°S 28°E) at a depth of 75 m. Bodotria glabra Jones, 1955 Bodotria glabra Jones, 1955: 282-284, figs 1-2. Remarks This species is known only from female and juvenile individuals in two plankton samples collected aboard the R.R.S. William Scoresby on the border of the southern African region. Distinguishing characters are: an elongate, flattish carapace (nearly two and a half times as long as deep, about one and a half times as long as wide) with a very faint lateral carina anteriorly; the basis of maxilliped 3 is stout, and the wide distal prolongation reaches the level of the carpus; the basis of pereiopod 1 is longer than the rest of the limb. The length of the ovigerous female is 4,5 mm, It is the only species occurring in Africa south of 20°S in which the endopod of the uropod is 1-segmented. Distribution Off Cape Frio (19-20°S 12°E), plankton, 0-100 m. Incertae sedis Iphinoe ? zimmeri (Stebbing, 1910) Figs. 47-48 Iphinoe zimmeri Stebbing, 1910: 411-412, pl. 44. Records sub- adult adult Ovig. no. of 3 6 @ juv. total records FAL & FBY 34°S18°E 17-44 m 3 D 5 4 Soul 34°S 21°F 15-80 m 2 bt yD, We) 3. AS 40 7 SCD 34°S 21°E-33°S 25°E 42-73 m 4 30) 2 eel Si ee Dei 5 Previous records East London (32°S 28°E), 75 m (Stebbing 1910). Holotype Adult male, deposited by Stebbing (1912) in the British Museum (Natural History). Type locality: 75 m, off East London (32°S 28°E). SOUTHERN AFRICAN CUMACEA: PART 2 281 Description Ovigerous female, length 7,0 mm, from the south coast, near Port Elizabeth. Integument poorly calcified, translucent, with small shallow pits. Carapace (Fig. 47A) elongate, more than twice as long as deep, pointed anteriorly. Antennal notch (Fig. 47B) moderately deep, triangular. Anterolateral angle acute, rounded. Carapace in dorsal view (Fig. 47C) narrower anteriorly, pseudorostral lobes meeting for a short distance in front of eyelobe. Eyelobe small, pigmented, Fig. 47. Iphinoe ? zimmeri. Ovigerous female. A. Lateral view. B. Detail of anterior tip of carapace. C. Dorsal view of carapace. D. Antenna 1. E. Maxilliped 3. F. Distal tip of maxilliped 3. G. Pereiopod 1. H. Pereiopod 2. I. Pereiopod 3. J. Telsonic somite and uropod. Scale line = 2 mm for A, C; 1 mm for B, E, G, J; 0,5 mm for D, F, H-I. 282 ANNALS OF THE SOUTH AFRICAN MUSEUM with very small scattered lenses. No middorsal carina—midline indicated by double rows of minute tubercles. First pedigerous somite invisible, second very wide. Carapace subequal in length to free thoracic somites together. Gonad clearly visible as triangular orange patch laterally beneath integument of second pedigerous somite (dotted in Fig. 47A). Cephalothorax subequal in length to first five abdominal somites together. Abdomen cylindrical, each somite slightly produced posterolaterally. Antenna | (Fig. 47D) short, first segment longer than next two together. Flagellum 2-segmented with two short aesthetascs. Accessory flagellum very small, 1-segmented. Basis of maxilliped 3 (Fig. 47E) more than twice as long as remaining segments together, abruptly angled proximal to its mid-point. Distal prolonga- tion (Fig. 47F) rather broad and short, reaching articulation of merus and carpus distally. Carpus slightly expanded. Pereiopod | (Fig. 47G) slender, elongate, reaching beyond tip of pseudo- rostrum with carpus. Basis longer than remaining segments together. Ischium very short, merus, carpus and dactyl subequal in length, propodus slightly longer. Pereiopod 2 (Fig. 47H) stout, 6-segmented. Basis equal in length to next three segments together. Carpus slightly longer than merus, well armed distally. Dactyl slender. Pereiopods 3 (Fig. 471) to 5 similar. Ischium, carpus and merus stout, propodus and dactyl very slender. All segments of pereiopods 2 to 5 distal to basis with very characteristic long, stout setae tipped with a brush of filaments. Telsonic somite (Fig. 47J) one and a half times as long as wide, very slightly produced between uropods, subequal in length to peduncle. Peduncle subequal in length to rami with two very small spines on inner edge. First segment of exopod less than half length of second, unarmed. Second with 10 plumose setae on inner edge and three terminal spines wih apical bristles. First segment of endopod two-thirds length of second with seven spines on inner edge and two terminally with apical bristles. Adult male, length 7,6 mm, from the south coast, near Port Elizabeth. As female, except as follows: integument very slightly wrinkled. Carapace (Fig. 48A) twice as long as deep, pseudorostral lobes (Fig. 48B) less pointed. Eye much larger, strongly pigmented, with six large lenses arranged in a ring. Antennal notch and anterolateral angle rounder and shallower. Carapace slightly depressed behind antennal notch and above posterior extremity of eyelobe. Gonad whitish, slightly visible through integument of second pedi- gerous somite. Sideplate of fourth pedigerous somite produced forward as linguiform process. Sideplates of abdominal somites defined ventrally. Ventral sternite (Fig. 48C) of third pedigerous somite produced into a crescentic ridge, of fourth with a rounded tubercle and of fifth with a larger, backward-pointing projection. Antenna | (Fig. 48D) stouter, with fifteen aesthetascs surrounding flagellum. SOUTHERN AFRICAN CUMACEA: PART 2 283 Basis of perelopod | longer. Pereiopods 3 (Fig. 48E) to 5 more slender, last four segments longer. Peduncle of uropod (Fig. 48F) armed with seven slender spines followed distally by about twenty-five serrated spines in two rows. Second segment of exopod with seventeen plumose setae on inner edge; first segment of endopod with ten serrate setae and second with twenty fine spines on inner edge. Length Adult male 7,9-8,4 mm Ovigerous female 5,8—-8,4 mm Remarks These animals are clearly the same as the one described by Stebbing (1910) as Iphinoe zimmeri. In almost all respects this species is typical of Jphinoe, particularly in the general appearance of the body, the anterior extension of the sideplate of the fourth pedigerous somite and the sternal processes of the males. However the fact that the first pedigerous somite is never visible in either sex excludes it from Iphinoe as the genus is now defined. The species does fit the Fig. 48. Iphinoe ? zimmeri Adult male. A. Lateral view. B. Detail of anterior tip of carapace. C. Ventral sternites of pedigerous somites. D. Antenna |. E. Pereiopod 3. F. Uropod. Scale line = 2 mm for A; 1 mm for B-C, E; 0,5 mm for D, F. 284 ANNALS OF THE SOUTH AFRICAN MUSEUM generic diagnosis of Bodotria, but is patently unlike any other members of the genus. Since the state of the first pedigerous somite is the only character which reliably distinguishes between Jphinoe and Bodotria, altering the generic diagnosis of Iphinoe to accommodate this species would be untenable. So for the present at least the generic position of the species must remain uncertain. Distribution A fairly common species found between False Bay and East London at depths from 15 to 80 m. DISTRIBUTION OF THE BODOTRIINAE The rate of endemism is very high within the Bodotriinae as well as in the family as a whole. Of approximately 183 species only 25 (14%) occur across more than one major climatic or zoogeographic zone and 9 of these in more than one ocean: 4 of the 9 are found in both the Indian and Pacific Oceans, linking the Australasian, Indo-Chinese and Japanese fauna into an Indo-West- Pacific group; the other 5 demonstrate the strong link between the Mediter- ranean, eastern Atlantic and Indo-West-Pacific faunas—2 occur in the Mediter- ranean, west Africa and Indo-China, | in the Mediterranean, the Red Sea and Ceylon and 1 in west Africa and India, while the most widely distributed species of all, Iphinoe crassipes, occurs in the Red Sea, west Africa, South Africa and India. The distribution of the species in the subfamily is detailed in Table 3. Each species may have more than one entry if it is known to occur in widely separate regions. It is clear that, as with the Vaunthompsoniinae (Day 1975), the Bodotriinae are negatively amphipolar (Zimmer 1941). Not a single species is yet known from latitudes greater than 70° and only 4 per cent of the records are from latitudes TABLE 3 Distribution of Bodotriinae according to depth and latitude (data mainly from Jones 1969) shore—5S m 5-200 m 200-2000m >+>2000m Total no. Yes no. Ye, no. A no. A no. YA N of 70°N pa me os Le — ass ae = oe ees 70°N-50°N — a AL <<} 1 <1 1, 200 7 — WnNwm BW fs It can be seen that the maximum depth at which each group occurs is very approximately double that of the previous group. The water pressure would also double at depths of 10, 20, 40, 80 m, etc., and these depths correspond fairly well with major changes in the fauna. It is not possible to say how well this relationship would hold at greater depths, due to the very small numbers of samples and individuals from these areas. Despite the correlation between pressure and faunal changes it is likely that temperature—or a temperature-related parameter—is of more significance in controlling depth distribution. This statement is borne out by the fact that many species occur at greater depths the further east they are found, because the temperature contours are deeper on the warmer east coast than on the cooler west coast. A similar effect is shown by Millard (1978) for the southern African Hydrozoa. A further factor which must be important in controlling the distribution of bodotriids is the size and composition of the substrate, since these animals are burrowing detritivores. Due to their very small size they presumably require fairly fine, well-sorted sediments. Particle size determinations for many of the UCT and NIWR samples may later become available for analysis, at which time this problem may be solved at least partially. The material from the collections at hand has added to our knowledge not only of the local cumacean fauna but also of the depth distribution of several genera. Cyclaspis scissa sp. nov. and C. australora sp. nov. are the first shallow- water species in the genus recorded from Africa. Bodotria and Eocuma have always been considered to be distinctly shallow-water genera, but B. tenuis sp. nov. has increased the known depth range for its genus from 120 m to 550 m, and Eocuma aculeatum sp. nov. has increased that of Eocuma from 108 to 550 m. Cyclaspoides pellucidus sp. nov. is the second species known in the genus and the first from the Southern hemisphere. Mossambicuma and Austrocuma are new genera from previously unsampled areas (a tropical estuary and a Cape 288 ANNALS OF THE SOUTH AFRICAN MUSEUM shore respectively). Alticuma establishes a new genus consisting of two deep- water species. The rate of endemism appears to be extremely high. In the southern African Vaunthompsoniinae it is of the order of 70 per cent (7 endemics out of 11 species), but in the Bodotriinae 28 (82 %) of the 34 species have been found south of 20°S only. The ranges of another four extend beyond this limit, but none of these is found in any other areas. One record (U. senegalensis) is a doubtful identification and only one species (/. crassipes) occurs from equatorial Africa to India. It should be stressed, however, that very little data is available from the regions due north of the area under consideration so that these figures must be treated with caution. Finally, 4586 specimens of 31 species in 607 records were examined in this study. This gives a figure of 7,5 individuals per record and a specimen : species ratio of 148: 1. In comparison, the Vaunthompsoniinae gave figures of 77 specimens, 11 species and 42 records with 1,8 individuals per record and a specimen : species ratio of 7:1. Thus the Bodotriinae exhibit much lower diversity and a much higher rate of occurrence than do the Vaunthompsoniinae in the same area. The ecological reasons for these differences are not clear, but the immediate cause is the very large number of specimens of some very success- ful species, particularly Iphinoe africana with 1 603 individuals and I. stebbingi with 1 186. These two species account for over half the number of individuals. The other numerically successful species are I. dayi (152 individuals), I. crassipes (143), Bodotria magna (200), B. nitida (238) and B. serica (108). Thus 7 species (about 20% of the total) account for 2 630 (nearly 60%) of individuals. In fact Iphinoe and Bodotria together account for 21 (about 65%) of the species and 93 per cent of the individuals. Without them the specimen : species and indivi- dual : record ratios would be much the same as they are in the Vaunthompsoniinae. In conclusion, the Bodotriidae are the most successful of the cumacean families in southern African waters, both in numbers of species and of indivi- duals. Preliminary estimates suggest that the Diastylidae will prove to be almost as diverse, although not as numerous, while the other families are relatively poorly represented. ACKNOWLEDGEMENTS I should like to thank the following people in South Africa for giving me the opportunity to examine and describe material: Dr Brian Kensley of the South African Museum; Mr Tim McClurg and Mr A. C. Connell of the National Institute for Water Research of the Council for Scientific and Industrial Research; Mr Larry Hutchings of the Sea Fisheries Branch; Dr J. Grindley and the late Dr Robin Boltt. Iam also very grateful to Miss Joan Ellis of the British Museum (Natural History), Dr. N. S. Jones and Dr P. LeLoeuff for allowing me to examine some west African material, and to Dr M. Bacescu of the Museé SOUTHERN AFRICAN CUMACEA: PART 2 289 d’Histoire Naturelle in Bucharest for providing some almost unobtainable literature. Iam grateful, too, to Mrs Sue Frost and Miss Judy Miller for technical and clerical assistance, and to many other people who have provided me with odd specimens found during their own collecting programmes. REFERENCES BAcsscu, M. 1950. Speciile de Iphinoe. Anal. Acad. rom. [Bucharest] 3(12): 1-54. BAcescu, M. 1961. Contributions a l’etude des Cumacés de la Méditerranée et particuliérement des cétes d’Israel. Rapp. P.-v. Réun. Comm. int. Explor. scient. Mer. Méditerr. 16: 495-502. BAcgscu, M. 1975. New Cumacea from the Red Sea. Trav. Mus. Hist. nat. “Gr. Antipa’’ 16: 35-69. BATE, S. 1856. On the British Diastylidae. Ann. Mag. nat. Hist. (2) 17: 449-465. BONNIER, J. 1896. Resultat scientifique de la Campagne du “‘Caudan” dans le Golfe de Gascogne. III. Annls Univ. Lyon. 26: 529-562. CALMAN, W. T. 1904a. Report on the Cumacea collected by Prof. Herdman at Ceylon in 1902. Ceylon Pearl Oyster Fish. suppl. Rep. 12: 159-180. CALMAN, W. T. 19045. The marine fauna of the west coast of Ireland. Scient. Invest. Fish. Brch. Tre. 1; 1-52. CALMAN, W. T. 1905. The Cumacea of the Siboga Expedition. Siboga Exped. Monograph 36: 1-23. CALMAN, W. T. 1907a. On new or rare Crustacea of the order Cumacea from the collection of the Copenhagen Museum. I. The families Bodotriidae, Vaunthompsoniidae and Leuconidae. Trans. zool. Soc. Lond. 18: 1-58. CALMAN, W. T. 19075. Sur quelques Cumacés des cétes de France. Bull. Mus. natn. Hist. nat., Paris 16: 116-123. Day, J. 1975. South African Cumacea. Part I. Family Bodotriidae, subfamily Vaunthomp- soniinae. Ann. S. Afr. Mus. 66: 177-220. ECKMAN, S. 1953. Zoogeography of the Sea. London: Sidgwick & Jackson. Face, L. 1928. Cumacés. Voyage de la Goellette ‘“‘Melita’’ au Senegal (1889-1890). Bull. Soc. Zool. Fr. 53: 331-339, Face, L. 1951. Cumacés. Result. scient. Expéd. oceanogr. belge Eaux cét. afr. Atlant. Sud 3: 1-9. FISCHER, P. 1872. In: FoLin A. G. L. DE & PERIER, L. Les Fonds de la Mer 2: 47. Paris GAmo, S. 1967. Studies on the Cumacea (Crustacea, Malacostraca) of Japan. Part I. Publs Seto mar. biol. Lab. 15: 133-163. Goopsir, H. 1843. Description of the genus Cuma and two new genera nearly allied to it. Edinb. New phil. J. 34: 119-180. GRIFFITHS, C. L. 1974. The gammaridean and caprellid Amphipoda of southern Africa. Unpublished Ph. D. Thesis, University of Cape Town. Hate, H. M. 1944a. Australian Cumacea. No. 7. The genus Cyclaspis. Rec. S. Aust. Mus. 8: 63-142. Hate, H. M. 19445. Australian Cumacea. No. 8. The family Bodotriidae. Trans. R. Soc. S. Aust. 68: 225-285. HALE, H. M. 1953. Two new Cumacea from South Africa. Trans. R. Soc. S. Aust. 76: 45-50. HANSEN, H. J. 1895. Isopoden, Cumaceen und Stomatopoden der Plankton-Expedition. Ergebn. Plankton-Exped. 2: 1-105. JONES, N. S. 1955. Cumacea of the Benguela Current. ‘Discovery’ Rep. 27: 279-292. Jones, N. S. 1956. Cumacea from the west coast of Africa. Atlantide Rep. 4: 183-212. JoNnEs, N.S. 1960. Cumacea from South Africa. Ann. Mag. nat. Hist. (13) 2: 171-180. JongEs, N. S. 1969. The systematics and distribution of Cumacea from depths exceeding 200 m. Galathea Rep. 10: 99-180. Jones, N. S. & SANDERS, H. L. 1972. Distribution of Cumacea in the deep Atlantic. Deep Sea Res. 19: 737-745. Kemp, S. 1916. Fauna of the Chilka Lake. Cumacea. Mem. Indian Mus. 5: 395-402. 290 ANNALS OF THE SOUTH AFRICAN MUSEUM KOssMANN, R. 1880. Zoologische Ergebnisse einer im Auftrage der Koniglichen Akademie der Wissenschaften zu Berlin ausgefiihrten Reise in die Kiistengebiete des Rothen Meeres. IJ. Halfte, I. Lief., II Malakostraken. 88-92. Leipzig. KurIAN, C. V. 1951. The Cumacea of Travancore. Bull. cent. Res. Inst. Univ. Travancore (C) 79° TALS KurRIAN, C. V. 1954. Notes on the Cumacea (Sympoda) in the Zoological Survey of India. Rec. Indian Mus. 52: 275-311. KurRIAN, C. V. 1961. Three species of Cumacea from the lakes of Kerala. Bull. cent. Res. Inst. Univ. Kerala (C) 8: 55-61. LELoEvuFF, P. & INTES, A. 1972. Les Cumacés du plateau continental de Cote d’Ivoire. Cah. Off. Rech. Sci. Tech. Outre-Mer. 10: 19-46. LELoeurF, P. & INTEs, A. In press. Les Bodotria (Crustacea, Cumacea) des mers d’Europe et des cdtes occidentales de |’Afrique tropicale. Bull. Mus. natn. Hist. nat. Paris. MILLARD, N. A. H. 1978. The geographical distribution of southern African hydroids. Aun. S. Afr. Mus. 74: 159-200. MARCUSSEN, J. 1894. Ueber ein neues Cumaceengenus Eocuma, Family Cumadae, aus Japan. Sber. Ges. naturf. Freunde Berl. 1894: 170-171. MontaGu, G. 1804. Description of several marine animals found on the south coast of Devonshire. Trans. Linn. Soc. 7: 61-85. Norman, A. M. 1867. On the Crustacea. ... Report of the committee exploring the coast of the Hebrides. Rep. Br. Ass. Advmt Sci. 36: 193-206. Sars, G. O. 1865. Om den aberrante Krebsdyrgruppe Cumacea og dens nordiske arter. Forh. VidenskSelsk. Krist. 1864: 128-208. - Sars, G. O. 1871. Beskrivelske of fire vestindiske Cumaceer opdagede of Dr A. Goes. Ofvers. K. Vetensk Akad. Forh. 28: 803-811. SARS, G. O. 1878. Middelhavets Cumaceer. Part 1. Arch. Math. Natur. 3: 461-512. Sars, G. O. 1879. Middelhavets Cumaceer. Part 2. Arch. Math. Natur. 4: 1-126. ScoTT, T. 1901. Notes on some gatherings of Crustacea collected for the most part on board the Fishery Steamer ‘‘Garland”’ and examined during the past years. Rep. Fishery Bd. Scotl. 19: 273. STEBBING, T. R. R. 1910. Sympoda. Ann. S. Afr. Mus. 6: 409-419. STEBBING, T. R. R. 1912. South African Crustacea. Part 6. The Sympoda. Ann. S. Afr. Mus. 10: 129-176. STEBBING, T. R. R. 1913. Cumacea. Tierreich 39: 1-210. ZIMMER, C. 1907. Neue Cumaceen von der Deutschen und der Schwedischen Siidpolar- expedition aus der Familien der Cumiden, Vaunthompsoniiden, Nannastaciden und Lampropiden. Zool. Anz. 31: 367-374. ZIMMER, C. 1908. Die Cumaceen der ,,Deutschen Tiefsee-Expedition’’. Wiss. Ergebn. dt. Tiefsee-Exped. ‘Valdivia’ 8: 155-196. ZIMMER, C. 1914. Cumacea. Fauna Siidw.Australiens. 5: 175-195. ZIMMER, C. 1916. Cumacea und Schizopoda. Beitr. Kennt. Meeresfauna Westafr., Crust. 4: 55-66. ZIMMER, C. 1921. Mitteilung iiber Cumaceen des Berliner Zoologischen-Museums. Mitt. zool. Mus. Berl. 10: 117-149. ZIMMER, C. 1936. California Crustacea of the order Cumacea. Proc. U.S. natn. Mus. 83: 423-439. ZIMMER, C. 1941. Cumaceen. Bronn’s KI. Ordn. Tierreichs 5 (1, Book 4): 1-222. ZIMMER, C. 1942. Die Gattung Iphinoe (Ord. Cumacea, Fam. Bodotriidae). Zool. Anz. 139: 190-200. iy yA es ya ai 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 9 6 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. JENNIFER DAY SOUTHERN AFRICAN CUMACEA PART 2 FAMILY BODOTRIIDAE, SUBFAMILY BODOTRIINAE VOLUME 75 PART 8 JUNE 1978 ISSN 0303-2515 CAPE TOWN Se a a. ey INSTRUCTIONS TO AUTHORS 1. MATERIAL should be original and not published elsewhere, in whole or in part. 2. LAYOUT should be as follows: (a) Centred masthead to consist of Title: informative but concise, without abbreviations and not including the names of new genera or species Author’s(s’) name(s) Address(es) of author(s) (institution where work was carried out) Number of illustrations (figures, enumerated maps and tables, in this order) (b) Abstract of not more than 200 words, intelligible to the reader without reference to the text (c) Table of contents giving hierarchy of headings and subheadings (d) Introduction (e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents (f) Summary, if paper is lengthy (g) Acknowledgements (h) References (i) Abbreviations, where these are numerous 3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced with 2,5 cm margins all round. First lines of paragraphs should be indented. Tables and a list of legends for illustrations should be typed separately, their positions indicated in the text. All pages should be numbered consecutively. Major headings of the paper are centred capitals; first subheadings are shouldered small capitals; second subheadings are shouldered italics; third subheadings are indented, shouldered italics. Further subdivisions should be avoided, as also enumeration (never roman numerals) of headings and abbreviations. Footnotes should be avoided unless they are short and essential. Only generic and specific names should be underlined to indicate italics; all other marking up should be left to editor and publisher. 4. ILLUSTRATIONS should be reducible to a size not exceeding 12 « 18 cm (19 cm including legend); the reduction or enlargement required should be indicated; originals larger than 35 x 47 cm should not be submitted; photographs should be rectangular in shape and final size. 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REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: ‘Smith (1969) describes .. .’ ‘Smith (1969: 36, fig. 16) describes...’ “As described (Smith 1969a, 19696; Jones 1971)’ “As described (Haughton & Broom 1927)...’ ‘As described (Haughton et al. 1927)...’ Note: no comma separating name and year Dagination indicated by colon, not p. names of joint authors connected by ampersand et al. in text for more than two joint authors, but names of all authors given in list of references. (b) Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, b, etc. to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a). For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list o, scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number (only if independently paged) in parentheses, pagination (first and last pages of article). Examples (note capitalization and punctuation) BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140. FiscHer, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. gén. 74: 627-634. Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320. KOHN, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51. THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zcologische 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) rr — ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 75 Band June 1978 Junie Part 8 Deel 4 a, S 4110, at VID nov X LOBEDU POTTERY By PATRICIA DAVISON & JUNE HOSFORD 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 aes), S93), SLD), ALS, B, than, SCS, S, TO), 6(1, t—p.i.), 71-4), 8, 9(1-2, 7), 10(1-3), GD, 8, Wh, bea), ISCS), BAO), D7, SIGS), BAS), 38 Copyright enquiries to the South African Museum Kopieregnavrae na die Suid-Afrikaanse Museum ISBN 0 908407 44 0 Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk., Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap LOBEDU POTTERY By PATRICIA DAVISON & JUNE HOSFORD South African Museum, Cape Town (With 28 figures and | table) [MS. accepted I February 1978] ABSTRACT In this paper the technique and the types of pottery in use among the Lobedu of the northern Transvaal are described. Recent examples are compared with museum specimens collected in the 1930s, and possible reasons for the continuity of the craft are discussed. CONTENTS PAGE lintiroclnctior, 5 » +o »« o ZAP TONS NOUS 5 4 5 6 6 5 | OMB Materials and technique . . 294 TINVOES BMG WSS o 5 6 ol CS CUB IDISCUSSIONN sen ee llG Acknowledgements Wecie e LS IRGIGMINCSS 5 5 0 « « o BY INTRODUCTION The Lobedu live in the northern Transvaal lowveld below the Drakensberg escarpment and are ruled by a female chief, Modjadji, the legendary ‘Rain Queen’. Culturally they are classified with the North Sotho branch of the Bantu-speaking peoples of southern Africa, but they show the marked influence of their northern neighbours, the Venda, with whom they have strong historical links (Van Warmelo 1974: 78). They have many cultural characteristics in common with other lowveld Sotho, for example the people of Phalaborwa, but relatively little in common with highveld North Sotho peoples such as the Pedi. Their social structure has been described in detail (Krige & Krige 1943; Monnig 1963: 49-64), but very little has been written on their material culture. Many of the people who recognize Modjadji as their chief do not, in fact, trace their descent from a Lobedu ancestor, but belong to other Sotho groups who have been assimilated through intermarriage, or to the Tsonga who have lived in close proximity to the Lobedu for many years. In the 1930s the Kriges (1943: 85) estimated that only 10 per cent of the Lobedu people were descended from the original founding group. From the late nineteenth century onwards missionaries and traders have been in the area, and their influence, together 291 Ann. S. Afr. Mus. 75 (8), 1978: 291-319, 28 figs, 1 table. 292 ANNALS OF THE SOUTH AFRICAN MUSEUM with the system of migrant labour, have made European wares increasingly accessible and desirable. Lobedu pottery is remarkably conservative and many potters still make pots in the traditional way. As in the case of all other Lobedu crafts, pottery is not a full-time occupation. It takes its place in the seasonal cycle of domestic and agricultural routine. During the dry winter months craftwork is at a peak, as Krige & Krige (1943: 32) comment: ‘From May, therefore, there are not only dances and much beer-drinking but there is time for the plying of crafts such as pottery, mat-making, basket-making. For the specialist in any craft is always primarily an agriculturist who plies his craft only when he has time, and it is in vain that anyone approaches even the most skilled craftsman for an object during the planting or weeding season’. A further practical reason which favours the making and sale of pottery in the dry season is that for a successful firing the ground must be completely dry. During the wet summer months, even if a potter does find time to make pots, she will keep them in her cooking-hut until she considers the conditions right for firing; it is common to find between twenty and sixty unfired pots piled up inside a potter’s hut awaiting suitable firing conditions (Fig. 1). The following technical account is based on data recorded between 1973 and 1976 from eleven potters in Modjadji’s village and in the surrounding areas. All the potters were observed at work on several occasions although the whole process could not be followed through in every case. Observation was supple- mented by asking the potters about their techniques. The description of the types of pottery is based on specimens in the ethno- Fig. 1. Unfired pots inside hut, Modjadji’s village, 1975. LOBEDU POTTERY 293 graphic collections at the South African Museum, specifically the Krige Collec- tion which was collected between 1936 and 1938, and comparative specimens collected by the authors between 1973 and 1976. THE POTTERS Among the Lobedu, pottery is always made by women. Not all women who make pots, however, are specialists. Almost every woman has seen pots being made and knows the technique, but in practice some women prefer to buy pots rather than make them. Many women make their own cooking-pots but buy the decorated pots for drinking beer. Even a good potter will buy pots from other potters if she needs a special pot for a particular purpose or simply if she likes the pot. Some potters take their pots to sell at the local market, but others keep them at home (Fig. 2) and wait for buyers to come to them. Traditionally pots were exchanged for the amount of grain they held or in return for performing a task like weeding (Krige 1941: 11, 17). This was still so in the 1930s but today pots are usually exchanged for money. Fig. 2. Fired pots for sale at potter’s home, Molototsi valley, 1973. Highveld Sotho potters sell their wares over a wide area and new ideas for shape and decoration are introduced in this way. Examples in the Krige Collec- tion show that potters are quick to imitate anything which appeals to them, from a jam-jar to the shape of a motor-car tyre. None of the potters put any identification mark or stamp on to their pottery, but almost every woman 294 ANNALS OF THE SOUTH AFRICAN MUSEUM questioned said she could always recognize the pots which she had made because ‘she had made them and knew them well’. Most of the skilled, specialized potters are older women whose children have grown up and who have daughters-in-law to help with the domestic chores. Indeed a good potter may become wealthy enough to acquire a ‘wife’ of her own to cook for her. Following the precedent set by the Queen who has ‘wives’, women of importance who have cattle of their own may use the cattle to marry a younger woman who will perform all the domestic duties of a young wife or daughter-in-law (Krige & Krige 1943: 144). Thus freed from some of the house- hold tasks, a potter has more time for other activities including pottery. Many of the potters had learned how to make pots by watching an experi- enced potter and then by experimenting themselves. The craft was not neces- sarily passed on from mother to daughter. Only one of the potters questioned had learned the technique from her mother. Daughters of the present potters show very little interest in learning the craft. MATERIALS AND TECHNIQUE There seem to have been no major changes in pottery processes over the past forty years. Recent fieldwork accords with E. Krige’s (South African Museum correspondence 3.3.66) description for the 1930s. The tools used today are similar to those collected in the 1930s and also recorded in the 1960s (Lawton 1967: 172). The present writers did not see any Lobedu potters using the method of continuous coiling which was recorded by Lawton (1967: CFE GR TT re C Ao gh . 4 - 26, 7h > : oe 2 Fig. 3. Collecting clay from river-bank, Molototsi valley, 1975. LOBEDU POTTERY 295 176) and stated by Shaw (1974: 116). As Lawton and Shaw based their account on a demonstration by a single potter, it seems possible that this was an excep- tional case. The clay Suitable clay, /edzuba, is found in the valley below the chief’s village. Two sources, one about a kilometre from the village, the other about 3 kilometres away, are used by the local potters. In both places the clay, probably kaolinite, is dug from the banks of dry river-beds (Fig. 3), but the two clays are of different colour and texture. The darker, finer clay, which comes from the further site, requires only kneading before use, but the distance from the village is a dis- advantage. The yellowish. coarser clay which occurs nearer has more impurities and requires more preparation before use. The dry clay is ground, ho seda, to a fine powder and all the large particles are removed. Thereafter it is covered with water and allowed to stand for a few days after which it is kneaded and pounded with a small pestle on a concave stone until a good workable consistency is achieved (Fig. 4A). No tempering material is added. Method (Figs 4B-D, 5) Like all other Bantu-speaking peoples, the Lobedu do not use a potter’s wheel. By working on a movable base (a potsherd, an enamel basin or plate) the potter is able to rotate the pot when necessary. For all shapes of vessel the method is the same. The start is made by placing a rough ring of clay on the base. From the ring the lower part of the pot is built up, by pinching, scraping, smoothing by hand, and adding pieces of clay. When it is roughly the required shape it 1s smoothed with a shell, thema, of the seed pod of the mothema creeper (Bauhinia kirkii Oliv.). The thema is soaked in water to become pliable and is then used in a variety of ways—the convex side for smoothing, the concave side for scraping, and the edge for shaping. At first the thema is used to smooth the outer wall in an upward, diagonal movement while the other hand supports the wall on the inside and presses lightly to thin out, heighten and shape the sides. Small rolls of clay are added to the sides and worked into the body of the pot. The number of pieces added depends on the size of the pot. For a very large pot it may be necessary to let the walls harden slightly before building it up to its complete height. Excess clay from the inner wall is scraped off and any irregularities in the clay are removed by hand or with an awl. The walls are smoothed by rubbing a small, smooth river pebble, thidelo, over the surface of the pot. Particular care is taken in making the rim as this is a potential area of weakness in a pot. In addition, a pot is judged by the neatness of its rim. The upper edge is flattened evenly, using the forefinger and the thumb, and a thin roll of clay is carefully added and smoothed into position. To give a very smooth finish a piece of wet calf-skin is folded over the rim and moved deftly round it (Fig. 6). The rim profile is defined by modelling with the thema. 296 ANNALS OF THE SOUTH AFRICAN MUSEUM A cloth is wrapped round the pot to protect it from draughts and changes in temperature which might cause the clay to dry unevenly and crack. It is taken indoors until it is dry enough to be inverted for the filling in of the base. Fig. 4. First stages in making a pot, Modjadji’s village, 1975. A. Preparing the clay. B. Positioning a rough ring of clay on the base. C. Starting to build the wall. D. Adding small pieces of clay. LOBEDU POTTERY 297 In the case of a small pot the base is usually finished on the following day, but a large pot may require up to a week before it is ‘strong enough’ to stand on its rim without the walls collapsing. Fig. 5. Shaping the walls. A. Scraping off excess clay. B. Smoothing the inner wall. C. Adding pieces of clay to give extra height. D. The pot has reached its final shape and must be left to harden before the rim can be made; the potter takes a pinch of snuff. 298 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 6. Smoothing the rim with a piece of wet calf-skin. If the pot is to be decorated, incised designs are applied before the pot is taken off its movable base (Fig. 7A—B). Geometric designs are outlined with a piece of sharpened metal and chosen areas are filled in with red ochre and graphite. The ochre, /edzogu, is powdered and mixed with water in a small bowl or sherd. It is applied with the finger-tip and then burnished with asmooth river stone (Fig. 7C). On the following day when the pot is drier, graphite, Phomu, is applied in the same way and it is burnished to a metallic sheen. When the pot is firm but still pliable, it is inverted and removed from the base. The thick edges of the open base are pinched inwards and excess clay is scraped off. Small pieces of clay are added (Fig. 8A) until the opening is com- pletely closed. The base is then smoothed with the thema (Fig. 8B) and patted into shape with a light wooden beater, khebato. At this stage uneven wall thickness may be corrected by scraping and smoothing the inner walls (Fig. 8C). Finishing touches may be applied to the decoration and thereafter the pot is set aside in a hut (Fig. 9) for at least a week, and often for much longer, until the firing takes place. Firing (Fig. 10) A number of pots are fired together, except in the case of a very large pot, which is fired singly. A calm, clear day is preferred and the site is generally sheltered from the prevailing wind. Firing usually takes place in the evening when there is said to be less wind. During the day the potter takes dry bark and kindling to the firing place, which is a shallow depression in the ground. Towards LOBEDU POTTERY 299 Fig. 7. Decorating a pot for drinking beer. A. Incising the design. B. Applying red ochre. C. Burnishing the colour with a river-stone. 300 ANNALS OF THE SOUTH AFRICAN MUSEUM C Fig. 8. Completing the base of a pot. A. Filling in the opening. B. Smoothing the closed base with a seed-pod; note beater and burnishing stone. C. Scraping off excess clay from inside. LOBEDU POTTERY 301 Fig. 10. Firing. A. Positioning the pots and placing bark inside and round them. B. Covering them with dry kindling. C. Old thatching grass is placed on top. D. The grass is set alight. 302 ANNALS OF THE SOUTH AFRICAN MUSEUM sunset the pots are positioned and supported on small stones. Bark is stacked inside and round the pots; kindling and old thatching grass are piled on top and ignited. As the grass burns, more is added until the flames are smothered and the burnt grass is insulating the smouldering coals within. The bark burns slowly and without a flame. After checking with a long stick that all the pots are surrounded by glowing coals, the potter leaves the fire to burn overnight. The pots are allowed to cool (Fig. 11) and are finally taken home. Fired pottery is porous and not fully waterproof. Pots for cooking and brewing beer are sealed through use. In pots for storing water porosity is an advantage as it allows the water to cool. On other pots the application of graphite or ochre, followed by burnishing, may make them less porous, but it was not asked whether burnishing was done consciously for this reason as well as for the more obvious decorative purpose. Breakages in the firing are usually explained in terms of poor quality clay or the excessive heat of the fire. None of the potters questioned put breakages down to supernatural causes. This accords with Krige & Krige’s (1943: 35) observation that technical success is based on practical knowledge of the environment, in this case the quality of the clay, rather than on manipulation of “mysterious forces’. Repair Cracked pots are repaired with beeswax. A well-made /edz6méla in the Krige Collection, SAM—9801, has been carefully caulked with beeswax and is held together with fine wire round the rim. Fig. 11. Pots cooling after a large firing. LOBEDU POTTERY 303 TYPES AND USES The Lobedu differentiate pottery vessels primarily according to function, which generally determines the shape, size and decoration of a pot. The pots in the Krige Collection cover the full range of Lobedu pottery except the very big beer-pots which were too large to remove from the locality. The 1930s specimens are described and illustrated here and are used as a standard of comparison for the specimens and data collected in the 1970s. As the number of pots in each group is very small, no statistical analysis has been attempted. Average dimensions and the range are given in centimetres; volume is given in litres and the number of pots measured in each group is indicated after the year. “Nkho’ (Fig. 12) The nkhoé is a very large, spherical beer-pot, usually decorated on the shoulder and round the mouth. Traditionally these pots are used on social occasions when many people are drinking beer. If a family does not have its own nkhé one is borrowed from relations or neighbours (E. Krige 1977, pers. comm.). In many places the half 44-gallon drum has replaced, or is used con- BWI Fig. 12. Beer being served from a relatively small nkhoé into a glass jar, Modjadji’s village, 1975. 304 ANNALS OF THE SOUTH AFRICAN MUSEUM Ag % 3 . el Fig. 13. The old and the new, Modjadji’s village, 1975. currently with, the nkho for brewing beer on a large scale (Fig. 13). The beer-hut of the Molokwane family (1975) contained six very large beer-pots partly embedded in the ground, two metal drums in which the beer had been brewed, and two smaller beer-pots from which the beer was served. Outer mouth Diam. Ht diam. Vol. 1970S (3) se semy fo ae ONO 37,0 26,1 not 40,0-51,9 32,0-43,3, 22,0-29,8 measured ‘Modzhéha’ (Fig. 14) This is a large, spherical pot usually decorated with bands of cross-hatching on the shoulder and incised geometric designs round the mouth; it is used for carrying and storing water. These pots are still much in use today as water keeps fresh and cool in the porous earthenware. A lighter tin container is widely used to carry water, but there is a modzhéha at every home for storing water (Fig. 15). The shapes of the pots of the 1930s and 1970s show great similarity, as does the decoration. Outer mouth Diam. Ht diam. Vol. 1930s'Q). 42,4 32,6 21,8 29,0 41,1-43,7 32, 1—33,0 21,5=22,2 25,6-32,4 1970s (8) . 41,4 3,3) Dp) I) 38,6—-45,0 30,0-37,5 17,8-30,8 23,9-32,0 LOBEDU POTTERY 305 WALMUMILLLLHIHUMIIE SS SSEIIIUE f 0 9g 18 _————— cm Fig. 14. Modzhéha, SAM-9712, collected 1930s. ‘Thukhwana’ (Figs 16-18) A fairly large, spherical or elliptical pot from which beer is served into drinking vessels. A thukhwana is always decorated, traditionally with incised designs coloured with graphite and red ochre but nowadays with enamel paints. Fig. 15. Modzhéha with calabash ladle close at hand, Modjadji’s village, 1975. 306 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 16. Thukhwana, SAM-9793, 9791, collected 1930s. Two examples showing variations in shape and decoration. A. Elliptical shape, inspired by a motor-car tyre. B. Spherical pot, decorated with incised textured triangles and bands. Size, shape and design vary greatly. These and the pots for drinking beer, which are the most ornately decorated pots, are used for entertaining. They may be considered ‘party-ware’ and are more susceptible to changes in fashion than other pots. In the 1930s the elliptical-shaped pot (Fig. 16A) was a recent innovation. This style, called ‘ya lebile’, which was said to have been inspired by the tread and shape of a motor-car tyre, was adopted from Dikxale potters on the highveld and became very popular among the Lobedu in the 1930s (E. Krige 1977, pers. comm.). Highveld Sotho influence is also seen in the arc design and the use of stamped lines on some of these pots (SAM-9797, LOBEDU POTTERY 307 UCT 38/83). An unusual feature is a stamped star design on the base (Fig. 17B). Outer mouth Diam. Ht diam. H9B0S(9) . .. . . . 30,0 19,9 14,7 22,4—38,5 15,5-23,3 12,2-17,5 MOSS), . « sw 6277 DED, Se 21,0—35,0 16,8—27,0 15,5-22,0 Fig. 17. Thukhwana, SAM-9797, collected 1930s. A. Spherical pot; decoration shows highveld Sotho influence in the arcs and in the stamped lines. B. Star design on base of pot. Vol. tE6 3,4-11,3 4,6 (only one measured) 308 ANNALS OF THE SOUTH AFRICAN MUSEUM ‘Ledzhoméla’ (Fig. 19) Small, decorated pot used for an individual serving of beer to an honoured guest. The smallest ones are also used to store baby food. Their shapes vary, but they are always decorated with incised, coloured designs. A 1930s specimen (Fig. 19B) in the Krige Collection is based on the shape of a jam-jar seen in Duiwelskloof, the nearest town. Many modern examples are flat-based. The ledzhéméla was borrowed from Shangaan neighbours, as was the term used to describe it (Lawton 1967: 177), and it has largely replaced the calabash vessel, phafa, which was the traditional Lobedu beer-cup. Outer mouth Diam. Ht diam. Vol. 1930s (@).. 3 2 1 oe Ble 12,6 9,4 ILS 15,4-17,4 10,5-15,1 8,8-10,0 1,2-2,1 L9OT0SO) 2 ey eee Sh) 12,8 12,9 1,8 16,9-18,6 12S) 12,5-13,5 1,6-2,0 *Bidzha ya khelalélé’ (Fig. 20) A spherical pot made in various sizes. The large ones are used mainly for cooking porridge and the smaller ones for preparing vegetable relish or meat. Decoration on these pots is minimal, usually a single line or band of cross- hatching on the shoulder and graphite colouring round the rim. They blacken Fig. 18. Pots in washing-up area —thukhwana, modzhéha, larger thukhwana, morifhi in foreground, Modjadji’s village, 1975. LOBEDU POTTERY eS OLR RPO COKER OE Cc Fig. 19. Ledzhéméla, SAM-—9800, 9801, col- lected 1930s; SAM-—10603, collected 1976. A. Small, decorated pot for drinking beer. B. This shape was inspired by a jar seen in Duiwelskloof. C. Modern /edzhémeéla, with flat base. 309 310 ANNALS OF THE SOUTH AFRICAN MUSEUM rapidly on the hearth and become seasoned and non-porous through use. The simplicity of decoration and lack of red ochre distinguishes cooking-pots from similar pots not used for cooking. Diam. 19308: GB) Gr oe Se Vass 31,7-41,4 LO7OS(3) ec Re eae ese 27,1-30,7 The dimensions for small pots, bidzhana, are: 193052) ee Ont 18,4-23,9 1970s) a a es ‘Morifhi’ (Figs 21-2) Ht 24,6 223-2950 202 19,3-21,4 1356 12,3-14,9 14,0 Outer mouth diam. DSO 22,8-28,5 D383) 22,0—24,6 16,3 12,7-19,8 Syl Vol. 17,0 11,3-26,4 6,9 (only one measured) 259 1,8-3,9 1,8 An open-mouthed bowl used for cooking certain relishes, such as termites and locusts, which do not need to be covered. It is also used as a lid for a cooking- pot and has many household uses. A morifhi is never decorated with incised designs. Occasionally graphite is applied to the rim as a finishing touch. Diam. L930SI(G)) 2 el OED 26,2-30,0 1970852) oe a ent i 9) 28,0-30,0 *‘Kheritswana’ (Fig. 23) Ht 10,3 10,0-10,5 10,4 9,8-11,0 Outer mouth diam. 26,1 24,8—-28,6 28,1 26,2-30,0 Vol. ml 2,9-3,6 2,8 (only one measured) A fairly small open-mouthed bowl used for serving relish and for drinking marula-cider, mokope. A kheritswana is always burnished with graphite both inside and out. This distinguishes it from a morifhi of similar shape. It is often, but not always, decorated with an incised, textured band near the rim. Modern examples are sometimes made with flat bases. Outer mouth Diam. Ht diam. Vol. 193086) 3 2) ee LOR 8,2 18,4 13 17,6-23,4 SO SIUL,33 16,0-21,8 0,8—2,0 LOTOS(@)) 3, ee ee ee 8,6 16,3 L@ 16,6-18,2 8,5-8,6 15,4-17,2 1,0-1,0 LOBEDU POTTERY Sil cm Fig. 20. Bidzha ya khelalél6, SAM-9706, 9785, collected 1930s. A. Large size for making porridge, with lid for steaming. B. Smaller size for cooking relish. 0 18 | cm Fig. 21. Morifhi, SAM-9706, collected 1930s. Fig. 22. Woman cooking relish in her courtyard. The pumpkin leaves in the bowl, morifhi, are cooked in a small pot, bidzhana ya moréhé, on the hearth, Lebiya’s place, 1975. Bile ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 23. Kheritswana, SAM-9788, collected 1930s; SAM-—10601, collected 1975. A. Traditional bowl for drinking marula cider. B. Pot for the same use, but with a flat base. Fig. 24. Lebéda, SAM-9787, collected 1930s. ‘Lebéda’ (Figs 24-25) A large open-mouthed pot, roughly half-spherical in shape, used for steeping maize prior to stamping and during the beer-making process. When woven beer-strainers were used, beer was always strained into a /ebéda (Krige & Krige 1943, fig. 11), but this is no longer done, as beer is now brewed on a much larger scale in tin drums. The /ebéda is still made and used for soaking grain, particularly during and just after the reaping season. In June 1976 a number of potters were making them for their own use and for sale, as the demand was great at this time of year. They are usually undecorated except for an incised band below the mouth. Outer mouth Diam. Ht diam. Vol. 19308 @)) = 2a. eee DO) 41,3 19,0 41,5-46,1 22,0-22,3 38,0-44,5 17,5-20,5 IS7Osi(Q)o2. pe, a le 27,0 Byeu 31,9 37,0-45,4 22,0-32,0 36,0-39,3 (only one measured) LOBEDU POTTERY 313 Na a ee Fig. 25. Pots in use; left to right modzhéha, nkho, thukhwana, lebéda; Modjadiji’s village, 1975. ‘Lesabélo’ (Fig. 26) A wide-mouthed bowl used as a wash-basin. It is always decorated with a broad, incised band of hatching round the rim. White chalk is rubbed into the incisions, and the bowl is burnished with graphite inside and outside. It was customary for a wife to bring her husband a /esabél6 of warm water for washing in the morning. In the 1930s Krige recorded that these bowls were rare and had been replaced almost entirely by enamel basins. For ritual purposes, however, the traditional bowl is said still to be used, for example, a person who is possessed by spirits must use a /esabélé for washing. In 1976 a potter at Modjadji’s village made an example on request as none had been seen in use. It differed in shape from the 1930s specimens in that it was not as large and the incised decoration was not coloured with white chalk. Fig. 26. Lesabélé, SAM—9804, collected 1930s. 314 ANNALS OF THE SOUTH AFRICAN MUSEUM Outer mouth Diam. Ht diam. Vol. 19308'(3) 3 Sue ee ee 10,9 iD 3,0 27,9-29,0 10,3-12,0 26,0—27,5 2,8—3, 1 TOPOS) wi ee er OL 9,1 DRS 0,9 ‘Khetsikhiyo’ (Figs 27-8) A deep bowl with a heavy, flat base and striated inner walls, used for grinding tobacco into snuff. Occasionally there is an incised band near the rim but many examples are undecorated. They are still in use, especially by old women who enjoy taking snuff. Potters make them on order. Outer mouth Diam. Ht diam. Vol. 1930812)" Ee ee eet n) IS 16,1 not 17,5-18,5 10,0-13,0 15,5-16,6 measured 1970s . . . . +. +. No specimens collected or measured but those seen did not appear to differ from the 1930s specimens. Fig. 27. Khetsikhiyo, SAM-—9802, collected 1930s. DECORATION The thematic features of Lobedu pot-decoration are incised patterning, often in the form of textured bands of hatching or cross-hatching alone or in combination with other geometric designs, and colouring with burnished graphite and red ochre. White chalk is used only on traditional wash-basins. Enamel paints are a recent innovation, rarely used by the potters at Modjadji’s village. LOBEDU POTTERY 315 Fig. 28. An old woman at the chief’s village making snuff, Modjadji’s village, 1973. The long stick grinds the tobacco in a rotating action. Together with shape and size the presence or absence of decoration charac- terizes the various types of pot. The lack of decoration on a morifhi distinguishes it from a kheritswana, which is of similar shape but burnished overall with graphite. Pots used on the hearth are very simply decorated and, by contrast, pots for serving and drinking beer are highly decorated and have the greater variation and innovation in design. Table 1 summarizes the types of Lobedu pottery in terms of function, shape, size and decoration. TABLE | Types of Lobedu pottery. Type Function Shape and size Decoration INGO) ox storing beer, spherical; very incised patterns between formerly for brewing large shoulder and rim; graphite beer and ochre colouring Modzhéha . carrying and storing spherical; large textured bands on shoulder, water geometric designs between shoulder and rim; graphite and ochre colouring 316 ANNALS OF THE SOUTH AFRICAN MUSEUM Type Function Shape and size Decoration Thukhwana . serving beer spherical or ellipti- incised bands, triangles, cal; wide range arcs and chevrons between rim and widest diameter; graphiteand ochrecolouring Ledzhoméla . serving beer, storing roughly spherical, incised bands, chevrons and baby food modern examples diamond shapes over most flat-based; small of the pot; graphite and ochre colouring Bidzha ya cooking pot (literally spherical; wide incised band on shoulder; khilalél6 ‘a pot of supper’) range graphite round mouth Morifhi . bowl for cooking bowl; wide range no incised design; some- relish; lid times a band of graphite round mouth Kheritswana drinking marula bowl; small band of cross-hatching, cider usually coloured with red ochre, near the rim; bur- nished with graphite inside and outside Lebéda steeping grain half-spherical ; little decorated; line of large incised pattern below mouth; graphite between rim and incising Lesabél6 wash-basin bowl; medium a wide textured band of hatching round the rim; white chalk rubbed into incisions; burnished with graphite inside and outside little decorated; occasional incised band and graphite colouring below rim Khetsikhiyo . deep bowl, striated inner wall, heavy flat base; small DISCUSSION Pottery continues to be made and used by the Lobedu despite availability of substitutes from local traders. This continuity merits discussion. Although the conservatism of pottery at Modjadji’s village could be partly explained by the fact that it is the chief’s village, for the same reason it has been less isolated than other villages and more exposed to outside influence. Krige & Krige (1943: 50) commented on the decline in craftsmanship and the impoverishment of traditional culture due to changing social and economic conditions in the 1930s. The migrant labour system meant that men were working in towns, acquiring new material needs and returning home with money to buy manufactured goods. Prestige was attached to the possession of bicycles, sewing machines, suitcases, furniture and modern tools and utensils. Changes in the material culture reflected major changes in the economy. In the case of pottery, the apparent lack of change is significant. Explanation probably lies in a number of sociological and economic factors. Traditionally pottery was not only of functional value. Objects associated with important sociological and ritual activities were prescribed by custom. ‘Material culture may be regarded as part of social relations, for material objects are chains along which social relationships run’ (Evans-Pritchard 1940: grinding snuff LOBEDU POTTERY SET) 89). The highly institutionalized rain-cult of the Lobedu, with the Queen at its centre, demanded a wide-ranging observance of custom and made for ritualized conservatism. Women played important roles in the social structure and mini- mized the disruptive effects of migrant labour. In a subsistence economy based primarily on agriculture, reciprocity in social relationships was stressed (Sansom 1974: 274). The social significance of beer exemplifies this clearly. Beer is a means of ‘thanking’ or paying for specialized services, of reconciliation after a dispute, of honouring a bridegroom, of showing hospitality, of offering tribute, and a necessary part of many rituals (Krige 1932: 343-57). In this context the pottery and calabash vessels used in making, transporting, serving and drinking beer are integral parts of customary practice. It is suggested that where the sociological significance of an activity is maintained, the use of associated traditional utensils is more likely to continue. The large metal drums, nowadays used for brewing beer, and the glass jars used for drinking, reflect a change in the pattern of beer-drinking. Beer is now brewed on a much larger scale, and it is offered for sale. In a traditional beer-drink a calabash is passed freely from person to person and from group to group, showing generosity and bonding ties of kinship and friendship. When beer is sold different principles operate. The maker brews a large quantity of beer. A glass jar or tin can is used in preference to a pot or calabash as it is a more standard measure of volume and the buyer drinks most of the beer because he has paid for it. In these circumstances the use of traditional utensils is not relevant. Today both kinds of beer-drinking co-exist (E. Krige 1977, pers. comm.). In addition to the sociological importance of the activities in which pottery is used, there are economic reasons why pottery has continued to be made and used. The availability of clay allows the craft to remain active. Two good sources of clay, fairly near to Modjadji’s village, provide potters with as much clay as they need and there are no restrictions on its use. Furthermore, the use of pottery has practical advantages. The porous quality of earthenware keeps water cool and fresh. A lighter plastic or tin container may be used for carrying water but a pot is invariably used for storing it at home. Cooking-pots have proved their usefulness over the years and efficient substitutes, like the heavy three-legged iron pot, are expensive. Today pottery is still cheaper than enamel- ware and plastic, if not as durable. For certain pots, such as the khetsikhiy6, for grinding snuff, no similar substitutes can be purchased. The fact that women are the makers and users of pottery has encouraged the continuity of the craft. In the 1930s women did not leave the villages to earn money and, although their agricultural work-load was increased through the absence of men, they were still able to do craftwork in their spare time. Most women could make pottery to supply their needs. A few Christian women made pots for sale, primarily to pay for school fees, clothing and things associ- ated with their westernized way of life. Forty years later the population has almost doubled and there is increased dependence on the earnings of migrant 318 ANNALS OF THE SOUTH AFRICAN MUSEUM labour to buy food. No family can produce enough food from its fields to support itself. In 1969 Modjadji’s area could produce only 12 per cent of its grain needs (Hattingh 1976: 360). Selling pottery is one of the few ways in which women can earn money at home. Many village women do casual labour on local farms and tea-plantations to make money and it is possible that their buying power has stimulated the trade in pottery. The active potters supply women who do not make pottery themselves but who still use it. Changing economic conditions have led to greater specialization and commercialization. The pottery continues to be made but it is meeting new needs. In the long term, if women continue to leave the villages and the process of westernization accelerates, it is probable that the demand for pottery will decrease in favour of factory-produced goods and the craft will decline unless it finds new markets. Continuity does not imply inflexibility. The viability of a craft is proved by its ability to adapt. Technically Lobedu pottery is remarkably consistent, but innovation in design and borrowing of ideas are not uncommon. Lobedu pottery is not unique and it did not develop in isolation—it is closely related to Venda pottery (Van der Lith 1972: 202-45; Schofield 1948: 178) and one can easily detect affinities with highveld Sotho and Shangaan pottery, both in appearance and in the vernacular terminology. A number of Lobedu pottery terms have equivalents in Venda, Sotho and Tsonga. This is not unexpected considering the heterogeneous composition of the Lobedu and their contact with neighbouring groups. A detailed analysis of linguistic affinities has not been attempted by the authors but might be rewarding. A number of the factors discussed above help to explain the continuity of the Lobedu pottery tradition over the past forty years. The flexibility shown in adapting to changing economic circumstances is as important as traditional conservatism. ACKNOWLEDGEMENTS The University of Stellenbosch is thanked for permission to publish data which will form part of Patricia Davison’s Master’s thesis in Anthropology, under the supervision of Prof. G. M. K. Schiiler. Our warmest appreciation goes to Prof. E. J. Krige for her help in planning the fieldwork and for her comments on the manuscript, to Simeon Modyjadji, our interpreter, for his willing and efficient guidance, and to all the potters, in par- ticular to Modabodi Molokwane and Madula Rabothada. Thanks are also due to Dr N. J. van Warmelo for checking the vernacular terms and to Mr V. Branco for the drawings. For permission to work in the area we are indebted to the Department of Plural Relations and Development and to the Lebowa Government. Financial assistance from the Human Sciences Research Council is gratefully acknow- ledged. Opinions expressed and conclusions drawn are those of the authors and should not be regarded as representative of the Human Sciences Research Council. LOBEDU POTTERY 319 REFERENCES EvANS-PRITCHARD, E. E. 1940. The Nuer. Oxford: Clarendon Press. HATTINGH, P. S. 1976. Population pressure, agriculture and planning: the example of the Modjadji Location. S. Afr. J. Sci. 72: 359-361. KriGE, E. J. 1932. The social significance of beer among the Balobedu. Bantu Stud. 6: 343-357. KriGE, E. J. 1941. Economics of exchange in a primitive society. S. Afr. J. Sci. 9: 1-21. KricE, E. J. & KRiGgE, J. D. 1943. The realm of the Rain Queen. London: Oxford University Press. Lawton, A. C. 1967. Bantu pottery of southern Africa. Ann. S. Afr. Mus. 49: 1-440. MOonnica, H. O. 1963. The structure of Lobedu social and political organisation. Afr. Stud. 22: 49-64. SANSOM, B. 1974. Traditional economic systems. Jn: HAMMOND-TOOKE, W. D., ed. Bantu- speaking peoples of southern Africa: 135-176. London: Routledge & Kegan Paul. SCHOFIELD, J. F. 1948. Primitive pottery. Cape Town: The South African Archaeological Society. SHaw, M. 1974. Material culture. Jn: HAMMOND-TOOKE, W. D., ed. Bantu-speaking peoples of southern Africa: 85-131. London: Routledge & Kegan Paul. VAN DER LitH, A. A. 1972. Die keramiek van die Venda. Jn: ELorr, J. F. & COERTZE, R. D. reds. Etnografiese studies in suidelike Afrika: 202-245. Pretoria: Van Schaik. VAN WARMELO, N. J. 1974. The classification of cultural groups. Jz; HAMMOND-TOOKE, W. D., ed. Bantu-speaking peoples of southern Africa: 56-84. London: Routledge & Kegan Paul. 6. SYSTEMATIC papers must conform to the International code of zoological nomenclature (particularly Articles 22 and 51). Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov., etc. An author’s name when cited must follow the name of the taxon without intervening punctuation and not be abbreviated; if the year is added, a comma must separate author’s name and year. The author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is transferred from its original genus. The name of a subsequent user of a scientific name must be separated from the scientific name by a colon. Synonymy arrangement should be according to chronology of names, i.e. all published scientific names by which the species previously has been designated are listed in chronological order, with all references to that name following in chronological order, e.g.: Family Nuculanidae Nuculana (Lembulus) bicuspidata (Gould, 1845) Figs 14-15A Nucula (Leda) bicuspidata Gould, 1845: 37. Leda plicifera A. Adams, 1856: 50. Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b). Nucula largillierti Philippi, 1861: 87. Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. Note punctuation in the above example: comma separates author’s name and year semicolon separates more than one reference by the same author full stop separates references by different authors figures of plates are enclosed in parentheses to distinguish them from text-figures dash, not comma, separates consecutive numbers Synonymy arrangement according to chronology of bibliographic references, whereby the year is placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable. In describing new species, one specimen must be designated as the holotype; other speci- mens mentioned in the original description are to be designated paratypes; additional material not regarded as paratypes should be listed separately. The complete data (registration number, depository, description of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.: Holotype SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973. Note standard form of writing South African Museum registration numbers and date. 7. SPECIAL HOUSE RULES Capital initial letters (a) The Figures, Maps and Tables of the paper when referred to 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.’ ace 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. 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