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ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
VOLUME 77 BAND
THE TRUSTEES OF THE DIE TRUSTEES VAN DIE
SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM
CAPE TOWN KAAPSTAD
1979
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ANNALS OF THE ANNALE VAN DIE
SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM
VOLUME 77 BAND 77
LIST OF CONTENTS
APPLETON, C. C.
The Unionacea (Mollusca, Lamellibranchiata) of south-central Africa. (Published
May 1979.) : a:
BARNARD, J. L.
Redescription of Plioplateia K. H. Barnard, a genus of amphipod (Crustacea) from
South Africa. (Published December 1978.) in = we is
Cooper, M. R. & KENNEDY, W. J.
Uppermost Albian (Stoliczkaia dispar Zone) ammonites from the Angolan littoral.
(Published June 1979.) : mac a oe é
De Freitas, A. J.
A new genus and species of the penaeoid family Solenoceridae (Crustacea, Deca-
poda) from south-east African waters. (Published February 1979.)
KENNEDY, W. J. see Cooper, M. R.
KENNEDY, W. J. & KLINGER, H. C.
Cretaceous faunas from Zululand and Natal, South Africa. A new genus and
species of Gastroplitinae from the Mzinene Formation (Albian). (Published
December 1978.) oh ae +
KENNEDY, W. J. & KLINGER, H. C.
Cretaceous faunas from Zululand and Natal, South Africa. The ammonite super-
family Haplocerataceae Zittel, 1884. (Published February 1979.)
KENSLEY, B.
The South African Museum’s Meiring Naude cruises. Part 8. Isopoda Anthuridea.
(Published October 1978.) 42 oe Shs er
KLINGER, H. C. see KENNEDY, W. J.
KLINGER, H. C. & Lock, B. E.
Upper Cretaceous sediments from the Igoda River Mouth, East London, South
Africa. (Published December 1978.) Bes ap oe ae
MILLARD, N. A. H.
Type specimens of Hydroida aun uraae in the South African Museum. oe
lished February 1979.) 2 .
Rau, R. E.
Additions to the revised list of preserved material of the extinct Cape Colony
quagga and notes on the relationship and distribution of southern plains
zebras. (Published December 1978.)
Page
151
47
175
123
57
85
71
133
27
NEW GENERIC NAMES PROPOSED IN THIS VOLUME
Alopecoceras Kennedy & Klinger, 1978
Cryptopenaeus de Freitas, 1979
‘OLUME 77 sail 1 OCTOBER 1978 ISSN 0303-2515
ae RY
DEC 27 1978
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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FiscHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
Fiscuer, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634. ‘
Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.
Koun, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
TurELe, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
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(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 77 Band
October 1978 Oktober
Part 1 Deel
THE SOUTH AFRICAN MUSEUM’S
MEIRING NAUDE CRUISES
PART 8
ISOPODA ANTHURIDEA
By
BRIAN KENSLEY
Cape Town Kaapstad
The ANNALS OF THE SOUTH AFRICAN MUSEUM
are issued in parts at irregular intervals as material
becomes available
Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000
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THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES
PART 8
ISOPODA ANTHURIDEA
By
Brian Kensley
South African Museum, Cape Town
(With 13 figures)
LMS. accepted 21 March 1978]
ABSTRACT
Ten species of anthurid and paranthurid isopods are recorded from deep water off Natal.
One is described as a new genus, viz. Natalanthura foveolata. A further five species are described
as new, viz. Apanthura insignifica, Neohyssura skolops, Colanthura uncinata, Leptanthura
minuta, and Leptanthura natalensis.
CONTENTS
PAGE
Introduction . . 3 ; : ; ; 1
Species list . : ; : : : : 1
Systematic discussion . ‘ ; ; : 2
Acknowledgements. : P : ae >,
References. : ‘ : 4 , mete ee
INTRODUCTION
The anthuridean isopods were excluded from the report on the isopods
of the 1975 and 1976 Meiring Naude cruises (Kensley 1977) either for lack of
specimens of some species or because of taxonomic uncertainty. With the 1977
cruise more anthurids were collected and some of the taxonomic problems
have been resolved. The anthurids of all three cruises are presented here.
Station data from the first two cruises may be obtained from Louw (1977),
and the South African Museum’s unpublished reports and station lists for the
1977 cruise.
SPECIES LIST
SM Station 3¢ 22 ~—so ovig. 92.—s jus
Family Anthuridae
Anthelura remipes Barnard ; ; : 123 os l — —
129 — — —
Apanthura insignifica sp. nov. . ; : 123 —- 2 — —
129 | 3 oe --
Mesanthura catenula (Stimpson) . 7 86 — 1 — l
Natalanthura foveolata gen. et sp. nov. . 86 = 6 os -—
123 — 1 — -
129 —_ _— l _
Neohyssura skolops sp. nov. : : : 129 — 1 — --
Ann. S. Afr. Mus. T7 (1), 1978: 1-25, 13 figs.
2 ANNALS OF THE SOUTH AFRICAN MUSEUM
SM Station oo cole) ovig. 22 juvs
Family Paranthuridae
Colanthura uncinata sp.nov. . ; : 86
103
Leptanthura minuta sp.nov. . : e 78
86
129
Leptanthura natalensis sp. nov. . : ‘ 53
Ww
9
12
1
|
Paranthura punctata (Stimpson) , 2 103
Pseudanthura tenuis Kensley ‘ é 53
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SYSTEMATIC DISCUSSION
Family Anthuridae
Apanthura insignifica sp. nov.
Figs 1-2
Description
Female
Integument thin. Body with following proportions: C<1=2=3<4=5>6>7.
Pereonite 7 half length of 6. Pereonites 4 to 7 with faint anterior constriction,
pereonite 4 with single, shallow, circular dorsal pit, pereonite 5 with transversely
elongate shallow pit, pereonite 6 with two small, shallow pits. Dorsolateral
grooves present. Pleonites | to 5 completely separate, pleonite 5 slightly longer
than preceding pleonites, pleonite 6 with middorsal incision in posterior margin.
Telson distally broadly rounded with several setae, widest at about midlength,
with low weakly developed middorsal ridge; two large statocysts present at
telsonic base. Cephalon with rostrum projecting beyond anterolateral corners;
eyes lacking.
Antennular peduncle 4-segmented, basal segment equal to second and
third segments together; fourth segment very short, flagellum of two articles
bearing three terminal aesthetascs.
Antennal peduncle 5-segmented, two basal segments fairly broad; flagellum
of four articles.
Mandibular palp 3-segmented, first and second segments subequal, third
segment shorter, with four distal serrate spines; incisor of three teeth, lacinia
plate with five teeth; molar process well developed, distally narrowly rounded.
Maxilla elongate, with one strong tooth and five smaller spines distally;
inner ramus tipped with single seta.
Lower lip complex tipped with single terminal hook-like lobe on each side.
THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 3
B
G
Fig. 1. Apanthura insignifica. A. Holotype in dorsal view. B. Antennule. C. Antenna.
D. Mandible. E. Maxilla. F. Lower lip complex. G. Maxilliped.
4 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 2. Apanthura insignifica. A. Pereonite 7 and pleon in lateral view. B. Pereopod 1.
C. Pereopod 2. D. Pereopod 7. E. Pleopod 1. F. Telson. G. Uropod.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 5
Maxilliped 5-segmented, terminal segment rectangular with several setae
on mediodistal corner.
Pereopod | unguis half length of dactylus, with small spine at base; pro-
podus broad, palm concave with strong, rounded lobe at about midpoint;
carpus triangular, distally narrowly rounded.
Pereopods 2 to 7 similar, with carpi barely underriding propodi, and
becoming slightly more elongate posteriorly.
Pleopod | exopod operculate, about three times wider and slightly longer
than the endopod, bearing numerous distal plumose setae; endopod with three
distal plumose setae; basis with three retinaculae.
Uropodal exopod folding over telson, oval in outline, reaching to end of
basis, bearing several sparsely plumose setae; endopod oval, fringed with
simple setae.
Submale
Antennular flagellum of about twelve articles extending posteriorly to
second pereonite, lacking whorls of aesthetascs.
Pereopod | as in female.
Stylet on pleopod 2 not yet detached from endopod.
Material
Holotype SAM-A15646 92° TL 5,9 mm SM 123 30°33’S 30°48’E 690 m
Paratype SAM-A15646 12 TL 5,2 mm SM 123
Paratypes SAM-A15647 1 sub. ¢ TL 5,6 mm 12 TL 4,5 mm SM 129
30°54’S 30°31’E 850 m
Paratypes USNM 170542 2 9° TL 5,6 mm 5,4 mm SM 129
Remarks
Of the twelve species of Apanthura described, only A. coppingeri Barnard
from the Dundas Straits (northern Australia) and A. africana Barnard from
South Africa lack eyes. The rhomboid-shaped telson, and the slender uropodal
endopod of A. coppingeri easily distinguish this species from A. insignifica.
A. africana, known from Saldanha Bay to Still Bay, is a much larger species
(up to 20 mm) recorded from shallow water. A. africana can further be separated
from A. insignifica by the maxilliped, which possesses a small terminal segment,
and an apically acute telson.
Etymology
The specific name refers to the small size of this species.
Natalanthura gen. nov.
Diagnosis
Integument heavily indurate and very obviously pitted. Mandible with
spike-shaped molar on right side, molar reduced to a spine on left side.
6 ANNALS OF THE SOUTH AFRICAN MUSEUM
Maxilliped 5-segmented with well developed endite. Carpus of pereopods
4 to 7 slightly underriding propodus. Pleopod | operculate. Pleonites 1 to 3
subequal, separate; pleonites 4 and 5 fused; pleonite 6 indistinguishably fused
with telson. Paired statocysts difficult to detect but probably present beneath
two pits at telsonic base.
Type species
Natalanthura foveolata
Etymology
The generic name Natalanthura is derived from the South African province
of Natal, in whose waters the species was caught, plus the usual ‘anthura’
suffix. Gender: feminine.
Remarks
The unusual fusion of the pleonites is not encountered in any of the
described genera of anthurids. The mandibular structure would also seem to
be unique, although reminiscent of Panathura. These two features demand
the erection of a new genus.
Natalanthura foveolata gen. et sp. nov.
Figs 3-4
Description
Female
Integument indurate and obviously pitted. Anterolateral corners of
cephalon extending slightly beyond rostrum. Eyes feebly pigmented, of three
or four ocelli. Pereonites with strong dorsolateral ridges and two large dorsal pits
per segment in addition to numerous smaller pits. Lateral walls of pereonites
pitted. Pereonites 1 to 4 increasing in width and length, pereonite 4 widest
and longest; posterior three pereonites decreasing in length, seventh half
length of sixth. Pereonites 3 to 5 with oblique dorsal grooves converging
anteriorly. Pleon with anterior three pleonites free and distinct; pleonite 4
and 5 fused, line of fusion marked by row of pits; pleonite 6 completely fused
with telson. Latter longer than pleonites 1 to 5 combined, distal margin evenly
convex, dentate, with strong mediodorsal rounded ridge.
Antennule with broadly curved basal segment plus six additional segments;
fourth segment short, terminal segment bearing setae and three or four
aesthetascs.
Antenna with 4-segmented peduncle; broad and relatively elongate basal
segment with distal triangular part folded over antennule; flagellum of six
articles.
Mandible with distal incisor tridentate, quite strongly sclerotized; lacinia
plate with row of twelve short teeth; right mandible with digitiform molar
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 7
process bearing row of small denticles and striations; molar process reduced
to small spine in left mandible; palp 3-segmented, middle segment longest and
widest, terminal segment bearing three stout setae.
Maxilla slender and elongate, with four terminal teeth.
Maxilliped 5-segmented, terminal segment rounded, bearing four setae;
endite elongate-lanceolate, apically acute, with single seta on median margin,
reaching just beond proximal margin of third segment.
See
F
Fig. 3. Natalanthura foveolata. A. Holotype in dorsal view. B. Antennule. C. Antenna.
D. Maxilla. E. Right mandible. F. Maxilliped.
8 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 4. Natalanthura foveolata. A. Pereopod 1. B. Pereopod 7. C. Pereonites 4 and 5 in lateral
view. D. Pleopod 1. E. Uropod.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 9
Pereopods | and 2 similar, unguis one-third length of dactylus, with single
spine at base; propodus with single distal spine and numerous setules on
palmar margin. Posterior pereopods with unguis one-third length of dactylus,
with short spine at base, propodus with strong distal spine and four or five
short, slender spines on ventral margin, and single serrate seta on upper distal
margin; carpus about half length of, and slightly underriding, propodus.
Brood pouch formed by four pairs of oostegites.
Pleopod | operculate, rami of equal length, endopod half width of exopod,
with four distal plumose setae; exopod with numerous plumose setae distally,
and with outer surface bearing scales; basis bearing two coupling hooks.
Uropodal endopod subequal to basis in length, entire margin strongly
dentate; exopod crescentic and curved, margin strongly dentate; basis triangular
in cross-section.
Material
Holotype SAM-A15648 2 TL 5,8 mm SM 86 27°59’S 32°40’E 550 m
Paratypes SAM-A15648 2 292 TL 3,5 mm 5,4 mm SM 86
Paratypes USNM-170543 3 92 TL 4,3 mm 4,3 mm 4,6 mm SM 86
Paratype SAM-A15649 1 2 TL3,4mm SM 123 30°33’S 30°48’E 690 m
Paratype SAM-A15650 | ovig. 2 TL 6,9 mm SM 129 30°53’S 30°31’E 850 m
Etymology
The specific name refers to the pitted nature of the integument.
Neohyssura skolops sp. nov.
Figs 5-6
Description
Female
Integument thin, only uropods and telson slightly indurate. Body pro-
portions: C<1<2=3<4=5=6>7. Dorsolateral grooves present on pereonites
but very difficult to see; dorsal pits absent. Anterior five pleonites separate,
equal in length and breadth; pleonite 6 fused with telson; latter with broad,
rounded base, tapering rapidly into cylindrical spike-like structure tipped with
setae; no statocysts visible. Eyes absent. Anterolateral corner of cephalon
extending beyond low rostrum.
Antennular peduncle 4-segmented, two basal segments broad, fourth
segment set obliquely into third, short; flagellum of five articles with few setae
and single apical aesthetasc.
Antennal peduncle 5-segmented, second and fifth segments subequal,
longer than third and fourth segments; second segment grooved to receive
antennule; flagellum of seven articles.
Mandibular palp 3-segmented, first and third segments subequal, about
one-third length of second segment; incisor of three teeth, lacinia plate bearing
five blunt teeth; molar process blunt, tapering.
10 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 5. Neohyssura skolops. A. Pereonite 7 and pleon in lateral view. B. Telson and uropods
in dorsal view. C. Antenna. D. Antennule. E. Maxilla. F. Maxilliped.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 11
Fig. 6. Neohyssura skolops. A. Pereopod 1. B. Mandible. C. Pleopod 1. D. Lower lip
complex. E. Pereopod 7.
12 ANNALS OF THE SOUTH AFRICAN MUSEUM
Maxilla with one strong, outer spine and five smaller spines distally.
Lobes of lower lip distally bearing two short acute processes.
Maxilliped 7-segmented, terminal segment short, bearing four setae;
thin-walled endite tapering, with single terminal seta.
Pereopods | to 3 similar, subchelate, pereopod 2 more robust than | or 3;
unguis one-third length of dactylus, propodus proximally broad, palm convex
between three short sensory spines, with few scattered setae on inner face;
carpus triangular, produced distally well beyond base of propodus to meet
dactylus. Pereopods 4 to 7 slender, unguis one-sixth length of dactylus; propodus
with strong posterodistal spine; carpus short, underriding propodus, with
strong sensory spine on posterior margin.
Uropods somewhat indurate, both rami reaching telsonic apex; exopod
situated dorsal to telson, with four strong teeth on medial margin, apically
rounded; basis with strong mediodistal spine visible in ventral view.
Material
Holotype SAM-A15651 2 TL 5,8 mm SM 129 30°53’S 30°31’E 850m
Remarks
Barnard (1925) noted that ‘Hyssura’ spinicauda Walker, 1901, did not
fit the generic diagnosis of Hyssura but he did not decide on the generic status..
Amar (1952), on examining a specimen from Corsica, erected the new genus
Neohyssura which he defined as follows: close to Hyssura but the maxillipeds
with seven segments, lacking endite, seventh pereopod present, pereopods 4
to 7 with triangular carpus underriding propodus.
The present specimen agrees with all except one of these characters. The
maxilliped does have a thin-walled endite (easily overlooked), but for the rest
is very similar to that of N. spinicauda. The shape of the telson also differs,
being narrowly triangular and bearing lateral spines in N. spinicauda but
narrowly spike-like in N. skolops. (See Walker 1901, pl. 27, figs 7-11; Monod
1925, pl. 44.)
Etymology
The specific name ‘skolops’ is taken from the Greek word meaning a sharp
object such as spike, and refers to the shape of the telson.
Family Paranthuridae
Colanthura uncinata sp. nov.
Figs 7-8
Description
Male
Entire integument covered with small imbricate scales. Rostrum extending
slightly beyond anterolateral corners of cephalon. Eyes ovate, lateral, consisting
of fourteen ocelli. Cephalon two-thirds length of pereonite 1. Pereonites | to 5
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 13
Ty
W/ wf f;
Y
Tf
Fig. 7. Colanthura uncinata. A. Holotype in dorsal view. B. Maxilla. C. Maxilliped.
D. Antenna. E. Pereopod 1 with spine further enlarged. F. Antennule 2. G. Antennule 3.
14 ANNALS OF THE SOUTH A8RICAN MUSEUM
F
Fig. 8. Colanthura uncinata. A. Pleopod 1. B. Pleopod 2 ¢ with apex of stylet further enlarged.
C. Pereopod 2. D. Pereopod 6. E. Telson. F. Uropodal exopod. G. Uropodal endopod j
and basis.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 15
increasing in width, but subequal in length; posterior margins of pereonites
1 and 2 bilobed. Pereonite 6 two-thirds length of pereonite 5; pereonite 7 very
short, subequal to anterior pleonites in length and width, lacking pereopods.
Ventral surface of pereonites formed by two longitudinal, low, rounded ridges,
conspicuously scaled. Pleon narrower than pereon; pleonites 1 to 5 subequal,
pleonite 6 posteriorly bilobed, almost twice length of preceding pleonite.
Telson equal to pleonites | to 6 in length, distally evenly convex; dorsally with
low, rounded median area widest proximally.
Antennular peduncle 4-segmented, basal segment largest; flagellum of six
articles, five proximal articles broad, bearing dense aesthetascs, terminal
article narrow, bearing setae.
Antennal peduncle 5-segmented, second segment largest and broadest;
flagellum reduced to setiferous terminal article about one-third width and half
length of penultimate segment.
Mandible reduced to blunt lobe lacking palp.
Maxilla very slender, acute, with about ten faint distal serrations.
Maxilliped elongate, consisting of a single segment, distally narrowed,
with about seven terminal setae.
Pereopod 1 dactylus strongly curved, unguis short; propodus broadly
oval, palm gently sinuous, with triangular proximal process and row of fourteen
fringed spines set back from the triangular process.
Pereopod 2 dactylus gently curved, unguis short; propodus proximally
broad, distally narrowed, palm with short spines.
Pereopods 3 to 7 similar, unguis of dactylus relatively longer than in
pereopods | and 2; propodus with three spines and numerous fine setules on
ventral margin, plus two distal fringed spines; carpus not underriding propodus,
bearing two ventral spines.
Pleopod 1 exopod broad and operculate, with scales on external face;
endopod as long as, but one-third width of, exopod.
Pleopod 2 exopod and endopod subequal in length and width, each with
few distal plumose setae; stylet of endopod extending well beyond apices of
rami, distally rounded with strong subapical hook.
Uropodal endopod just extending to telsonic apex, oval; exopod narrowly
ovate, apically narrowly rounded.
Ovigerous female
Antennule consisting of five segments, penultimate segment very short.
Pereopod | similar to that of male, but with spine row of six (rather than
fourteen) spines. Brood pouch formed by four pairs of oostegites, containing
four eggs or four larvae.
Material
Holotype SAM-A15652 3 TL 3,9 mm SM 103 28°31’S 32°34’E 680m
Allotype SAM-A15652 ovig. 2 TL 4,5 mm SM 103
16 ANNALS OF THE SOUTH AFRICAN MUSEUM
Paratypes USNM 170544 2 3g 1 ovig. 2 SM 103
10 gd 2 ovig. 92 12 immature SM 103
Paratypes SAM-A15653 2 gg 1 ovig. 29 SM 86 27°59’S 32°40’E 550 m
1g lovig.2 9 immature SM 86
Remarks
Four species of Colanthura have been described. These are C. tenuis
Richardson, 1902, from Bermuda; C. squamosissima Menzies, 1951, from
California; C. nigra Nunomura, 1975, from Japan, and C. caeca Mezhov, 1976,
from Russia. The present material most closely resembles the Californian
species, this similarity being most marked in the character of the integument,
both species being almost covered with imbricate scales. Several differences
between these two species are apparent. These include pleonites | to 5, which
in C. uncinata are subequal, while in C. squamosissima pleonite 5 is four times
the length of the preceding segment; the telson, which is more obviously tapered
in C. uncinata; and the apex of the stylet of pleopod 2 3, which has a swollen
apical lobe in Menzies’s species, but is hooked in C. uncinata. The stylet of
pleopod 2 3 in C. caeca is also apically hooked, but possesses three smaller
supplementary apical spines (Mezhov 1976, fig. 3). The shape of the uropodal
exopod and telson, and the propodal spination of the pereopods of the Russian
species differs from C. uncinata.
Etymology
The specific name is taken from the hooked stylet of pleopod 2 2.
Leptanthura minuta sp. nov.
Figs 9-10
Description
Male
Body slender, elongate, not indurate. No dorsal pits or dorsolateral
grooves present. Cephalon subequal to pereonite | in length; eyes lacking.
Body proportions: C=1<2=3=4<5>6>7. Pleon equal to pereonites 6 and 7
together in length; all pleonites distinct, pleonites | and 5 slightly longer than
2 to 4; pleonite 6 with convex distal margin. Telson elongate-oval in outline,
distally evenly rounded, with four median setae; ventrally concave; dorsally
with gently raised central area; prominent statocyst present with single medio-
dorsal pore.
Antennule with 4-segmented peduncle, basal segment largest, fourth
segment very narrow; flagellum of eight articles each with dense whorl of
aesthetascs.
Antenna with 5-segmented peduncle, second segment longest, with disto-
dorsal triangular part folding over basal antennular segment; flagellum of
four articles.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 17
Mandible narrowly triangular, apically acute; 3-segmented palp with
middle segment two and a half times length of basal segment, terminal segment
short and curved, with single terminal seta.
Maxilla lanceolate, apically acute, with about twelve faint distal serrations.
Maxilliped 3-segmented, with few distal setae; second segment six times
longer than wide.
Pereopod | dactylus fairly stout, unguis one-third length of rest of segment;
propodus proximally broad, palm straight with low triangular proximal
D
Fig. 9. Leptanthura minuta. A. Holotype in dorsal view. B. Mandible. C. Maxilla.
D. Maxilliped. E. Antennule ¢. FF. Antennule 2. G. Antenna. H. Pereopod | 9.
I. Pereopod 1 ¢.
18 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 10. Leptanthura minuta. A. Pleopod 1. B. Pleopod 2g. C. Pereopod 2. D. Pereopod 7.
E. Telson and uropod.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 19
tooth, armed with six short serrate spines and numerous simple setae; carpus
triangular, small, with two spines and four setae on ventral margin.
Pereopod 2 slightly less robust than pereopod 1, propodus with six spines
on palm, lacking dense setae found on pereopod 1; carpus with single ventral
spine. Pereopods 4 to 7 similar, slender; propodi with three serrate spines on
ventral margin; carpus short, triangular, underriding propodus, with single
ventral spine.
Pleopod | basis with four retinaculae on median margin, exopod operculate,
two and a half times longer than broad with about ten to twelve distal elongate
plumose setae; endopod one-third width and two-thirds length of exopod, with
eight distal plumose setae.
Pleopod 2 with three or four retinaculae on basis; rami subequal in length;
exopod with transverse articulation at midlength and about eight distal plumose
setae; endopod with four distal plumose setae, stylet on median margin extend-
ing well beyond apex of ramus, apically blunt.
Uropodal exopod extending to base of endopod, triangular-oval, median
margin crenulate and setose; basis with single elongate plumose seta at medio-
distal angle; endopod with several distal setae, apically rounded.
Female
Antennule with 4-segmented peduncle, basal segment longest and broadest,
fourth segment very narrow; flagellum reduced, consisting of three articles.
Pereopod | very similar to pereopod 1 3, but lacking numerous simple
setae on palm.
Brood pouch of four pairs of oostegites, anterior pair smallest.
Material
Holotype SAM-A15654 3 TL 4,6 mm SM 86 27°59’S 32°40’E 550 m
Allotype SAM-A15654 ovig. 9 TL 4,5 mm SM 86
Paratypes SAM-A15654 1 gl ovig.2 12 SM 86
Paratypes USNM 170545 1g 1 ovig.2 12 SM 86
SAM-A15655_ 1 ovig. 2 SM 7S. 2731S. 32°S0'E
750 m
SAM-A15656 2g3¢ 12 SM 129 30°53’S 30°31’E
850 m
Remarks
The present small blind species with its characteristic elongate-oval telson
and crenulate uropodal exopod bears no close resemblance to any of the
approximately eighteen described species of Leptanthura. Certainly it is very
different from the described South African species, viz. L. agulhasensis,
L. laevigata, and L. urospinosa. The shape of the telson of L. minuta to some
extent resembles that of L. antarctica Kussakin, 1967, but the shape of the
uropodal endopod and exopod, the antennular peduncle, and the first pereopod
easily separate these species.
20 ANNALS OF THE SOUTH AFRICAN MUSEUM
Etymology
The specific name derives from the small size of the adult animals.
Leptanthura natalensis sp. nov.
Figs 11-12
Description
Male
Body very slender and elongate. No dorsal pits. No dorsolateral grooves
present, but dorsolateral ridge developed on cephalon and pereonites | and 2.
Cephalon lacking eyes. Body proportions: 1<2<3<4>5>6>7. Pereonite 5
in small males (6-7 mm) with two rounded posteroventral lobes; holotype
male (20,6 mm) with similar lobes on pereonite 7. Pleonites free. Telson distally
broadly rounded, not indurated, dorsally slightly concave, with single, large
proximal statocyst.
Antennule with 4-segmented peduncle, basal segment largest, fourth seg-
ment short; flagellum of thirteen articles.
Antenna 8- or 9-segmented, of which three distal segments are probably
flagellum; second segment longest.
Mandible acutely triangular, palp 3-segmented, basal segment two-thirds
length of middle segment, latter with single elongate distal seta, terminal seg-
ment short and curved.
Maxilla slender, with twelve serrations and three barbs distally.
Maxilliped 4-segmented, second segment four and a half times longer
than wide, with short distal lobe bearing two setae at mediodistal angle; two
distal segments narrowing, bearing few setae.
Pereopod | dactylus with very short unguis; palm of propodus almost
straight, with low proximal triangular lobe, and row of fourteen short sensory
spines, several elongate setae, and dense border of short simple setae.
Pereopod 2 propodus narrower than that of pereopod 1, with nine or ten
sensory spines on palm, lacking dense border of simple setae; carpus small,
triangular, with two spines on ventral margin.
Pereopod 7 with short carpus bearing two spines and underriding propodus;
latter with three spines on ventral margin.
Pleopod | operculate, not indurated; endopod considerably shorter than
exopod, both rami bearing distal plumose setae; basis bearing about six
retinaculae on medial margin.
Pleopod 2 endopod shorter than exopod, with apically blunt stylet on
medial margin; exopod with distinct transverse articulation at about midlength,
both rami with distal plumose setae; basis armed with five or six retinaculae
on medial margin.
Uropodal basis triangular in cross-section, longer than endopod; latter
narrower than basis, with tuft of apical setae; exopod broadly oval, with few
marginal setae, exopods overlapping telson dorsally.
ae
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 21
Fig. 11. Leptanthura natalensis. A. Holotype in lateral view. B. Pereonite 5 of 6,8 mm ¢.
C. Antennule g. D. Antenna. E. Mandible. F. Maxilla. G. Maxilliped.
22 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 12. Leptanthura natalensis. A. Pleopod 1. B. Pleopod 2 3. C. Pereopod 1 3, with palm
enlarged (short, simple setae omitted). D. Pereopod 2. E. Pereopod 7. F. Telson.
G. Uropodal exopod. H. Uropodal basis and endopod.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 23
Material
Holotype SAM-A15657_ 3 TL 20,6 mm SM 60 27°09’S 32°58’E 800 m
Paratypes SAM-A15658 2 gg TL6,8 mm 6,7 mm 1 juv. 4,8 mm
SM 53 26°51’S 33°12’E 720m
Paratypes SAM-A15659 2 gg TL 6,7 mm 7,5 mm SM 61 27°10’S
32°58’E 820 m
Paratypes USNM 170546 1 ¢ TL 8,0mm 1 2 5,7mm SM 117 30°17’S
31°10’E 820 m
SAM-A15660 1 ¢ 9,9 mm 1 juv. SM 123 30°33’S 30°48’E
690 m
SAM-A15661 7 922 complete but damaged SM 129 30°53’S
30°31’E 850m
Remarks
The general structure of the pereon and pleon, as well as the mandible,
maxilliped, antennae, and pereopods of the present material, agrees with
Leptanthura, and quite closely resembles L. lJaevigata and L. glacialis. The
broadly rounded telson is similar to that of L. /aevigata (Stimpson) (Barnard
1925), but L. natalensis lacks the characteristic notch in the uropodal exopod,
and is a more attenuated species both in body shape and pereopodal propor-
tions. The terminal maxillipedal segment is longer than that of L. laevigata.
L. glacialis Hodgson (Kussakin 1967) has a uropodal exopod similar to
L. natalensis but possesses an apically pointed telson, and shorter and broader
antennal segments. L. glacialis appears to have a 3-segmented maxilliped,
rather than the 4-segmented structure of L. natalensis.
Etymology
The specific name is taken from the province, Natal, from whose waters
the species was taken.
Pseudanthura tenuis Kensley
Fig. 13
Pseudanthura tenuis Kensley, 1978: 222, figs 1-2.
Description
Male
Antennular peduncle 4-segmented; segments broad, basal segment equal
in length to three distal segments combined; basal flagellar article broad and
almost as long as remaining seven flagellar articles together; six distal articles
each with pair of aesthetascs.
Pleopod 2 stylet of endopod extending well beyond apex of ramus, straight,
apically rounded.
24 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 13. Pseudanthura tenuis. A. Pleopod 2 3. B. Antennule ¢.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 25
Material
Le TE9 9mm SM53 26°5S1'S 33°1E 720m
692 TL6,0-25,33mm SM 103 28°31’S 32°34’E 680m
be TE 20,7 mm SM 129 30°53’S 30°31’E 850m
Remarks
P. tenuis Kensley was described from seven females from localities very
close to that of the present material. Now that a single male is available, the
two main sex-linked characters are described and figured.
ACKNOWLEDGEMENTS
In addition to the people and institutions thanked in the introductory
paper in this series, I should like to thank Mr R. Downes of the South African
Museum for sorting the samples of the 1977 cruise. My sincere thanks are
also due to Dr T. E. Bowman of the Smithsonian Institution for useful dis-
cussions and for critically reading the manuscript.
REFERENCES
Amar, R. 1952. Isopodes marins du littoral Corse. Bull. Soc. zool. Fr. 77: 349-355.
BARNARD, K. H. 1925. A revision of the family Anthuridae (Crustacea Isopoda) with remarks
on certain morphological peculiarities. J. Linn. Soc. 36: 109-160.
KENSLEY, B. 1977. The South African Museum’s Meiring Naude cruises. Part 7. Marine
Isopoda. Ann. S. Afr. Mus. 74: 125-157.
KENSLEY, B. F. 1978. Two new species of the genus Pseudanthura Richardson (Crustacea,
Isopoda, Anthuridea). Proc. biol. Soc. Wash. 91: 222-233.
KussAKIN, O. G. 1967. Fauna of Isopoda and Tanaidacea in the coast zones of the Antarctic
and Subantarctic waters. Biol. Rep. Sov. Antarct. Exped. (1955-58) 3: 220-380.
Louw, E. 1977. The South African Museum’s Meiring Naude cruises. Part 1. Station data
1975, 1976. Ann. S. Afr. Mus. 72: 147-159.
Menzies, R. J. 1951. New marine isopods, chiefly from northern California, with notes on
related forms. Proc. U.S. natn. Mus. 101: 105-156.
MEzHov, B. V. 1976. New species of Anthuroidea from the upper part of the sublittoral zone
of the Middle Kurile Islands. Biol. Sea 5: 19-27.
Monon, T. 1925. Tanaidacés et Isopodes aquatiques de l’Afrique Occidentale et Septentrionale.
Bull. Soc. Sci. nat. Maroc 5: 233-247.
NunomurA, N. 1975. Marine Isopoda from the rocky shore of Osaka Bay, Middle Japan.
Bull Osaka Mus. nat. Hist. 29: 15-35.
RICHARDSON, H. 1902. The marine and terrestrial isopods of the Bermudas, with descriptions
of new genera and species. Trans. Conn. Acad. Arts Sci. 11: 277-310.
WALKER, A. O. 1901. Contributions to the Malacostracan fauna of the Mediterranean.
J. Linn. Soc. 28: 290-307.
6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.
An author's name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nickles, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.
In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes shou!d be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and date.
7. SPECIAL HOUSE RULES
Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. *... the Figure depicting C. namacolus .. .’; *. . . in C. namacolus (Fig. 10)...’
(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene
(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian
Punctuation should be loose, omitting all not strictly necessary
Reference to the author should be expressed in the third person
Roman numerals should be converted to arabic, except when forming part of the title of a
book or article, such as
“Revision of the Crustacea. Part VIII. The Amphipoda.’
Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.
Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.
BRIAN KENSLEY
THE SOUTH AFRICAN MUSEUM’S
MEIRING NAUDE CRUISES
PART 8
ISOPODA ANTHURIDEA
a AL. Caplou—
OLUME 77 PART 2 DECEMBER 1978 MUS. COMP. ZOOS
LIBRARY SN 0303-2515
FEB 15 07)
HARV far
UNIVERSITET?
ANNALS
OF THE SOUTH AFRICAN
MUSEUM
CAPE TOWN
1. MATERIAL should be original and not published elsewhere, in whole or in part.
2. LAYOUT should be as follows:
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INSTRUCTIONS TO AUTHORS
Centred masthead to consist of
Title: informative but concise, without abbreviations and not including the names of new genera or species
Author’s(s’) name(s)
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Abstract of not more than 200 words, intelligible to the reader without reference to the text
Table of contents giving hierarchy of headings and subheadings
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Summary, if paper is lengthy
Acknowledgements
References
Abbreviations, where these are numerous
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5. REFERENCES cited in text and synonymies should all be included in the list at the end of
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(a) Author’s name and year of publication given in text, e.g.:
(b)
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within each name, with suffixes a, 5, etc. to the year for more than one paper by the same
author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a).
For books give title in italics, edition, volume number, place of publication, publisher.
For journal article give title of article, title of journal in italics (abbreviated according to the World list o,
scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part
number (only if independently paged) in parentheses, pagination (first and last pages of article).
Examples (note capitalization and punctuation)
BULLouGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FiscHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FiscHErR, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann, Mag. nat. Hist. (13) 2: 309-320.
Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
TurELe, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 77 Band
December 1978 Desember
Part 2 Deel
BODITIONS TO THE REVISED LIST OF PRESERVED
maALERIAL OF THE EXTINCT CAPE COLONY
QUAGGA AND NOTES ON THE RELATIONSHIP AND
DISTRIBUTION OF SOUTHERN PLAINS ZEBRAS
By
cE; EY RAY
Cape Town Kaapstad
The ANNALS OF THE SOUTH AFRICAN MUSEUM
are issued in parts at irregular intervals as material
becomes available
Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000
Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
word uitgegee in dele op ongereelde tye na beskikbaarheid
van stof
Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 8000
OUT OF PRINT/UIT DRUK
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6(1, t.-p.i.), 711-4), 8, 911-2, 7), 10(1-3),
11(1-2, 5, 7, t.—p.i.), 15(4-5), 24(2), 27, 31(1-3), 32(5), 33
Copyright enquiries to the South African Museum
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ISBN 0 908407 57 2
Printed in South Africa by In Suid-Afrika gedruk deur
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap
ADDITIONS TO THE REVISED LIST OF PRESERVED MATERIAL
OF THE EXTINCT CAPE COLONY QUAGGA
AND NOTES ON THE RELATIONSHIP AND DISTRIBUTION OF
SOUTHERN PLAINS ZEBRAS
By
R. E. RAu
South African Museum, Cape Town
(With 9 figures)
LMS. accepted 7 September 1978]
ABSTRACT
Previously unpublished photographs and data of two mounted skins of the extinct
quagga (Equus quagga quagga) and a further relic are added to the list of preserved material.
Photographs of the destroyed KGnigsberg quagga are reproduced.
Examination of plains zebra (Equus ‘burchelli’) skins from southern Africa and comparison
with the true Burchell’s zebra (Equus ‘burchelli burchelli’) and the quagga (Equus quagga
quagga) suggest that all belong to one species (Equus quagga) and are forms which are dominant
in certain areas within the continuous distribution, rather than subspecies. In addition, the true
Burchell’s zebra is considered to be still extant in Zululand and possibly Swaziland, if the
generally accepted but vague range of that ‘subspecies’ is ignored.
CONTENTS
PAGE
Introduction . ; : : : 2 : eT
Additional material : ; : : : eaves
Notes on Burchell’s zebra and quagga : Ay si
Reintroductions of zebras. . : : Ae: |
Acknowledgements . : d ‘ : : . 44
References . : : , : : : = . 44
INTRODUCTION
Since the publication of the revised list of preserved material of the quagga
(Equus quagga quagga) a photograph of and additional data on the quagga at
the Zoological Museum, State University at Kazan, Soviet Union, have now
been obtained. In addition, a mounted quagga has been discovered at the
Muséum d’Histoire Naturelle Lyon, France, and a mounted foot at the Royal
Albert Memorial Museum at Exeter, England. These new data are presented in
accordance with the method followed by Rau (1974). The photographs included
in the description of the Kénigsberg quagga (Hilzheimer 1930), which was
destroyed at the end of the Second World War, are reproduced to complete the
illustrations of mounted skins which existed into this century.
27
Ann. S. Afr. Mus. 77 (2), 1978: 27-45, 9 figs.
28 ANNALS OF THE SOUTH AFRICAN MUSEUM
In a monograph Antonius (1951), a world authority on zebras, included the
quagga and the Burchell’s zebras in the single species Equus quagga, as had been
done earlier (Pocock 1904; St Leger 1932). This view is supported by
Rzasnicki (1951), Mohr (1964) and Klingel (1969, 1972) amongst others.
There is, however, a second school of thought which divides them into two
species, Equus quagga and Equus burchelli (Ansell 1971). As the present study
has revealed that there is no definite division between the quagga and Burchell’s
zebras, the former view is supported.
ADDITIONAL MATERIAL
KAZAN— MOUNTED SKIN
Fig. 1
Catalogue number: 109
Sex: female
Locality: unknown
Date of acquisition: 1843
Remarks on acquisition: bought in Hamburg from a Mr Brandt by Professor
Eversman for the Zoological Museum of Kazan University
Fig. 1. Kazan quagga.
ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 29
History of mount: bought as mounted specimen in bad condition and remounted ;
a second remounting (‘restoring’) was performed in 1969 by M. A. Zaslavski
in Leningrad, U.S.S.R.
Description of striking features: face clearly striped; shadow-stripes on neck;
posterior part of feet (fetlock-hoof) dark; the photograph seems to show
faint striping on buttock
Measurements: head-body 2,36 m* (2,42 m)*
tail 0,54m (0,53 m)
ear 0,14m (0,17 m)
hind foot 0,56 m (0,54 m)
shoulder height 1,29 m_ (1,28 m)
State of preservation: appears to be good
Further material of same individual: none
Remarks: exhibited open in gallery
References: none known
LYON— MOUNTED SKIN
Fig. 2
Catalogue number: 6102
Sex: female
Locality: Kaffraria, South Africa
Date of acquisition: unknown
Remarks on acquisition: none
History of mount: appears to be the original mount
Description of striking features: face not clearly striped
Measurements: head-body 2,10 m
tail 0,40 m
ear 0,17 m
hind foot 0,50 m
shoulder height 1,15 m
State of preservation: fair
Further material of same individual: none
Remarks: exhibited; specimen not mentioned in previous literature; V. Eisen-
mann of the Muséum d’Histoire Naturelle, Paris, discovered this specimen
early in 1975 and kindly informed the author
References: M. Philippe, ‘La zoologie au Muséum d'Histoire Naturelle de Lyon’
(in preparation)
* Measurements received through two intermediaries are not identical.
30 ANNALS OF THE SOUTH AFRICAN MUSEUM
—_
Fig. 2. Lyon quagga.
EXETER— FOOT TROPHY
Fig. 3
Catalogue number: A758
Locality: Bontebok Flats, British Kaffraria, South Africa (between present
Cathcart and Queenstown)
Date of acquisition: after 1869 )
Remarks on acquisition: from the collection of W. S. M. D’Urban, first curator
of Exeter Museum
Measurements (hoof only): anterior height 6,0 cm
width 6,7 cm \
length 8,4 cm .
References: Howes & Bamber (1970)
ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 3]
Fig. 3. Exeter quagga foot; anterior and lateral views.
KONIGSBERG QUAGGA
Fig. 4
The K6nigsberg quagga was destroyed at the end of the Second World War
when Schloss Waldhof, about 20 km from KGnigsberg, where museum specimens
had been taken for safe-keeping, was shelled and completely destroyed by fire
(pers. comm. Prof. Dr O. Koehler (1972), director of the K6nigsberg Museum
until 1945).
The preserved material of Equus quagga quagga, including the above
specimens, now consists of:
skins—23, plus 1 mounted head and neck
complete skeletons—7
skulls—20, including 7 from complete skeletons; at least 3 of the mounted
skins appear to contain skulls
loose skeletal parts—2 sets of foot bones; upper and lower incisivae in
portion of jaws
other—dried connective tissue, flesh, etc., removed from skin; nasal
cartilages and palate removed from skull (in alcohol) (Rau 1974: 58)
NOTES ON BURCHELL’S ZEBRA AND QUAGGA
During February to October 1977 approximately 400 privately owned skins
of plains zebra Equus quagga (burchelli) and mountain zebra Equus zebra
hartmannae were processed at a Cape Town tannery and examined by the
author. These originated from many different localities in southern Africa.
Skins of 113 plains zebras were photographed, the initial 88 non-selectively and
32 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 4. Konigsberg quagga (destroyed), adapted from Hilzheimer (1930).
ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 35
the remaining 25 selectively. The proportion of the two species processed
during a two-month period (February and March) was 88 plains zebra:
62 mountain zebra.
A great number of ‘subspecies’ and even ‘species’ of plains zebra have been
described. In bestowing names to the many variations, the stripe pattern has
been the principal, and often the only, character considered (Ansell 1971). This
is certainly true in the case of the ‘subspecies’ burchelli.
The diagnostic characteristics of the true Burchell’s zebra Equus quagga
burchelli are usually given as follows: whitish, unstriped legs except occasionally
some traces of stripes on the hocks (Pocock (1897: 52) accepts some stripes on
the ‘knees’ as well); basic colour of dorsal parts white to ochre and brownish;
interspaces with shadow-stripes, which may be broad and intense, or less
conspicuous; body-stripes not meeting the ventral midline.
Equus quagga antiquorum is defined by the striping extending on to the legs
to at least the ‘knees’ and hocks, but sometimes to the pasterns.
Equus quagga burchelli (E. burchelli burchelli) is one of the few still recog-
nized as a valid subspecies. The accepted distributional range of this subspecies
included the Orange Free State, the north-eastern Cape Province, southern
Botswana and the south-western Transvaal. As zebras no longer occur in these
regions, other than reintroduced specimens, the true Burchell’s zebra is usually
considered to have been extinct since about 1910.
The locality of the type specimen on which Gray (1825) based the descrip-
tion of the subspecies is between Campbell and the Vaal River. The specimen
figured as Equus quagga burchelli by Antonius (1951, fig. 22) originated from
British Bechuanaland (Botswana) and lived at Hagenbeck’s Tierpark, Germany,
during 1907-9. However, most preserved specimens of this subspecies lack
locality data, as do those individuals that lived at menageries or zoos and are
known from illustrations and/or descriptions.
It appears, therefore, that the accepted range is based on old reports rather
than collected material, and that the true range is not known. Indeed, different
opinions as to the range of Equus quagga burchelli have been expressed. Matschie
(1894) has included Zululand while Shortridge (1934) and Eloff (1967) extend
the range to Great Namaqualand (South West Africa). The concept of the
range of a subspecies within the continuous distribution of a species is open to
question, especially in the case of fast-moving and migrating animals such as
the plains zebra. With regard to zebra distribution, Tegetmeier & Sutherland
(1895) wrote: ‘Disputing, therefore, as to whether two closely allied animals are
specifically or sub-specifically distinct is almost a waste of words. We know that
a species spread over a wide area will change according to the conditions of life,
until at last the two extremes are so diverse as to be regarded as distinct species,
but no one can say where one species ends and the other begins, for they merge
gradually into each other.’
The uncertainty about the former range of the subspecies Equus quagga
burchelli has been aggravated by the indiscriminate use of the names ‘quagga’
34 ANNALS OF THE SOUTH AFRICAN MUSEUM
and ‘Burchell’s zebra’, and by the difficulty in distinguishing between those two
forms as well as between the true Burchell’s zebra and its northern relatives.
Accurate assessment of old documents is, therefore, practically impossible.
Pocock (1897: 42) wrote 13 years before the alleged extinction of Equus quagga
burchelli: *. .. and on account of the general application of the term Burchelli
to any or all of its subspecies, we are in ignorance of the exact area that it now
occupies in S. Africa.’
If all the mounted zebra specimens preserved in museums had precise
locality data, there would be in all probability antiquorum-types from the
burchelli ‘range’ and vice versa. Since most older mounted zebras do not,
however, have conclusive locality data, it is accepted that apparent burchelli
specimens originate from the accepted burchelli range while antiquorum
patterned specimens are thought to be from further north. Since the plains zebra
has a continuous distribution, recent interferences by man excepted, within
which certain forms or variations predominate in certain areas, it follows that,
in intergradational areas between two forms, the one form will gradually
decrease in percentage of total numbers as the area of the next form is
approached.
Despite the fact that for most of the preserved or known plains zebra
specimens, which conform basically to the description of Equus quagga burchelli,
the place of origin is not known, they are accepted as belonging to that sub-
species, as is shown in the figures by Cabrera (1936) (Fig. 5) and Antonius (1951).
The subspecies Equus quagga antiquorum is presently considered to include
all the extant southern forms of the plains zebra, ranging from southern Angola
and (historical) Damaraland through Botswana and Transvaal to Zululand.
Consequently, individual variation within antiquorum is considerable (Fig. 6).
Eloff (1967) stated that the three subspecies antiquorum, chapmani and wahlbergi
occur together in Zululand. Antonius (1951) had observed that in Damara zebra
and especially in Wahlberg zebra burchelli-like individuals do occur.
However, such individuals originate from outside the presumed range of
the true Burchell’s zebra and are consequently not accepted as such. Neverthe-
less, some authors are cautious. Cabrera (1936: 94), while saying that the
subspecies ‘appears to be extinct’, also states, ‘in fact, I regard as typical
burchelli all members of the species having all four legs white from the elbow
and the stifle joint, except for the occasional presence of a few short markings
across the hocks, and with the thighs free of complete and well-defined dark
stripes behind the stifle stripe’. Dorst & Dandelot (1970) list the subspecies
burchelli as ‘probably extinct’.
Amongst the extremely variable skins (many with numerous small, light
dots in stripes and shadow-stripes) examined at the tannery, were several
specimens from a Zululand game farm with a surprising combination of light
basic colour and few and/or weak shadow-stripes and considerable stripe
reduction (Fig. 7A). It is generally believed that the change in coloration of
plains zebras to the south of their distributional range is achieved by stripe-
ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 35
Fig. 5. Examples of true Burchell’s zebra, Equus quagga burchelli, shown in standardized outline.
A. Type of ‘Equus burchelli paucistriatus’ Hilzheimer, 1912, at Mainz Museum. B. Animal that
lived at Jardin des Plantes, Paris, 1826. C. Type of subspecies from Little Klibbolikhonnifontein,
British Museum (Natural History) (destroyed). D-E. Animals that lived at Knowsly Park,
about 1845. F. Animal that lived at Zoological Garden, Dresden. (Adapted from Cabrera
(1936).)
36 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 6. Examples of Equus quagga antiquorum, shown in standardized outline. A. From
Rietfontein West, Munich Museum. B. Type of subspecies from Angola (Smith 1841).
C. From eastern Transvaal. D. From west of Lake Ngami. E. From Zululand.
F. From eastern Transvaal. (Adapted from Cabrera (1936).)
ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 37
reduction and the darkening of basic colour and the occurrence of shadow-
stripes. However, the above specimens indicate that these processes do not
necessarily occur together. Indeed, there are specimens from other areas which,
in spite of more extensive striping, display dark basic colour (Fig. 8A), some
with numerous strong shadow-stripes (sometimes three in one interspace). From
the Zululand skins (Fig. 7) and additional photographs it appears that stripe-
reduction is the dominant feature in this eastern population, while to the west
on the interior plateau the darkening in both basic colour and through shadow-
stripes is more pronounced.
In terms of stripe reduction these Zululand skins certainly do not fit into
the concept of Equus quagga antiquorum (Fig. 6), but they do agree with the
description of Equus quagga burchelli. Indeed, a few are less striped than some
of the burchelli specimens figured by Cabrera (1936) (Fig. 5) and others. It
appears, therefore, that Equus quagga burchelli is, in fact, not extinct but still
survives in Zululand (and possibly Swaziland), and that its range does extend
eastward into Zululand, as Matschie (1894) stated. (The ‘myriads of Quaggas’
which Chapman (1868, cited in Antonius 1951: 96) encountered in the extreme
north-eastern Orange Free State could thus have been true burchelli, and not as
Antonius (1951: 96) considered, Wahlberg zebra.) Otherwise the subspecies
burchelli would have to be considered as part of the antiquorum complex, a
thought similarly expressed by Rzasnicki (1951) and Roberts (1951), cited in
Ansell (1971). The only other alternative would be not to recognize such
individuals (Figs 7B—D, 9B, D) as members of the subspecies burchelli. However,
had they been old mounted specimens, they would be regarded as true Bur-
chell’s zebras.
It must be noted that the farm where these specimens had been shot is a
hunting farm which was originally stocked with 300 zebras from Hluhluwe
Game Reserve, and that hunters select the most striped specimens (C. Tinley,
pers. comm.). Selective breeding is thus exercised unintentionally and this might
have ‘re-created’ burchelli, although similar specimens do occur in Hluhluwe
and Umfolozi Game Reserves, the only area in Zululand where the original
zebra population has not been interfered with through reintroductions
(P. M. Brooks, pers. comm.).
Zululand undoubtedly is an inter-gradational area between the two forms
burchelli and antiquorum. Observations made during a recent visit to that area
and Swaziland suggest that the percentage of burchelli types may be as high as
15 in Zululand, although P. M. Brooks, biologist at Hluhluwe, estimates it to be
5-10 per cent (pers. comm.).
The north-eastern and eastern parts of Swaziland also retain an endemic
population of plains zebra (T. E. Reilly, pers. comm.). There is no geographic
barrier between Zululand and the Swaziland lowveld, and it is not surprising
that some of the zebras observed there fit or approach the description of Equus
quagga burchelli, though the percentage of such specimens must be expected to
be lower than further south.
38 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 7. Skins of plains zebras from Zululand, 1977. A. Note stripe-reduction, few shadow-
stripes. B-D. Note shadow-stripes on neck and stripe-reduction. (B now at South African
Museum, ZM38819.)
ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 39
ys _
es
Petey.
Td | - |
ww 7g
_- *
mm <<
Fig. 8. Skins of plains zebras from southern Africa. A. Precise locality unknown; note dark
basic colour of upper parts compared to lower leg. B. Precise locality unknown; note numerous
shadow-stripes and extensive striping. C. From game farm in Zululand; note dark basic colour
of upper parts. D. From southern Kaokoveld, South West Africa; note little contact between
body-stripes and dorsal midline.
40 ANNALS OF THE SOUTH AFRICAN MUSEUM
Despite the uncertainty whether the ‘quaggas’ of old reports were, indeed,
the true quagga, Equus quagga quagga, the northern boundary of its range is
usually considered to have been formed roughly by the Orange and Vaal rivers.
The eastern limit might have been about the boundary between the Orange Free
State and Natal (Grout, cited in Shortridge 1934: 402). It is also generally
accepted that the ranges of quagga and true Burchell’s zebra overlapped in the
Orange Free State (Antonius 1951; Liebenberg 1964; Ansell 1971).
Antonius (1951) was of the opinion that the quagga, being essentially a form
adapted to the Karoo and possibly the most distinct of all the plains zebra
subspecies, expanded its range secondarily in a northerly direction and met and
overlapped along a broad front with the southward expanding Burchell’s zebra.
He obviously did not believe in reports about the social and reproductive
separation of the two forms, since he considered one of the preserved quagga
specimens (the more striped specimen of ‘Equus burchelli paucistriatus’ at Mainz
(Fig. 9A and Rau 1974, fig. 13) as a possible cross between the two forms
(Antonius 1951: 26).
It is difficult to accept that separate herds of the two forms existed in the
same locality. Even if they did represent separate species, as some are inclined to
believe, herds formed by members of both must be expected, since this is known
to occur in extant zebra species where their ranges overlap. For example, Equus
zebra hartmannae overlaps with Equus quagga antiquorum in South West Africa
(Antonius 1951: 82) and Equus grevyi forms mixed herds with Equus quagga
boehmi in Kenya (Antonius 1951: 82; Bartlett 1963). Even true quaggas were
seen together with the Cape mountain zebra at Geelbeckfontein, at the southern
limit of the Karoo, by Barrow (1801, cited in Antonius 1951: 82).
As shown previously (Rau 1974) the quagga and the true Burchell’s zebra
approach one another in basic colour and in stripe reduction, so much so that
some of the preserved specimens (e.g. Rau 1974, fig. 12 right) have variously
been regarded as one or other of the two. Furthermore, some of the charac-
teristics previously considered to be typical of the quagga (Rau 1974) have been
observed in the skins at the Cape Town tannery and in a specimen from Kruger
National Park. These characteristics include narrow interspaces (Smuts 1974,
fig. 1.4.5), dorsal midline being flanked by a light band on either side, and body-
Stripes not in contact with dorsal midline except sometimes via narrow bridges
(Fig. 8D).
Antonius (1951: 29) has already pointed out that the quagga had reddish
dark-brown stripes which, especially on the body, merge into the reddish basic
colour. It has previously been explained why the coloration of the quagga has
been incorrectly interpreted as light stripes on a dark background (Rau 1974: 43).
Careful examination reveals beyond all doubt that the quagga, like all the other
plains zebras, is dark-striped and that the interspaces darken progressively from
the shoulder in a posterior direction until they become equal in tone with stripes
and shadow-stripes to produce the ‘uniform’ posterior portion. In other southern
plains zebras the interspaces are also lightest on head and neck. Ansell (1971: 5)
ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 4]
and others are wrong in defining the striping in the quagga as ‘whitish on dark
background’.
Since identification is therefore complex, and since it is impossible to
observe the stripe pattern of each individual when looking at a herd, the alleged
recognition of separate herds of quaggas and Burchell’s zebras in the same
locality must be treated with suspicion.
The grouping of plains zebras into subspecies is thus somewhat vague, or,
as it is expressed by Ansell (1971) ‘has never been satisfactory’. In an attempt to
sort out this confusion, Pocock (1897) pointed out that throughout the distri-
bution area of the plains zebra ‘intermediate types’ do exist between particular
forms which prevail in particular localities. In addition, as mentioned earlier,
typical representatives of the two forms occur in intergradational areas, the
closer to the area of the one form the lower the percentage of the other form.
The high degree of variability, both between geographically distant
populations and within one herd, has been attributed by Antonius (1951) to the
process of eliminating the contrasting stripe-pattern. It should be noted that
Antonius’s ‘Riickbildung der Bein- und Rumpfstreifung’ has been misinterpreted
by Eloff (1966, 1967) as being a sign of genetic pauperization.
As shown above, the gradual change from one form to another in plains
zebras appears to include, at least in exterior characteristics, the ‘separation’ into
Equus quagga and Equus burchelli. Relatively few skulls of the true quagga are
available and some of these lack sufficient data to establish their authenticity.
The fact that some skulls may have been obtained by breaking up an old stuffed
-quagga skin (as has happened at Berlin, where the skull was later identified as
that of Equus caballus (Antonius 1951: 35)) is no guarantee of their identity as
Equus quagga quagga.
However, some zoologists believe that there are specific morphological
differences between skulls of the quagga and the plains (Burchell’s) zebra
(e.g. Cooke 1950; Lundholm 1951; Gentry 1975; V. Eisenmann, pers. comm.).
The recent study by V. Eisenmann (pers. comm.) has shown that in some skull
characters there is a cline from northern to southern plains zebras. In other
words, in some respects at least skulls of southern plains zebras approach those
of the quagga more closely than do northern forms. This conforms to the
well-documented cline in colour and stripe-pattern.
To conclude, it would appear that the Burchell’s zebras and the quagga
should be regarded as one species, Equus quagga. Furthermore, since the
grouping into subspecies, either by appearance or by distribution, is not clear-cut
in the case of the plains zebras, it might be preferable to refer them to forms,
types or variations prevailing in certain areas, rather than to subspecies.
REINTRODUCTIONS OF ZEBRAS
The reintroduction of animal species into areas where natural populations
have been exterminated is increasingly common. There are, however, problems
with this practice, some of which have been discussed by Greig (1977).
42 ANNALS OF THE SOUTH AFRICAN MUSEUM
V BRANCO
Dis A h -
( V BRANCO
Fig. 9. Stripe patterns and tone of basic colours of various plains zebras, shown in standardized
outline. A. Female quagga at Mainz Museum. B. Burchell’s zebra from Zululand (same as
Fig. 7B). C. Male Burchell’s zebra at Leiden Museum. D. Burchell’s zebra from Zululand
(same as Fig. 7D). E. Quagga at Tring Museum (near London). F. Type of true Burchell’s zebra,
British Museum (Natural History) (destroyed). G. Quagga at Berlin Museum. H. Male
quagga at Mainz Museum. I. Burchell’s zebra at Mainz Museum (type of ‘Equus burchelli
paucistriatus’ Hilzheimer, 1912).
= —--—-
ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 43
44 ANNALS OF THE SOUTH AFRICAN MUSEUM
Both plains and mountain zebras are amongst those South African
mammals whose ranges have been much restricted during the past three
centuries. Now, and in the recent past, zebras are being translocated from areas
where populations survive to areas which they formerly inhabited. In the case
of the plains zebras such reiniroductions have involved the movement into
certain areas of forms dissimilar to those which previously occurred there, even
though the original form, or one close to it, still exists.
As indicated above, the ‘subspecies’, Equus quagga burchelli, apparently
survives in Zululand and thus, when reintroductions of zebras into the former
range of the true Burchell’s zebra are contemplated, it would be desirable that
selected specimens from Zululand be used, rather than animals of the antiquorum
type. It would even be desirable to eliminate former indiscriminate reintro-
ductions and to replace them with individuals which are closer to the original
population. With a minimum of initial selective breeding in order to eradicate
recessive antiquorum characteristics, which are present also in burchelli-like
individuals from Zululand (an intergradational area between the two forms),
a ‘pure’ population of plains zebras resembling the former endemics could be
achieved within a relatively short period.
ACKNOWLEDGEMENTS
My sincere thanks are due to Dr Vera Eisenmann of the Muséum National
d’Histoire Naturelle, Paris, for kindly informing me of the mounted quagga
which she discovered at the museum at Lyon, France. I also thank Dr C. A.
Howes for drawing my attention to the quagga foot which is housed at the
Royal Albert Memorial Museum, Exeter, England, Dr I. Sokolov of the
Academy of Sciences, Leningrad, U.S.S.R., and Mr E. Grangqvist of the
Zoological Museum, Helsinki, Finland, for being instrumental in obtaining
photographs and data of the mounted quagga in the Museum at Kazan, U.S.S.R.
I am indebted to Mr V. Branco for producing the drawings, Mr A. Byron for
some of the photographic reproductions and to Prof. G. H. Findlay, and
Drs Q. B. Hendey and P. A. Hulley for their co-operation.
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Smuts, G. L. 1974. Growth, reproduction and population characteristics of Burchell’s zebra in
the Kruger National Park. Unpublished D.Sc. thesis, University of Stellenbosch.
St Lecer, J. 1932. On Equus quagga of south-western and eastern Africa. Ann. Mag. nat.
Hist. (10) 10: 587-593.
TEGETMEIER, W. B. & SUTHERLAND, C. L. 1895. Horses, asses, zebras, mules and mule breeding.
London: Horace Cox.
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FiscHer, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.
Konun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.
Koun, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
TutELe, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: SCHULTZE, L. Zoologische
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ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 77 Band
December 1978 Desember
Part 3 Deel
REDESCRIPTION OF PLIOPLATEIA K. H. BARNARD,
A GENUS OF AMPHIPOD (CRUSTACEA) FROM
SOUTH AFRICA
By
J. LAURENS BARNARD
Cape Town Kaapstad
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REDESCRIPTION OF PLIOPLATEIA K. H. BARNARD,
A GENUS OF AMPHIPOD (CRUSTACEA) FROM SOUTH AFRICA
By
J. LAURENS BARNARD
Department of Invertebrate Zoology, Smithsonian Institution, Washington
(With 4 figures)
[MS. accepted 12 September 1978]
ABSTRACT
Plioplateia K. H. Barnard (1916) is removed from Phliantidae to form the type genus of
a new family demonstrating evolutionary outflow from the southern Pacific family Ceinidae
towards the circumtropical family Phliantidae. At least nine major characters constrain
Plioplateia from assignment to Phliantidae. Plioplateia appears to be the last living relict of
what may have been a diverse group of taxa standing between ceinids and phliantids. It joins
many other South African amphipods now considered to be relicts.
CONTENTS
PAGE
Introduction . . ee ce ed ee A,
Plioplateidae, fam. nov. . . . . . 48
Plioplateia K. H. Barnard . 49)
Plioplateia triquetra K. H. Barnard . : OO
Acknowledgements . .. . . . S55
Retercncest Paie® Fhe eres, Legett 55
INTRODUCTION
Plioplateia, a gammaridean amphipod, was described by K. H. Barnard
(1916) and placed in the family Phliantidae. Since that time many genera in
this evolutionary vicinity have been described and then later realigned, mainly
by J. L. Barnard (1972a, 19726) and Griffiths (1975), into families Phliantidae,
Temnophliidae, Kuriidae and Ceinidae, with subfamily Chiltoniinae. Plioplateia
appears to stand between Ceinidae and Phliantidae but differs in so many ways
from either family group that a new family is established for Plioplateia. All of
these families are included in the superfamily Talitroidea.
The Plioplateidae join the Temnophliidae of South Africa as the narrowest
relict families known in the Gammaridea except for the Kuriidae from
Abd-el-Kuri near the Gulf of Aden. South Africa is noted for other isolated or
relict amphipods, mainly the genera Calliopiella Schellenberg, Cypsiphimedia
K. H. Barnard, Dikwa Griffiths, Triodos K. H. Barnard, Hoplopleon K. H.
Barnard, Exampithoe K. H. Barnard, Macropisthopus K. H. Barnard, Unciolella
Chevreux, Elasmopoides Stebbing, Cunicus Griffiths and Phoxostoma K. H.
Barnard.
47
Ann. S. Afr. Mus. T7 (3), 1978: 47-55, 4 figs.
48 ANNALS OF THE SOUTH AFRICAN MUSEUM
Plioplateidae fam. noy.
Type genus
Plioplateia K. H. Barnard, 1916.
Diagnosis
Talitroidea with laterally compressed tall, cuspidate bodies but coxae
poorly splayed and excavate or cuspidate; coxa 4 small. Cuticle humped and
rough. Head dorsoventrally compressed, complexly cuspidate. Antennae
disparate, cuspidate, flagella elongate. Anterior part of body dorsally cuspidate.
Mandibular molar huge, granular, not distinctly triturative. Inner lobes of
lower lip well developed. Outer plate of maxilla | with eight spines. Inner plate
of maxilla 2 much narrower and shorter than outer. Outer plate of maxilliped
not larger than inner plate, palp articles | and 4 elongate, palp thin.
Gnathopods subchelate, hands expanded, lacking giant facial setae.
Article 2 of pereopods 5-7 narrow, small.
Pleon small, deeply flexed below thorax, urosomite 3 distinct but vestigial.
Pleopods elongate, peduncles thin, rami elongate, thin, uni-articulate. Uropod 3
formed of leaf-like peduncle lacking ramus. Telson forming erect cowl, deeply
cleft.
Relationship
On first sight Plioplateia appears to belong to the Phliantidae as originally
placed by K. H. Barnard, but Plioplateia differs from phliantids in the elongate
flagella of the antennae, the elongate, thin pleopods with equal and uniarticulate
rami, the giant mandibular molar, the presence of well-developed inner lobes
on the lower lip, the small inner plate of maxilla 2, the small outer plate of the
maxilliped, the thin maxillipedal palp with elongate article 1, the subchelate
gnathopods lacking giant setae, and the cowl-like cleft telson. Plioplateia bears
the bidentate pereonite | similar to that of many phliantids but would appear to
be much more primitive and to share many features of the plesiomorphic
Ceinidae (see J. L. Barnard, 1972a, 1972b). To a great extent Plioplateia fits
many aspects of a model ancestor to phliantids with evolutionary outflow
from ceinids, and, in addition, its isolation in South Africa fits the role of a
relict.
Plioplateia shares the tall body and subchelate gnathopods of ceinids, and
in many other characters (to follow) appears to have a foundation in Taihape
J. L. Barnard (1972a) or Waitomo J. L. Barnard (1972a): size and flagellar
condition of antennae in Waitomo; posterior dorsal cuspidation of Waitomo;
molar of Waitomo; outer plate of maxilla 1 in both genera; maxillipedal palp
of Taihape; gnathopods (less giant setae) and telson of either genus.
Plioplateia differs from Ceinidae in the dorsoventrally depressed head with
complex cuspidation, the anterodorsal body cuspidation, the cuspidation and
excavation of coxae and antennae, the special form of maxilla 2, the lack of
REDESCRIPTION OF PLIOPLATEIA K. H. BARNARD 49
large setae on the gnathopods and the presence of inner lobes on the lower lip,
wholly unique to Pliop/ateia in the Ceinid—Phliantid scheme.
The reduction in segmentation on the pleopods suggests that Plioplateia
has progressed far from a perfect intergrade between ceinids and phliantids and
this is also supported by the extra cuspidation on the head and coxae, the
development of inner lobes on the lower lip, the reduction in size of the inner
lobe on maxilla | and the outer plate of the maxilliped, the enlargement of the
molar, the loss of giant setae on the gnathopods. The brood plates appear to
be much larger than in ceinids and phliantids whereas the gills are much smaller.
The weakly developed multispination on the apices of uropods 1-2 would
appear to be a remnant of ancestry in ceinids where these spines are well
developed.
The Kuriidae, composed only of Kuria Walker & Scott (1903), from
Abd-el-Kuri, appear also to be in a level of evolution similar to Plioplateia.
For example, the gnathopods of Kuria are subchelate, though weakly, the
palms being almost transverse, and the outer plate of the maxilliped is small
as in Plioplateia. Plioplateidae differ from Kuriidae in the ornamentation of the
head, body and coxae, the small coxa 4, small article 2 of pereopods 5-7,
expanded hands of the gnathopods, the disparity in sizes of antennae, and the
extremely large mandibular molar. In minor ways, Plioplateidae differ from
Kuriidae in the larger dactyls of the maxillipeds, and the absence of a ramus on
uropod 3. Pleopods, maxillae and lower lip have not been reported for Kuria.
Plioplateia K. H. Barnard
Plioplateia K. H. Barnard, 1916: 155.
Type species
Plioplateia triquetra K. H. Barnard, 1916 (monotypy).
Diagnosis
With the characters of the family.
Description
Rostrum well developed, erect, thorn-like, bearing bilateral subsidiary
tooth. Antenna | much longer than antenna 2, flagella of both pairs highly
articulate, those of antenna | bearing 1-2 aesthetascs each. Epistome rounded
anteriorly, upper lip deeply incised, asymmetrical. Mandibular incisor well
toothed, right lacinia mobilis, if present, composed of three fused spines (or
these actually rakers and lacinia mobilis absent), left lacinia mobilis large and
well toothed, left mandible with three raker spines; molar very large, broad,
blunt, tumid, poorly triturative, mainly granular apically; palp absent or
possibly represented by small leaf. Lower lip with well-developed inner lobes.
Outer plate of maxilla 1 with eight spines, palp uniarticulate, of medium size,
bearing one medium apical seta. Inner plate of maxilla 2 much shorter and
50 ANNALS OF THE SOUTH AFRICAN MUSEUM
narrower than outer, inner subconical, outer subrectangular, both poorly
setose but inner with one medial and outer with two lateral setae. Inner plate
of maxillipeds leaf-like, with tapered base, poorly armed, outer plates as large
as inner; palp thin, article 1 elongate, article 4 unguiform, greatly elongate,
with three apical setae. Pleurae of pereonites produced, rugose, humped or
cuspidate.
Plioplateia triquetra K. H. Barnard
Figs 1-4
Plioplateia triquetra K. H. Barnard, 1916: 156, pl. 26 (figs 18-24); Griffiths, 1974: 328.
Diagnosis
With the characters of the family and genus.
Description
Head depressed but with erect thorn-like, rostrum with smaller basal
tooth on each side, lateral lobes with weak dorsal and strong ventral tooth,
middle of lobe bulging laterally and containing small but multifaceted ommati-
dial eye, antenna | inserted by sleeve into pocket anterior to ocular lobe,
anteroventral area of head extended forward through fusion of articles 1 and 2
of antenna 2, lateral surface of article 2 with large cusp, article 3 also with large
lateral cusp, gland cones emerging ventrally from fused article 2. Mouthpart
field from lateral view, apart from maxillipeds, dominated by outer surface of
lower lip.
Antenna | elongate, articles 1 and 2 cuspidate, article 3 shorter than
article 1 of flagellum, latter 12-articulate, each article of flagellum with 1-2
aesthetascs and several curled setae. Accessory flagellum absent but marked by
weak sinuate stripes inside of article 3. Antenna 2 small and slender, article 4
weakly cuspidate, article 5 slightly longer than article 4 of peduncle on article 1
of flagellum, latter 9-articulate, with short stiff curled setae.
Upper lip scarcely distinct from epistome, together rounded anteriorly,
upper lip deeply bilobed. Each mandibular molar with large setule, right incisor
with 8-9 teeth, left with 3, left lacinia mobilis with 7 teeth, right either absent
or formed of 3 fused spines, left mandible with 3 distinct rakers each independent
and mostly fused to mandible. Mandibular lobes of lower lip well developed,
inner lobes distinct, thin across faces, broad, widely separating outer lobes.
Inner plate of maxilla | linguiform, of medium size.
Wrist of gnathopod | longer than hand, shorter on gnathopod 2, neither
lobate, palm well developed, oblique, defined by pair of spines, armed with
pairs of wire-setules.
Coxa 1 apically expanded, with deep posteroventral notch, coxae 2-4
somewhat tapered, each with weak or moderate notch, coxa 4 smaller than
coxa 1, not excavate posteriorly; coxae 5-7 short, coxae 5 and 7 bilobed and
acuminate, coxa 6 trilobed and acuminate.
REDESCRIPTION OF PLIOPLATEIA K. H. BARNARD 51
Fig. 1. Plioplateia triquetra K. H. Barnard. h, holotype, female ‘h’ unmeasured; p, male
‘p’ 5,72 mm. A. Prebuccal outline, left lateral. B. left mandibular rakers. C. Ventral view
of head, upper lip hatched; a, antenna 2, g, gland cone. D. Head, lateral; a, antenna 2
broken off. E. Prebuccal, left lateral. F. Body, scale = 1 mm. G. Right mandibular rakers.
H. Head. I. Pleon, left lateral; u, uropod 3. J. Upper lip, anterior.
52
ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 2. Plioplateia triquetra K. H. Barnard. male ‘p’ 5,72 mm. A. Antenna 2. B. Antenna 1.
C. Left mandible. D. Mouthparts, left side; e, epistome, i, lower lip, 1, upper lip,
m, mandible, x, maxilla. E. cuticle. F. Right mandible. G. Left mandible. H. Lower lip,
oral side. I. Outer plate of maxilla 1. J. Maxilla 1. K. Palp of maxilliped, flattened.
L. Maxilla 2. M. Lower lip, aboral side. N. Maxilliped.
REDESCRIPTION OF PLIOPLATEIA K. H. BARNARD 53
Fig. 3. Plioplateia triquetra K. H. Barnard. male ‘p’ 5,72 mm, 0, outer ramus. A. Gnathopod
1. B. Gnathopod 2. C. Pleopod 1, rear. D. Pleopod 2, rear. E. Pleopod 3, rear. F. Pereo-
pod 7. G. Pereopod 5. H. Pereopod 6. I. Pereopod 4.
54 ANNALS OF THE SOUTH AFRICAN MUSEUM
Article 2 of pereopods 3-4 uncuspidate, that of pereopods 5—7 with bicuspi-
date posteroventral process; defining armaments on article 6 composed of
setae. Coxal gills very small, attached to segments 2-6, somewhat triquetral
(see appendages attached to figures of coxae 2-4). Brood plates very large,
lamellar, attached to coxae 2-5, densely furnished with coil-tipped setae. Male
penes of pereonite 7 highly lateral, just basal to coxae, very elongate and sausage-
shaped.
Epimera tapering distally, poorly armed and unornamented. Pleopods
tightly clumped, decreasing in size from front to rear slightly, inner rami scarcely
shortened, each with 2 apical setae longer than ramus, except inner ramus of
pleopod 3 with only one apical seta, peduncles elongate, only pleopod 3 with
pair of apicomedial coupling spines.
Urosomite 3 vestigial, represented only by ventral plate, telson on dorsal
side appearing attached directly to urosomite 2, telson formed of bifid cowl
lacking macroscopic armament. Uropods 1-2 short, stout, poorly armed, outer
Fig. 4. Plioplateia triquetra K. H. Barnard. male ‘p’ 5,72 mm, g, gill, o, outer ramus.
A. Uropod 3. B. Uropod 2. C. Coxa 2. D. Coxa 1. E. Coxa 3. F. Pleonite 6, ventral,
telson hatched. G. Coxa 4. H. Uropod 1. I. Uropod 1, rami enlarged. J. Uropod 2.
K. Pleonites 1-2 dorsal showing telson, t.
REDESCRIPTION OF PLIOPLATEIA K. H. BARNARD yes
ramus of uropod | shortened, each ramus of both uropods with apical jewel-nail
plus accessory spine except on inner ramus of uropod 2, inner ramus of uropod
1 with third spine fixed subapically. Uropod 3 ovate, bearing weak apical
spinule.
Cuticle densely pebbled, each pebble usually complex, either with apparent
pit or appearing ring-shaped (with central vacuole) bulbar setules sparse, pipes
often abundant; no pits typical of ceinids have been observed.
Pereonite 1 with large tooth dorsally, remaining pereonites each with
single tooth or hump, pleonites 1-3 each with increasingly smaller dorsal
hump, urosomite | with slightly larger hump, this segment covering urosomites
2-3 dorsally; pereonites 1-7 from front to rear with increasingly complex
distolateral rugosities or cusps.
Holotype
South African Museum A174, female ‘h’ (not measured to prevent damage)
lacking right antennae, uropods 2-3, telson, right uropod 1.
Other material
South African Museum, University of Cape Town SCD 310 F, male ‘p’
5,72 mm (dissected and illustrated herein).
Distribution
South Africa, 50-91 m.
ACKNOWLEDGEMENTS
I thank Dr C. L. Griffiths of the C.S.I.R. Oceanographic Research Unit,
University of Cape Town, for locating this material, and Dr T. H. Barry
Director of the South African Museum, for his kind assistance. Carolyn L. Cox
of Smithsonian Institution inked and prepared the illustrations for publication;
she also created several of the original drawings.
REFERENCES
BARNARD, J. L. 1972a. Gammaridean Amphipoda of Australia, Part I. Smithson. Contr.
Zool. 103: 1-333.
BARNARD, J. L. 19726. The marine fauna of New Zealand: Algae-living littoral Gammaridea
(Crustacea Amphipoda). Mem. N.Z. oceanog. Inst. 62: 7-216.
BARNARD, K. H. 1916. Contributions to the crustacean Fauna of South Africa. 5.—The
Amphipoda. Ann. S. Afr. Mus. 15: 105-302.
GriFFitHs, C. L. 1974. The Amphipoda of southern Africa. Part 4. The Gammaridea and
Caprellidea of the Cape Province East of Cape Agulhas. Ann. S. Afr. Mus. 65: 251-336.
GriFFITHS, C. L. 1975. The Amphipoda of southern Africa. Part 5. The Gammaridea and
Caprellidea of the Cape Province west of Cape Agulhas. Ann. S. Afr. Mus. 6: 91-181.
WaLker, A. O. & A. Scott. 1903. Decapod and sessile-eyed crustaceans from Abd-el-Kuri.
Nat. Hist. Sokotra and Abd-el-Kuri: 216-232.
7
dt ‘oe oo
»
ee aon pe.
HRA G: @
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REDESCRIPTION OF PLIOPLATEIA
K. H. BARNARD, A GENUS OF AMPHIPOD
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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FiscHer, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.
Koun, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
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(continued inside back cover)
ee Se ie ee
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 77 Band
December 1978 Desember
Part 4 Deel
CRETACEOUS FAUNAS FROM ZULULAND AND
NATAL, SOUTH AFRICA
A NEW GENUS AND SPECIES OF GASTROPLITINAE
FROM THE MZINENE FORMATION (ALBIAN)
By
WILLIAM JAMES KENNEDY
&
HERBERT CHRISTIAN KLINGER
Cape Town Kaapstad
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are issued in parts at irregular intervals as material
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap
CRETACEOUS FAUNAS FROM ZULULAND AND NATAL,
SOUTH AFRICA
A NEW GENUS AND SPECIES OF GASTROPLITINAE FROM THE
MZINENE FORMATION (ALBIAN)
By
WILLIAM JAMES KENNEDY
Geological Collections, University Museum, Oxford
&
HERBERT CHRISTIAN KLINGER
South African Museum, Cape Town
(With 7 figures)
LMS. accepted 20 September 1978]
ABSTRACT
Alopecoceras gen. nov., type species Alopecoceras ankeritterae sp. nov., is a unique
Southern hemisphere homoeomorph of the Boreal genus Neogastroplites McLearn, 1930,
which occurs in the low Middle Albian Mzinene Formation (Albian III) of the Mzinene River,
Zululand. Whereas Neogastroplites evolved from Gastroplites McLearn, 1930, by acquisition
of umbilical and ventral nodes and is of late Albian age, A/opecoceras is believed to be derived
from Hatchericeras Stanton, 1910, by a similar acquisition of nodes. The genus also shows
similarities to Tetrahoplitoides Casey, 1954.
CONTENTS
PAGE
AMtrOdUCHION: so we ee ee ews) on) ST
Systematic Palaeontology . . . . 58
Acknowledgements . . .. . . 68
References ay eee Pee ee Le 68
INTRODUCTION
The Mzinene Formation exposed along the Mzinene River | 200 m north-
east of the farm Amatis, north of Hluhluwe, Zululand (locality 35 of Kennedy
& Klinger (1975: 28), 27°58’03”S, 32°18'34’E) has yielded a series of remarkable
specimens which closely resemble the exclusively boreal late Albian ammonite
genus Neogastroplites McLearn, 1930 (see Reeside & Cobban 1960 for extensive
illustration of the genus) in association with a fauna of low Middle Albian age
in part described by the authors in previous publications (Kennedy & Klinger
1977, 1978, in press; Klinger 1976; Klinger, Wiedmann & Kennedy 1975), and
including Carinophylloceras collignoni Klinger, Wiedmann & Kennedy, Ammo-
noceratites (Ammonoceratites) mahadeva (Stoliczka), Anagaudryceras buddha
Ann, S. Afr. Mus. 77 (4), 1978: 57-69, 7 figs.
57
58 ANNALS OF THE SOUTH AFRICAN MUSEUM
(Forbes), Protanisoceras (Rossalites) aff. superbus Collignon, Lyelliceras and
Douvilleiceras species.
Further study has shown that the material represents a new genus and
species which is a heterochronous homoeomorph of Neogastroplites, and is
described below as Alopecoceras ankeritterae sp. nov.
All specimens are housed in the British Museum (Natural History) and
the Geological Survey of South Africa, designated BMNH and SAS respectively.
SYSTEMATIC PALAEONTOLOGY
Superfamily HOPLITACEAE H. Douvillé, 1890
Family Hoplitidae H. Douvillé, 1890
Subfamily Gastroplitinae Wright, 1952
Genus Alopecoceras Kennedy & Klinger gen. nov.
Type species
Alopecoceras ankeritterae gen. et. sp. nov., low Middle Albian, Zululand.
Derivation of name
Alopex (Greek) fox.
Diagnosis
Initially depressed, whorl section trapezoidal with sparse conical or bullate
umbilical nodes giving rise to pairs of ribs with additional ribs intercalated; all
ribs bearing conical ventral tubercles and connected across the fastigiate venter
by a much thickened rib. Whorl section becomes compressed and lanceolate in
later growth, venter narrowly rounded and ornament reduced to low falcoid
ribs with interspaces sometimes accentuated into feeble constrictions. Suture
line with moderately incised bifid elements.
Discussion
Alopecoceras closely resembles Neogastroplites McLearn, 1930, but is of
low Middle Albian age whereas that genus, a descendant of Gastroplites
McLearn, 1930, is of late Albian age. Because of the great intraspecific variation
in Neogastroplites (see Reeside & Cobban 1960) it is a little difficult to make
overall statements of difference, but in general Alopecoceras is consistently more
evolute, tubercles are lower on the flank, ribs fewer and less flexuous in juveniles,
the ventral ribbing thicker and less convex.
Neogastroplites typically develops a siphonal tubercle and is rostrate;
neither of these features are seen in Alopecoceras, whilst the suture lines are
more intricately subdivided.
When adult, Neogastroplites may become smooth and very compressed
(e.g. Reeside & Cobban 1960, pl. 17 (figs 17, 22), pls 18-19), but lack the
CRETACEOUS FAUNAS FROM SOUTH AFRICA 59
flexuous branching ribs and feeble constrictions of A/opecoceras. Furthermore,
these individuals have slender, feebly ornamented inner whorls; individuals of
a similar degree of inflation to Alopecoceras ankeritterae retain strong ribs and
nodes to a great size (Reeside & Cobban 1960, pl. 11 (figs 16, 20), pl. 12 (figs
11-12, 16), pl. 36 (figs 1, 5, 9-11)).
Alopecoceras can be easily separated from Gastroplites McLearn, 1930 (see
Reeside & Cobban 1960 pls 8-9), for that genus does not possess umbilical
bullae and is usually more involute.
Lemuroceras Spath, 1942, never develop strong umbilical nodes, shoulder
tubercles or a fastigiate venter (see illustrations in Collignon 1963).
Arcthoplites Spath, 1925 (see illustrations in Casey 1965, text-fig. 177), and
Subarcthoplites Casey, 1954, both have strong, narrow ribs, round venters and
lack tubercles, as does Cymahoplites Spath, 1922.
Alopecoceras shows closer similarities to Tetrahoplitoides Casey, 1954 (see
Fig. 7H—J); both have a trapezoidal whorl, although, in the type species at least,
the whorls are slender, the umbilical tubercles are bullate and not conical as in
Alopecoceras. The venter is narrower in Tetrahoplitoides, flat rather than
fastigiate, and lacks ventrolateral tubercles.
Evolutionary origins
Alopecoceras occupies an isolated geographic position, and its evolutionary
origins are far from obvious in the remaining faunas of both the South African
and Madagascar Albian. C. W. Wright has, however, called our attention to
its similarities to the genus Hatchericeras Stanton, 1901. Originally described
from Patagonia, this genus was referred to the Neocomitinae Spath, 1924, in
the Treatise, and regarded as of possibly Lower Hauterivian age. Subsequent
publication by Leanza (1970) has show it to be an early Albian gastroplitinid.
During middle growth (Fig. 7A—D) it differs from Alopecoceras very obviously
in the lack of umbilical nodes and in possessing numerous flexuous ribs. There
are, however, blunt thickenings of the ribs—incipient tubercles—at the ventro-
lateral shoulder in some species, and a broad, blunt rib crosses the venter.
Adults (Leanza 1970, fig. 3la—b) are compressed, with a rounded venter,
feeble flexuous ribs and striae, plus feeble constriction-like folds. The sutures
(Fig. 6C) of the two genera are also constructed on the same plan, with a
similar degree of incision.
The authors would conclude, therefore, that A/opecoceras is an offshoot of
Hatchericeras which has developed prominent umbilical nodes, accentuated
the ventral tubercles present in some of the latter, and developed a fastigiate
rather than flattened venter: it is a homoeomorphous Southern hemisphere
analogue of Neogastroplites rather than a close relative.
Occurrence
Alopecoceras is known only from the low Middle Albian of Zululand.
60 ANNALS OF THE SOUTH AFRICAN MUSEUM
Alopecoceras ankeritterae gen. et. sp. nov.
Figs 1-4, 5A-B, 6, 7E-G
Holotype
BMNH C81266 from locality 35, cliff and stream sections extending over
several hundred metres along the Mzinene, approximately 1 200 m north-east
of the farm Amatis, north of Hluhluwe, Zululand, 27°58’03”S 32°18’31"E
Mzinene Formation, Albian III.
Fig. 1. Alopecoceras ankeritterae sp. nov. The holotype, BMNH C81266. x 1.
CRETACEOUS FAUNAS FROM SOUTH AFRICA
Fig. 2. Alopecoceras ankeritterae sp. nov. Inner whorls of the holotype, BMNH C81266.
61
62 ANNALS OF THE SOUTH AFRICAN MUSEUM
Paratypes
BMNH C81267 to C81274, SAS H54a—b, SAS A584, all from the same
horizon and locality as the holotype; SAS EM152 from locality 171, on the
hill slopes west-south-west of Mlambongwenya Store, Zululand, Mzinene
Formation, Albian II-III, and SAS A589 from locality 53, a derelict dam site
on Indambana, south of Izwehelia Farm, north of Hluhluwe, Zululand, Mzinene
Formation, Albian II.
a Sa ey ae
= 2
Fig. 3. Alopecoceras ankeritterae sp. nov. Body chamber of the holotype, BMNH C81266. x 1.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 63
Fig. 4. Alopecoceras ankeritterae sp. nov. Paratypes BMNH C81267-C81271. A-C, E-G,
I-K, P-R x 2; D, H, L-O x 1.
64 ANNALS OF THE SOUTH AFRICAN MUSEUM
C | A |
D
B
Fig. SA-B. Whorl sections of Alopecoceras ankeritterae sp. nov. A. BMNH C81274. x 12.
B. BMNH C81268. x 6. C—D. Early whorls of BMNH C81274. x 12.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 65
oo
Fig. 6. A-B. Alopecoceras ankeritterae sp. noy. External sutures of BMNH C81268, x 12,
and BMNH C81266, x 3. C. Hatchericeras semilaeve Leanza. External suture of a specimen
in C. W. Wright's collection from Santa Cruz, Argentine. x 6.
66 ANNALS OF THE SOUTH AFRICAN MUSEUM
Derivation of name
The species is named for Anke Ritter of Gauting.
Diagnosis
As for genus.
Dimensions
All dimensions are in millimetres. D = diameter, Wb = whorl breadth,
Wh = whorl height, U = umbilical diameter, R = ribs per whorl.
Figures in parentheses are dimensions expressed as a percentage of diameter.
D Wb Wh Wb:Wh U R
Holotype
BMNH C81266 .« §37,5(100) 35026) 61,2(45) 0,57 34,0(25) =
at 97,9(100) 26,4(27) 42,8(44) 0,62 24,3(25) 32
at 67,2(100) 21,8(32). 30,0(45) 0,72 18,7(28) —
SAS A584 .. «+ 41,9(100) 19,5(47) 18,5744) 1,05 10,224) @
BMNH C81267 .. 25,5(100) 11,8(46) 10,5(41) = ‘1,12 7,2(28) 26
SAS H54a => on “23,9(100) . 10,046) —10;9%46) — im 6,0(25) 26
BMNH C81268 .. 22,7(100) 10,4(46) 85738) - AL9 6,3(28) 26
Description
The early developmental stages are seen in BMNH C81274. At a diameter
of 3,8 mm, the juvenile shell is cadicone, the umbilicus comprising approxi-
mately 30 per cent of the diameter, with a whorl breadth to height ratio of 1,4
(Fig. SC-D). Ornament consists of blunt, low, rounded umbilical nodes, which
first appear at a diameter of c. 3 mm. These occur at a rate of seven per whorl
up to a diameter of 6 mm, and in this interval, low, broad folds pass across the
flanks and venter, although never developing into clearly delineated ribs. By
6 mm diameter, the whorl breadth to height ratio has become 1,2.
From 6 mm (Fig. 4A—-R) the whorl section changes from depressed and
reniform to trapezoidal. The umbilicus is shallow, and the wall slopes gently
outwards. The greatest breadth is at the umbilical bullae, and the whorl! breadth
to height ratio is commonly reduced to 1,!. In intercostal section the outer
flanks are flattened, the ventrolateral shoulders broadly rounded and the venter
very broadly rounded. In costal section, the flanks are concave at mid-flank and
flattened on the outer flank, and the venter fastigiate.
There are eight to ten prominent umbilical bullae per whorl, placed well
out from the umbilical seam, and these give rise to groups of two or three broad
ribs, weakened at mid-flank, where additional short ribs are intercalated, and
recti- to feebly rursiradiate in some cases. All ribs bear a bullate to conical
tubercle at the ventrolateral shoulder, and there are commonly twenty-five to
twenty-eight ribs per whorl. The ribs pass across the venter with a faint con-
vexity, broadening and diminishing in elevation over the siphonal line.
i
CRETACEOUS FAUNAS FROM SOUTH AFRICA 67
Fig. 7. A-D. Hatchericeras semilaeve Leanza. Inner whorls of a specimen in C. W. Wright's
collection from Santa Cruz, Argentina. x |. E-G. Alopecoceras ankeritterae sp. nov. Para-
type, SAS H93/3. x 1. H-—J. Tetrahopliloides stantoni (Anderson) cast of the holotype, from
Texas Springs, Shasta County, California. H = 1, I-J x 2.
68 ANNALS OF THE SOUTH AFRICAN MUSEUM
This general style of ornament extends to a diameter of c. 50 mm; as size
increases, the whorls eventually change from depressed to compressed, the ribs
coarsen, and, on the venter, become very thick (Fig. 2C). There is some variation
in strength and number of ribs, bullae and whorl section at this stage, as can
be seen from the figures.
The features of later growth are shown by the holotype (Figs 1-3). Here,
from 50 mm onwards, all ornament declines, the ribs become low, broad and
flexuous, arising in pairs from the bullae, whilst the whorl becomes compressed
and lanceolate in section with a narrowly rounded venter which is crossed by
ribs and has, as a result, an undulose profile.
On the mature body chamber (Fig. 3), coiling has become more evolute,
with a low, outwards sloping, flattened umbilical wall. There is a somewhat
abrupt umbilical shoulder and high, lanceolate whorls (whorl breadth to height
ratio is 0,57 to 0,62) with a narrowly arched venter.
Ornament consists of irregular falcoid ribs and striae, arising from feeble
umbilical bullae, separated by interspaces which may be strengthened into
feeble, irregular constrictions.
The suture line (Fig. 6B) at maturity is moderately divided. E/L is broad
and asymmetrically bifid, L broad and asymmetrically bifid. L/U, is large,
although narrower than E/L, and asymmetrically bifid. U, is narrow and quite
deeply divided.
Discussion
Features which separate Alopoceceras ankeritterae from species of other
genera are fully covered in comments in the generic discussion on p. 58.
Occurrence
Low Middle Albian of Zululand only.
ACKNOWLEDGEMENTS
Our best thanks are due to Mr C. W. Wright for drawing our attention to
the similarities between Alopecoceras and Hatchericeras, as well as providing
useful discussion. We are also grateful to Dr M. R. Cooper (Oxford), Dr M. K.
Howarth and Mr D. Phillips (British Museum, Natural History) and the staff
of the Geological Collections, Oxford, and the South African Museum for
technical assistance.
REFERENCES
Casey, R. 1965. A Monograph of the Ammonoidea of the Lower Greensand. Palaeontogr.
Soc. (Monogr.) 6: 399-546.
CoLLIGNoN, M. 1963. Atlas des Fossiles caractéristiques de Madagascar (Ammonites), Fasicule X
(Albien). Tananarive: Service Géologique.
KENNEDY, W. J. & Kuincer, H. C. 1975. Cretaceous faunas from Zululand and Natal, South
Africa. Introduction, Stratigraphy. Bull. Br. Mus. nat. Hist. (Geol.) 25: 263-315.
KENNEDY, W. J. & KLINGER, H. C. 1977. Cretaceous faunas from Zululand and Natal, South
Africa. The Ammonite Family Phylloceratidae. Bull. Br. Mus. nat. Hist. (Geol.) 27:
347-380.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 69
KENNEDY, W. J. & KLINGER, H. C. 1978. Cretaceous faunas from Zululand and Natal, South
Africa. The ammonite family Lytoceratidae Neumayr, 1875. Ann. S. Afr. Mus. 74:
257-333.
KENNEDY, W. J. & KLINGER, H. C. In press. Cretaceous faunas from Zululand and Natal,
South Africa. The ammonite family Gaudryceratidae Spath, 1927. Bull. Br. Mus. nat.
Hist. (Geol.).
KLINGER, H. C. 1976. Cretaceous heteromorph ammonites from Zululand. Mem. S. Afr
geol. Surv. 69: 1-142.
KLINGER, H. C., WIEDMANN, J. & KENNEDY, W. J. 1975. A new carinate phylloceratid from the
early Albian (Cretaceous) of Zululand, South Africa. Palaeontology 18: 657-644.
LeANZA, A. F. 1970. Ammonites nuevos o pocos conocidos del Aptiano, Albiano y Ceno-
maniano de los Andes Australes, con notas acera de su posicion estratigraphica. Revta
Assoc. geol. argent. 25: 197-261.
ReesipE, J. B. & CoBBAN, W. A. 1960. Studies of the Mowry Shale (Cretaceous) and con-
temporary formations in the United States and Canada. Prof. Pap. U.S. geol. Surv. 355:
1-126.
6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.
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Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
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Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
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WILLIAM JAMES KENNEDY
&
HERBERT CHRISTIAN KLINGER
CRETACEOUS FAUNAS FROM ZULULAND AND
NATAL, SOUTH AFRICA
A NEW GENUS AND SPECIES OF GASTROPLITINAE
FROM THE MZINENE FORMATION (ALBIAN)
OLUME 77 PART 5 DECEMBER 1978 MUS. COMP. zoo. !SSN 0303-2515
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BuLLouGu, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
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FiscHER, P.-H., DuvaLt, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gen. 74: 627-634.
Konun, A. J. 1960a. Ecological notes = ead (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309.
Konn, A. J. 19606. Spawning hate te oe egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
Tues, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 77 Band
December 1978 Desember
Patt 5 Deel
PPreR CRETACEOUS SEDIMENTS FROM THE
IGODA RIVER MOUTH, EAST LONDON,
SUU TE APRICA
By
HERBERT CHRISTIAN KLINGER
&
BRIAN E. LOCK
Cape Town Kaapstad
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UPPER CRETACEOUS SEDIMENTS FROM THE IGODA RIVER
MOUTH, EAST LONDON, SOUTH AFRICA
By
HERBERT CHRISTIAN KLINGER
South African Museum, Cape Town
&
BRIAN E. LOCK,
The University of Southwestern Louisiana, Lafayette
(With 7 figures)
[MS. accepted 20 September 1978]
ABSTRACT
The name Igoda Formation is proposed for a sedimentary sequence consisting mainly of
calcareous sandstones and arenaceous limestones exposed on the bluffs overlooking the
Igoda River Mouth near East London, South Africa. The invertebrate fauna indicates a Late
Cretaceous age, probably Late Campanian to Early Maastrichtian. Faunal paucity precludes
detailed comparison with other areas in southern Africa, but common faunal elements occur
at Lower Needs Camp (here regarded as a lateral facies equivalent of the Igoda Formation),
Pondoland, Zululand, Madagascar and Angola. Affinities with Madagascar are strongest.
CONTENTS
PAGE
IMtTOMUCIONNE Ee kes ee ees ee fe) aes 71
The Igoda River Mouth section. . . ... . 73
ithe roadsection™. = = | me Mew eet ek is)
Fauna and age of the Igoda Formation ... . 75
Comparison with other areas in southern Africa . . 81
SURIUMAY; Mee ooo SP ee a ty Gee a) ce. Bech 82
Acknowledgements =. . 5 6 4 «a 83
IREfEDeD CES Twat Pe un ei tcweils, Say Wee eters 83
INTRODUCTION
Late Cretaceous fossiliferous limestones have long been known from the
eastern Cape from one of the small quarries on the farm Needs Camp (the
East or Lower Quarry) near East London (Lang 1908; Woods 1908; Chapman
1916). McGowran & Moore (1971) established a probable Upper Senonian
(Campanian to Maastrichtian) age for this deposit on the basis of the micro-
faunal content. Contrary to a report by King (1972), the second quarry at
Needs Camp (the West or Upper Quarry) is excavated in limestones of Tertiary
age (Lock 1973 and in preparation). Microfaunal investigations confirm this
Ann. S. Afr. Mus. 77 (5), 1978: 71-83, 7 figs.
71
TZ ANNALS OF THE SOUTH AFRICAN MUSEUM
(W. G. Siesser pers. comm.), although reworked Cretaceous material occurs in
the limestones.
Until 1974 no other outcrops of Late Cretaceous marine sediments had
been reported from south of the Umzamba Estuary (Pondoland, Transkei).
Mountain (1974: 19) briefly described a new fossiliferous locality less than
15 km from Needs Camp as follows: The rock occurs in a ‘tiny exposure on
the East London—Kidd’s Beach road just west of the Goda River valley as the
road climbs up on to the coastal plain. It can be seen in the present road cutting
and consists of a friable limestone with abundant echinoid spines, Ostrea shells
and gastropod casts. It is possibly Cretaceous but no one has examined the
microfossils.’
Following landslides during the last few years, an extension of this occur-
rence became exposed in the bluffs overlooking the western shore of the lagoon
at Igoda Mouth, less than a kilometre from the roadside section mentioned by
Mountain (Fig. 1).
Tertiary and ?Pleistocene
calcareous sands
Igoda Formation
Beaufort sedimentary rocks
and Karroo dolerite
Fig. 1. Locality map, Igoda Formation. A. Roadside exposure reported by Mountain (1974);
note the new road under construction. B. Type section of the Igoda Formation. C. Reference
section. (See also Fig. 2.)
UPPER CRETACEOUS SEDIMENTS FROM THE IGODA RIVER MOUTH te
THE IGODA MOUTH SECTION
The succession in the new exposures is summarized in Figure 2.
The Cretaceous sequence, here named the Igoda Formation, rests uncon-
formably on sandstones and mudstones of the Permian/Triassic Beaufort Group,
which have been intruded by dolerites of later Karoo age. The contact is one
of some relief. The type section (sections BI, B2) is a composite one (see Fig. |
for location and Fig. 2 for sections), and has at its base up to 50 cm of matrix-
supported small-pebble conglomerate, comprising well-rounded and well-
sorted brown-stained pebbles with a mean diameter of about 2 cm but with
individual pebbles up to about 5 cm long. Pebble lithologies consist of vein-
quartz and quartz arenites for the most part, although one pebble was a single,
well-rounded crystal of orthoclase 1,2 cm long.
The matrix is an arenaceous limestone with abundant glauconite. Some
shell fragments are present in this limestone matrix. The basal conglomerate
passes up into a sequence of arenaceous limestones and calcareous sandstones,
all with a high content of glauconite. As this section is followed up the slope,
exposures become very poor, and only a few of the more resistant calcareous
beds form ledges. At an altitude some 20 m above the small-pebble conglomerate,
a second unconformity, at the base of the overlying Alexandria Formation (of
probable Tertiary age), is reached.
The middle portion of the Igoda Formation is better exposed some 30 m
to the south. In this second section a brown, small-pebble conglomerate,
indistinguishable from that just described, lies at the base of a similar sequence
of well-exposed glauconitic arenaceous limestones and calcareous sandstones.
As in the first section, these strata are fossiliferous, yielding a shelly fauna
dominated by ostreids which litter the surface. At this point, however, the
calcareous sequence has a much reduced thickness of just over 7 m, and the
small-pebble conglomerate overlies a 3-metre-thick sequence of white, poorly
consolidated, unfossiliferous sandstone and white, small-pebble conglomerates
which are in all other respects identical to the brown ones already described.
These three lithological associations are regarded as informal members,
and are known as the ‘white member’, ‘brown, small-pebble conglomerate
member’ and ‘calcareous member’ respectively.
The base of the Alexandria Formation is marked by another conglomerate
consisting of much larger pebbles (up to 30 cm in diameter) mostly of Beaufort
Group sandstone and siltstone. These pebbles are generally less resistant than
those found in the conglomerates of the Igoda Formation. The matrix of the
Alexandria Formation basal conglomerate is a coarse bioclastic limestone
containing fossil gastropods, especially Patella sp. and Conus sp., usually as
moulds. Glauconite is absent to rare. This unit is about 1,5 m thick and is
overlain by 10 m of cross-bedded, well-indurated, coarse bioclastic limestones.
Characteristically, the cross-bedding comprises a single tabular set, with
seaward dips of about 25°. Above this scarp-forming unit is a considerable
thickness (at least 30 m) of poorly consolidated calcareous aeolian sands.
ANNALS OF THE SOUTH AFRICAN MUSEUM
74
C] ‘Sty Osye 99g) “y7eW J9}eM YsIY A[oJeUITXOIdde ‘[OAo] 19}eM UOOSR] SAO SOIJOU UI OB
sopnyn[y “(MOjaq) suOT}OES S1Yde.ASIVvI}S pue (QAOQK) 1X9} Ul Poqiiosop Ssdoidjno JO UOTINGIAWsSIP SUIMOYS ‘UOCOSL] BPOST JO yURQ UID}SOAA “7 “BIT
~ 330) JOQGUIBUWI BLIWUM
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“| -—uod uMojq
sequiow
snoasesjeo
NOILVINHOA4 VdODOI
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NOILVINHOS
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UPPER CRETACEOUS SEDIMENTS FROM THE IGODA RIVER MOUTH 75
A second section was measured, 500 m closer to the sea (section C). Here
the white member is no longer present, and the calcareous member is even
further reduced in thickness (5,5 m). In other respects the sequence is similar
(see Fig. 2).
THE ROAD SECTION
The original outcrop described by Mountain (1974, see above) was identi-
fied, and the adjective ‘tiny’ found to be very appropriate. Only a few tens of
centimetres of very weathered glauconitic arenaceous limestone can be seen
overlying Beaufort Group sandstones and merging upwards into soil and surface
debris in the bank at the inland side of the road. The outcrop is very overgrown
and easily overlooked. Broken specimens of Rhynchostreon decussata and
Lopha spp., common forms in the main outcrop area, establish a correlation
with the lagoon-side exposures.
At the time of the most recent visit (May 1977), roadwork was in progress
for the straightening and improving of the main coastal road. Where this new
road reaches the approximate altitude of Mountain’s outcrop, it runs about
100 m inland of the latter, and cuts through a mass of dolerite. At the top of
the cutting at this point, a few rounded pebbles of Beaufort Group sandstone
in a calcareous cement were found. The size of these pebbles (20-30 cm) suggests
that they are remnants of the conglomerate from the base of the Alexandria
Formation. This would imply that the Igoda Formation pinches out at this
point (area A on map, Fig. 1).
FAUNA AND AGE OF THE IGODA FORMATION
In comparison with the Cretaceous sediments of Zululand and Transkei
(Kennedy & Klinger 1975 onwards), the invertebrate fauna of the Igoda Forma-
tion is meagre and poorly preserved. Bivalves are the commonest group,
especially the ostreid forms, which have been preserved by virtue of their
unique shell mineralogy, followed by brachiopods, baculitid ammonites,
echinoids, and rare, normally coiled, ammonites, in that order of abundance.
At present, the material in the authors’ collections is too scant to merit
formal description and discussion, but a preliminary examination of the fauna
has revealed the presence of the following faunal elements:
FORAMINIFERA
Textularia sp.
COELENTERATA
“Caryophyllia’ cf. arcotensis Forbes (Fig. 3)
ECHINODERMATA
Unidentifiable cidarid with uniserial pore pairs (Fig. 4B)
Crinoid stem ossicles of Pentacrinus type (Fig. 4A)
76 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 3. ‘Caryophyllia’ arcotensis Forbes (Geology Department, Rhodes University.)
A-B. Lateral view. Scale bar 0,5 cm long. C. Dorsal view, stereopair. Scale bar 0,5 cm long.
UPPER CRETACEOUS SEDIMENTS FROM THE IGODA RIVER MOUTH tif
Fig. 4. A. Crinoid stem ossicle of the /socrinus type. B. Cidaris sp. indet.
Scale bars 0,5 cm long.
MOLLUSCA—CEPHALOPODA
Baculites subanceps Haughton (Fig. 5)
Eupachydiscus ? sp. (Fig. 6)
Pachydiscid sp. indet (compressed)
Saghalinites sp. cf. S. cala (Forbes) (Fig. 7A—B)
MOLLUSCA—GASTROPODA
Turritella (Zaria) cf. T. (Z.) besairiei Basse
MOLLUSCA-BIVALVIA
Rhynchostreon decussata (Goldfuss)
Lopha (Actinostreon) schnaebelei Basse (Fig. 7C—H)
‘Trigonia sp.
Spondylus douvillei Basse
Panopea cf. orientalis (Forbes)
‘Inoceramus’ spp.
BRYOZOA
cf. Ceriopora micropora Goldfuss
“Membranipora’ cf. plebicola Brydone
BRACHIOPODA
cf. Terebratulina relicta Stoliczka
cf. Terebratula manuaensis Muir-Wood
cf. Rhynchonella natuans Stoliczka
Eolacazella affine (Bosquet)
78
ANNALS OF THE SOUTH AFRICAN MUSEUM
J K
Fig. 5. Baculites subanceps Haughton. A, C. SAM-PCI5721. x 1,2.
B. SAM-PCI5723. « 1,0. D. SAM-—PCI5728. « 1,0. E-H. SAM-—PCI5906.
x 1,0. I-L. SAM-PCI5720. x 1,0.
UPPER CRETACEOUS SEDIMENTS FROM THE IGODA RIVER MOUTH 79
Fig. 6. Eupachydiscus ? sp. indet. SAM-—PCI5719. x 1,0.
Of the species identified, only a few can be used for accurate dating of the
Igoda Formation.
Baculites subanceps s.s. has been firmly dated as Late Campanian in
Angola (Howarth 1965), whilst the Pacific subspecies B. anceps pacificus occurs
in the Late Campanian of Japan and California. Saghalinites ranges from the
Santonian to Maastrichtian stages of the Late Cretaceous (Kennedy & Klinger
1977). S. cala, which the Igoda specimen resembles most, ranges from Cam-
panian IV to Maastrichtian II (sensu Kennedy & Klinger 1975) in Zululand but
is also known to occur in slightly older sediments in Pondoland (Transkei)
80 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 7. A-B. Saghalinites sp. cf. S. cala (Forbes). C-H. Lopha (Actinostreon) schnaebelei
(Basse). C. SAM-PCI5773. x 1,0. D. SAM-PCI5749. x 1,0. E. SAM-PCI5736. x 1,0.
F. SAM-PCI5750. x 1,0. G. SAM-PCI5752. x 1,0. H. SAM-PCI5744. x 1,0.
UPPER CRETACEOUS SEDIMENTS FROM THE IGODA RIVER MOUTH 81
(Middle Santonian—Lower Campanian) and southern India (Santonian or
Campanian). The ammonite genus Eupachydiscus is typically Late Cretaceous,
ranging from Coniacian to Maastrichtian, but the Igoda material is too poorly
preserved for specific identification.
Ceriopora micropora Goldfuss has been described from the type locality
of the Maastrichtian stage, but is also common in the Santonian and Campanian.
M. plebicola is from the Maastrichtian.
Eolacazella affine (Bosquet) occurs in the Maastrichtian of Western Europe,
and was also found at the Lower Needs Camp Quarry now dated as Upper
Campanian/Lower Maastrichtian by McGowran & Moore (1971).
These data all seem to suggest an age of Campanian/Maastrichtian for the
Igoda Formation with only a slight possibility of Santonian elements present.
COMPARISON WITH OTHER AREAS IN SOUTHERN AFRICA
The nearest onshore sediments of similar age occur at the Lower or Eastern
Quarry at Needs Camp. Eolacazella affine occurs at both the Igoda River Mouth
and at Lower Needs Camp, thus suggesting the two outcrops to be temporal
equivalents, though of different lithologies. In places, the outcrops at Lower
Needs Camp consist virtually of bryozoan limestone only. Exposures of similar
lithologies have since been found by one of the authors (H. C. K.) in roadside
excavations approximately half-way between the Upper and Lower Quarries,
illustrating that the Lower Needs Camp lithology has a much greater aerial
extent than previously suspected, and may yet prove to be a mapable unit to
conform with the requisites for formal recognition as a separate Formation by
the South African Committee for Stratigraphy.
The Lower Needs Camp sediments probably grade laterally into the slightly
deeper water facies of the Igoda Formation, but outcrops connecting the two
areas are, as yet, unknown.
Along the east coast of southern Africa, the closest Senonian sediments
occur at the Umzamba Estuary (Transkei), which have been firmly dated as
Mid-Santonian to Lower Campanian (Klinger & Kennedy 1977). Rare Saghali-
nites cala in the Umzamba Formation are comparable with Saghalinites sp. in
the Igoda Formation.
The closest, and thus far only known, onshore Upper Campanian/Lower
Maastrichtian sediments on the east coast of southern Africa occur in the
False Bay/St Lucia region of Zululand (Kennedy & Klinger 1975). Here Saghali-
nites cala occurs quite frequently with Nostoceras sp. and Eubaculites sp. in the
Upper Campanian and Lower Maastrichtian respectively. S. cala appears to
be the only identifiable faunal element in common between the two areas.
Lower Maastrichtian sediments are known from Cheringoma further north in
Mozambique, but to date only poorly-preserved Eubaculites species are known
(Crick 1923).
Much further north, the Campanian and Maastrichtian faunas of Mainti-
rano, Madagascar, as described by Basse (1931), strongly resemble those from
82 ANNALS OF THE SOUTH AFRICAN MUSEUM
Igoda especially as far as the thick-shelled ostreid forms are concerned, and to
a lesser extent with the Late Senonian fauna of the east coast Province of
Vatomandry described by Cottreau (1922), which is characterized by the paucity
of ammonites, similar to the Igoda Formation.
Towards the southern and western coast, the closest sediments of com-
parable age are found in the off-shore Alphard Group between Cape St Francis
and Cape Recife (Klinger ef a/. in prep.). Here, however, the dominant faunal
element is Eubaculites latecarinatus indicative of Early Maastrichtian age.
No macrofossils are common to both outcrops.
From here westwards, the first known Cretaceous on-shore sediments occur
near Bogenfels (South West Africa) (Klinger 1977), but these have been dated
definitely as Cenomanian, although some doubt still exists as to whether
Senonian strata are present on-shore or not.
The closest comparable deposits occur at Carimba in Angola, the type area
of Baculites subanceps, the latter being the only faunal element in common
between Igoda and Angola. The Cretaceous sediments at Carimba have been
firmly dated as Late Campanian by Howarth (1965).
SUMMARY
The name Igoda Formation is proposed for a sedimentary sequence
consisting mainly of calcareous sandstones and arenaceous limestones exposed
on the bluffs overlooking the Igoda River Mouth. The Formation is informally
divided into ‘white member’, ‘brown, small-pebble conglomerate member’ and
‘calcareous member’.
In comparison with the Cretaceous sediments of Natal and Transkei, the
Igoda fauna is meagre, consisting mainly of thick-shelled ostreid bivalves,
followed by brachiopods, baculitid ammonites, echinoids, and rare, normally
coiled ammonites, in that order of abundance.
These faunas point to an age of Late Campanian to Early Maastrichtian.
Due to the paucity of the faunas, comparisons with other temporally
equivalent areas are tenuous. The Lower Needs Camp Quarry deposits are of
similar age, and probably represent a shallower water and more restricted lateral
facies equivalent of the Igoda Formation. Saghalinites sp. cf. cala connects
the Igoda Formation to the Cretaceous deposits on the east coast of southern
Africa at the Umzamba Estuary and in the False Bay/Lake St Lucia region of
Zululand.
As far as the abundance of thick-shelled ostreids and relative scarcity of
ammonites is concerned, the Cretaceous deposits of Maintirano and Vato-
mandry in Madagascar show greatest affinity, but this should probably be
ascribed to the depositional environment rather than to more favourable
migration routes.
On the west coast of southern Africa, comparable deposits occur at Carimba
in Angola which provide a definite date for Baculites subanceps, i.e. Late
Campanian.
UPPER CRETACEOUS SEDIMENTS FROM THE IGODA RIVER MOUTH 83
ACKNOWLEDGEMENTS
We wish to express our gratitude towards the following persons for identi-
fying and commenting on some of the material: Mr I. McMillan (SOEKOR,
Johannesburg), Mr D. Salmon (Geological Survey, Cape Town) (Foraminifera),
Prof. Dr E. Voigt (Geologisch-Palaontologisches Institut, Hamburg—Bryozoa),
and Dr E. Owen (British Museum (Natural History), London—Brachiopoda).
Mr L. Matthews collected some of the specimens while a graduate student.
REFERENCES
Basse, E. 1931. Monographie Paléontologie De Crétacé de la Province de Maintirano, Mada-
gascar. Tananarive. Gouv. Gén. Madagascar et Dépend., Service des Mines.
CHAPMAN, F. 1916. Foraminifera and ostracoda from the Upper Cretaceous of Needs Camp,
Buffalo River, Cape Province. Ann. S. Afr. Mus. 12: 107-118.
CoTTREAU, J. 1922. Paléontologie de Madagascar. X. Fossiles crétacés de la cote orientale.
Annls Paléont. 11: 111-192.
Crick, G. C. 1923. Appendix A. On Upper Cretaceous Cephalopoda from Portuguese East
Africa. In: Teale, E. O. The Geology of Portuguese East Africa between the Sambezi and
Sabie Rivers. Trans. geol. Soc. S. Afr. 26: 130-140.
HowartH, M. K. 1965. Cretaceous ammonites and nautiloids from Angola. Bull. Br. Mus.
nat. Hist. (Geol.) 10: 337-412.
KENNEDY, W. J. & KLINGER, H. C. 1975. Cretaceous faunas from Zululand and Natal, South
Africa. Introduction, Stratigraphy. Bull. Br. Mus. nat. Hist. (Geol.) 25: 265-315.
KENNEDY, W. J. & KLINGER, H. C. 1977. Cretaceous faunas from Zululand and Natal, South
Africa. The ammonite family Tetragonitidae Hyatt, 1900. Ann. S. Afr. Mus. 73: 149-197.
Kina, L. C. 1972. Geomorphic significance of the Late Cretaceous limestones at Needs Camp,
near East London. Trans. geol. Soc. S. Afr. 75: 1-3.
K.uincer, H. C. 1977. Cretaceous deposits near Bogenfels, South West Africa. Ann. S. Afr.
Mus. 73: 81-92.
K.uincer, H. C. & KENNEDY, W. J. 1977. Upper Cretaceous ammonites from a borehole near
Richards Bay, South Africa. Ann. S. Afr. Mus. 72: 69-107.
LANG, W. D. 1908. Polyzoa and anthozoa from the Upper Cretaceous limestones of Needs
Camp, Buffalo River. Ann. S. Afr. Mus. 7: 1-11.
Lock, B. E. 1973. Tertiary limestones at Needs Camp, near East London. Trans. geol. Soc.
S. Afr. 76: 1-5.
McGowran, B. & Moore, A. C. 1971. A reptilian tooth and Upper Cretaceous microfossils
from the Lower Quarry at Needs Camp. Trans. geol. Soc. S. Afr. 74: 103-105.
Mountain, E. 1974. The geology of the area around East London, Cape Province. An explanation
of sheet map 3227 D (East London), 3228 (Kei Mouth). Pretoria: Geological Survey.
Woops, H. 1908. Echinoidea, Brachiopoda and Lamellibranchiata from the Upper Cretaceous
limestone of Needs Camp, Buffalo River. Ann. S. Afr. Mus. 7: 13-19.
s¢ - 5
siragst der 6 aie
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Biological Abstracts.
HERBERT CHRISTIAN KLINGER
&
BRIAN E. LOCK
UPPER CRETACEOUS SEDIMENTS FROM THE
IGODA RIVER MOUTH, EAST LONDON,
SOUTH AFRICA
| s- cf ‘ Ca pelow a
VOLUME 77 PART 6 FEBRUARY 1979 . ISSN 0303-2515
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BuLLouGn, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FISCHER, P_-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FiscHer, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gen. 74: 627-634.
Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.
Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
TurEte, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: SCHULTZE, L. Zoologische
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ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 77 Band
February 1979 Februarie
Part. [email protected] » Deel
CRETACEOUS FAUNAS FROM ZULULAND
AND NATAL, SOUTH AFRICA
THE AMMONITE SUPERFAMILY
HAPLOCERATACEAE ZITTEL, 1884
By
WILLIAM JAMES KENNEDY
&
HERBERT CHRISTIAN KLINGER
Cape Town Kaapstad
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CRETACEOUS FAUNAS FROM ZULULAND AND NATAL,
SOUTH AFRICA
THE AMMONITE SUPERFAMILY HAPLOCERATACEAE ZITTEL, 1884
By
WILLIAM JAMES KENNEDY
Geological Collections, University Museum, Oxford
&
HERBERT CHRISTIAN KLINGER
South African Museum, Cape Town
(With 20 figures)
[MS. accepted 18 October 1978]
ABSTRACT
Representatives of the superfamily Haplocerataceae Zittel, 1884, are locally common in
rocks of Barremian age in northern Zululand, where Sanmartinoceras (Sanmartinoceras)
africanum sp. nov. occurs at all stages of development from larval shells to discs which are still
septate at a diameter of 120 mm, and represent the largest known specimens of the genus.
Sanmartinoceras (Theganeceras) nodosum sp. nov., of Lower Aptian age, Sanmartinoceras
(Sinzovia) trautscholdi (Sinzow), and an Aconeceras sp. of Upper Aptian age are other repre-
sentatives of the Aconeceratidae, whilst the Binneyitidae are represented by specimens of
Borissiakoceras of Middle Cenomanian age. In addition to systematic descriptions, the
problems of recognizing dimorphism in these genera are discussed, and certain resultant
taxonomic problems noted, and an annotated list of species referred to the two families is
included.
CONTENTS
PAGE
Introduction . ‘ ‘ ‘ : SD
Location of specimens : 3 : . 86
Field localities . ; é : : - 86
Dimensions of specimens . : : «286
Suture terminology . ‘ ; 5 ~ 3S6
Systematic palaeontology : 87
Annotated list of species referred to the
Aconeceratidae and Binneyitidae . a bY
Acknowledgements. : ; : ~ 1a9
References! \.., 228 aie cri tee » 120
INTRODUCTION
The Haplocerataceae are the longest ranging of the Ammonitina, first
appearing in the Middle Jurassic (Bajocian) and ranging to the Upper Cretaceous
(Coniacian). As Casey has noted, two broad morphologies recur within the
superfamily: firstly, smooth platycones, typified by the Haploceratidae and
Binneyitidae, and secondly, variously ribbed or strigate oxycones with entire or
85
Ann. S. Afr. Mus. 77 (6), 1979: 85-121, 20 figs.
86 ANNALS OF THE SOUTH AFRICAN MUSEUM
crenulate keels, typified by the Oppeliinae, Strigoceratidae and Aconeceratidae
(Casey 1961a: 118). In the Cretaceous of Zululand both groups are represented,
in the Upper Barremian and Aptian the Aconeceratidae are not uncommon at
some levels, whilst the Binneyitidae occur as a great rarity in rocks of Middle
Cenomanian age. Although only a few species are represented in the authors’
collections, they are of some significance, for their specimens of Aconeceratidae
allow, for the first time, the description of the detailed ontogeny of the widely
occurring genus Sanmartinoceras Bonarelli, 1921, as well as clarifying some
aspects of its stratigraphic distribution, whilst the present records of the
binneyitid Borissiakoceras extend the known distribution of the group.
The following species are described below: Aconeceras sp., compared to
A. walshense (Etheridge) ; Sanmartinoceras (Sanmartinoceras) africanum sp. Nov. ;
Sanmartinoceras (Theganeceras) nodosum sp. nov.; Sanmartinoceras (Sinzovia)
trautscholdi (Sinzow); and Borissiakoceras sp.
To this is added an annotated list of genera, subgenera and species referred
to the two families.
LOCATION OF SPECIMENS
The following abbreviations are used to indicate the repositories of the
material studied:
BMNH British Museum (Natural History), London
GSM Institute of Geological Sciences, London
MNHP Muséum d’Histoire Naturelle, Paris
OUM = Oxford University Museum, Oxford
SAM _ South African Museum, Cape Town
SAS South African Geological Survey, Pretoria
FIELD LOCALITIES
Details of localities mentioned in the text are given by Kennedy & Klinger
(1975); fuller descriptions of sections are deposited in the Palaeontology
Department of the British Museum (Natural History), London; Geological
Survey, Pretoria; and South African Museum, Cape Town.
DIMENSIONS OF SPECIMENS
All dimensions given below are in millimetres:
D = diameter, Wb = whorl breadth, Wh = whorl height, U = umbilical
diameter.
Figures in parentheses are dimensions as a percentage of the total diameter.
SUTURE TERMINOLOGY
The suture terminology of Wedekind (1916), recently reviewed and dis-
cussed by Kullman & Wiedmann (1970) is followed here:
Is = Internal lobe with septal lobe, U = Umbilical lobe, L = Lateral lobe,
E = External lobe.
|
,
CRETACEOUS FAUNAS FROM SOUTH AFRICA 87
SYSTEMATIC PALAEONTOLOGY
Phylum MOLLUSCA
Class CEPHALOPODA Cuvier, 1797
Subclass AMMONOIDEA Zittel, 1884
Order AMMONITIDA Zittel, 1884
Suborder AMMONITINA Zittel, 1889
Superfamily HAPLOCERATACEAE Zittel, 1884
Family Aconeceratidae Spath, 1923
Discussion
The Aconeceratidae Spath, 1923, is a group of small to medium-sized,
compressed, involute, commonly oxycone ammonites with flat or convex sides
and a fastigiate to keeled venter. Ornament consists of weak to strong flexuous,
commonly biconvex ribs and growth striae and a keel which may be minutely
crenulate.
The following genera and subgenera have been placed in the family:
Genus Protaconeceras Casey, 1954, Hauterivian
Genus Aconeceras Hyatt, 1903, Barremian to Lower Albian
Genus Sanmartinoceras Bonarelli, 1921 (in Bonarelli & Nagera 1921),
Barremian to Lower Albian
Subgenus Sanmartinoceras s.s., Barremian to Aptian
Subgenus Sinzovia Sazonova, 1958, Aptian
Subgenus Theganeceras Whitehouse, 1926, Lower Aptian
Genus Doridiscus Casey, 1961, Aptian
Genus Nothodiscus Casey in Collignon, 1962, Aptian
Genus Gyaloceras Whitehouse, 1927, Upper Aptian.
Aconeceratids have a wide geographic distribution extending from
Greenland to Antarctica, although their occurrence is sporadic. In Europe at
least, they occur in enormous numbers in some clay facies as in southern
France, where the blue marls of the Fosse Vocontienne yield thousands of
Aconeceras and Casey (19616: 122) regarded them as an open sea group which
invaded the neritic zone only in periods favourable for their growth. As will be
seen below, however, their local abundance in carbonaceous nearshore sand-
stones of the South African Barremian is scarcely compatible with such a
generalization.
The Haplocerataceae as a whole are characterized by striking dimorphism
(Makowski 1962, Callomon 1963, Kennedy & Cobban 1976 with references),
but within the Aconeceratidae our knowledge is such that it remains most
difficult to recognize dimorphic pairs. In Aconeceras itself, the many hundreds
of specimens from the French Aptian examined are all pyritic phragmocones
and it would appear that juveniles, at least, are most difficult to split into
dimorphs, as Palframan (1969) found in some Jurassic haploceratids. The
authors do know, however, of a few specimens referrable to the family in which
88 ANNALS OF THE SOUTH AFRICAN MUSEUM
apertures are preserved: the original material of S. (Sanmartinoceras)
groenlandicum Rosenkrantz, 1934 (in Bogvad & Rosenkrantz), illustrated here
as Figure 1A-C, is an obvious microconch with prominent lappets and a
rostrum. Equally, Gyaloceras smithi Whitehouse, 1927, appears to be a female
or macroconch. The specimen is reproduced here as Figure 2A. In most cases,
however, the shell ornament of aconeceratids (as opposed to apertural form)
gives little clue to dimorphism and a markedly biconvex growth line, indicating
a long rostrum and blunt lappets, occurs, in the present material at least, in both
small and large (if not mature specimens). The authors would, however, suggest
that the presence of a strong spiral depression may indicate the possible presence
of a long lappet (Figs 1A-B, 5C). The topic is returned to below under the
discussion of Sanmartinoceras.
Genus Aconeceras Hyatt, 1903
Type species
Ammonites nisus d’Orbigny, 1841.
Discussion
Aconeceras is represented by a single, poor specimen from the Aptian of
Zululand ; in consequence the reader is referred to Casey’s (19615: 123) extensive
remarks on the genus.
Occurrence
The genus ranges from Barremian to Lower Albian and is widely distributed
from western Europe and the U.S.S.R. to east Africa, Madagascar, Nepal,
Antarctica and eastern Australia.
Aconeceras sp.
Material
One specimen only, BMNH C80002, from the Makatini Formation,
Aptian III, locality 166, Mfongosi Spruit, northern Zululand.
Description
The specimen is an external mould of an oxycone individual with an
original maximum whorl height of over 20 mm. Coiling is very involute with a
small, pit-like steep-sided umbilicus. The whorl section is high, compressed with
the greatest breadth well below mid-flank. The flanks are distinctly flattened and
the venter fastigiate.
The specimen is somewhat worn and there is no obvious trace of ornament
preserved.
Discussion
Although poorly preserved, this specimen is clearly referrable to the genus
Aconeceras. Amongst described species it is perhaps to be compared with the
89
CRETACEOUS FAUNAS FROM SOUTH AFRICA
‘TX = “(pE6I) ZULIYUOSOY wz peaseg jo AdoD “pEeG] ‘zZJUBIYUOSOY WndIpunjuaosd (SpsaIOUIJADULUDG) SDAaIOUIJADULUDS *| “BL{
90 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 2. A. Gyaloceras smithi Whitehouse, 1926. B-D. Aceneceras walshense (Etheridge), 1892.
After Whitehouse (1926, 1927a). x1.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 91
Fig. 3. A-C. Sanmartinoceras (Sanmartinoceras) fontinale (Hudleston), 1890.
D-G. Sanmartinoceras (Sanmartinoceras) olenae (Tenison-Woods), 1883. After Whitehouse
(1927a). x1.
92 ANNALS OF THE SOUTH AFRICAN MUSEUM
F
Fig. 4. Sanmartinoceras (Sanmartinoceras) africanum sp. nov., paratypes.
A. SAS H54/41/e. B-D. BMNH C79988. E-H. BMNH C79982. x1.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 93
Australian Aconeceras walshense (Etheridge) (see Whitehouse 1927a: 114, pl. 16
(figs 2-3); text-figs 1, 6-7) (Fig. 2B—D herein).
Occurrence
Upper Aptian of Zululand.
Genus Sanmartinoceras Bonarelli, 1921 (in Bonarelli & Nagera 1921)
Type species
Sanmartinoceras patagonicum Bonarelli, 1921 (in Bonarelli & Nagera 1921).
Diagnosis
Stout oxycones, initially smooth, with falciform striae and ribs, the latter
weak on the inner flank, sometimes bunched, and strengthening into distinctive
concave ribs on the outer flank. Keel finely denticulate.
Discussion
Sanmartinoceras was originally based upon crushed specimens from the
Aptian of Lago San Martin, Argentina. Leanza (1970, text-fig. 14) (see Fig. 5SA-B
herein) has figured uncrushed topotypes, whilst Howarth (1958) and Thomson
(1974) have figured other material from the sub-Antarctic Islands. Specimens
are illustrated here as Figure SC_D. Specimens of S. patagonicum with apertures
preserved are unknown, but Rosenkrantz (in Bogvad & Rosenkrantz 1934: 20,
pl. 4 (fig. 3), pl. 5 (figs 1-5)) figured a species, S. groenlandicum Rosenkrantz,
with a distinctive rostrum and lappets. These features, indicative of a micro-
conch, have been incorporated into the generic diagnosis. Examination of these
examples shows that the lateral lappets are associated with a distinct spiral
groove (Fig. 1A-B). In the Antarctic specimens (Fig. SC—D) similarly sized
individuals show this groove developed to varying degrees at the same diameter,
and the authors suspect it may prove a criterion for differentiation of immature
males and females at similar diameters, in the type species at least. In the case
of Australian Sanmartinoceras the problem is even more tantalizing. Whitehouse
(1926; 1927a) described four aconeceratids from the Aptian of Walsh River.
Two, Sanmartinoceras fontinale (Hudleston) (Fig. 3D-G, 7A—H) and S. olenae
(Tenison-Woods) (Fig. 3A—C) have strong falcoid ribs and prominent rostra.
The others are very feebly ornamented. Aconeceras walshense (Etheridge)
(Fig. 2B—D) has falcoid growth striae, whilst Gyaloceras smithi Whitehouse
(Fig. 2A) has an inflated whorl and fastigiate venter. 1he authors strongly
Suspect that these species are dimorphs but again cannot resolve the problem
fully.
Within Sanmartinoceras, three subgenera have been recognized. In addition
to Sanmartinoceras sensu stricto, Theganeceras Whitehouse, 1926, was treated
as a subgenus by both Wright (1957) and Casey (19616). Only three species,
S. (T.) falcatum (von Koenen) (Fig. 6I-J) S. (7.) scalatum (von Koenen)
(Fig. 6H) from the Lower Aptian of northern Germany, and S. (7.) grande
94 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 5. Sanmartinoceras (Sanmartinoceras) paragonicum Bonarelli, 1921.
A-B. Topotype, after Leanza (1970). C. BMNH C49055. D. BMNH C49057.
C_D. from Alexander Land. x1.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 95
H 1 J
Fig. 6. A. Sanmartinoceras (Sinzovia) aff. trautscholdi (Sinzow), 1870. B—D. Sanmartinoceras
(Sinzovia) stolleyi Casey, 1961. BMNH C14365 and 14362, from the Upper Aptian of Bekum,
near Hildesheim, Germany. E-—F. Sanmartinoceras (Sinzovia) trautscholdi (Sinzow), 1870.
Copy of Trautschold (1865). G. Sanmartinoceras (Theganeceras) grande Thomson, 1974.
After Thomson (1974). H. Sanmartinoceras (Theganeceras) scalatum von Koenen, 1902.
I-J. Sanmartinoceras (Theganeceras) falcatum von Koenen, 1902. H-J after von Koenen
(1902). All x1.
96 ANNALS OF THE SOUTH AFRICAN MUSEUM
Thomson (Fig. 6G) from the Aptian of Antarctica, have thus far been ascribed
to the subgenus, which Casey (19616: 132) diagnosed as lacking the smooth
juvenile stage of Sanmartinoceras sensu stricto and having finer, more numerous
dense ribs. As is described below, the single Zululand specimen referred to the
subgenus suggests that dimorphism may also present taxonomic problems.
The subgenus Sinzovia Sazonova, 1958, differs from Sanmartinoceras sensu
stricto (according to Casey 19616: 133) in having a very low keel and absence or
poor development of a spiral groove. It differs from Theganeceras in having a
longer juvenile smooth stage and greater tendency to smoothness on the lower
flank. The type species is illustrated here as Figure 6E-—F, related German species
as Figure 6A—D, and the only South African representative, S. (S.) trautscholdi
(Sinzow), as Figure 7I-K.
Occurrence
Sanmartinoceras and its subgenera range from Upper Barremian to Aptian
and are known from Greenland, western Europe, the U.S.S.R., Zululand,
Madagascar, Nepal, Australia, Antarctica, Argentina and Papua.
Subgenus Sanmartinoceras sensu stricto
Sanmartinoceras (Sanmartinoceras) africanum sp. nov.
Figs 4A-H; 8A-F; 9A-I; 10A-F; 11A-B; 12A-C; 13A-F; 14A-—C; 15F-J;
16-17; 19D-E
Sanmartinoceras Kennedy & Klinger 197: 274.
Types
The holotype is SAS H54; paratypes are BMNH C79977-C80001, SAS
H54/4la-d, SAS ZO(i), SAS LJE13la, SAS LJE112, SAS H54/17, SAS
H54/33, all from the Makatini Formation, Barremian I-II, locality 170,
Mlambongwenya Spruit, northern Zululand.
Diagnosis
A large (up to 120 mm diameter phragmocone) species of Sanmartinoceras
in which the inner ‘haft’ of the ribs is narrow, straight and prorsiradiate and the
outer ‘blade’ broad and markedly concave.
Dimensions
D Wb Wh Wb: Wh U
Holotype, SAS H54 —(—) 20,5(—) 42-5(R—) 0,48 —(—)
C79985 we ne 27,3(100) 6,6(24) 14,8(54) 0,45 4,1(15)
C79978 ae .. 120,0(100) 28,8(24) 69,8(58) 0,41 TA
C79984 .. ~—..-—«:115,0(100) 24,0(21) 65,5(57) :0.37-~——«11,3(9, 8)
CRETACEOUS FAUNAS FROM SOUTH AFRICA 97
K
Fig. 7. Sanmartinoceras (Sanmartinoceras) fontinale (Hudleston), 1890. Holotype, BMNH
C5306, Upper Aptian, Primrose Springs, north of Lake Eyre, south Australia. A-D x1;
E-H x2. I-K. Sanmartinoceras (Sinzovia) trautscholdi (Sinzow, 1870). SAM-—PCZ5919. x1.
ANNALS OF THE SOUTH AFRICAN MUSEUM
98
Fig. 8. Sanmartinoceras (Sanmartinoceras) africanum sp. nov.
A-C. Holotype SAS 54; D-F paratype BMNH C79989. x1.
CRETACEOUS FAUNAS FROM SOUTH AFRICA
Fig. 9. Sanmartinoceras (Sanmartinoceras) africanum sp. nov., paratypes.
A-D. SAS H54/416. E-H. SAS H54/41a. I. BMNH C79996. All x2.
99
100 ANNALS OF THE SOUTH AFRICAN MUSEUM
C D E F
Fig. 10. Sanmartinoceras (Sanmartinoceras) africanum sp. nov., paratypes.
A-B. BMNH C€79984. C-F. SAS H54/41c. A-B x1; C-F x2.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 101
Fig. 11. Sanmartinoceras (Sanmartinoceras) africanum sp. noy., paratypes.
A. BMNH C79984. B. SAS H54/41d. A x1; B x2.
102 ANNALS OF THE SOUTH AFRICAN MUSEUM
Description
The barrel-shaped protoconch is succeeded briefly by rather stout whorls,
beyond which, up to a diameter of 0,5-0,8 mm, the shell is moderately involute,
slightly compressed, with flattened flanks and a rounded venter. All the available
specimens are partially or wholly exfoliated, and any ornament, if present, must
have been weak. A few specimens begin to show traces of a ventral ridge, but
not a keel at this diameter, although these features may be an artefact of
preservation.
From 0,8 mm onwards the shell is an oxycone, the coiling is very involute,
with a tiny, crater-like umbilicus. The umbilical wall is flat and subvertical, with
an abruptly rounded shoulder. The whorl section varies markedly; stout
individuals have a whorl breadth to height ratio of 0,6; in slender individuals the
figure is 0,4. Maximum whorl breadth is low on the flanks; the inner flanks are
flattened, the outer flanks convergent; a distinct ventrolateral shoulder is
developed, and there is a sharp keel which, when well preserved, can be seen to
be minutely crenulated (Fig. 9E—-H).
Ornament varies greatly. In some individuals (Figs 11B, 13C—D) there are
only the finest growth striae. These arise at the umbilical seam, but are very
weak. They pass forwards across the inner flank, strengthening as they do so,
and sometimes splitting (Fig. 15G—J). They are at their maximum development
at mid-flank, where they flex gently backwards, giving rise to a clear concavity
which extends across the outer third of the flank (Fig. 13D). Striae decline on
the ventrolateral shoulder and project strongly forwards to meet the keel at an
acute angle. Individual striae correspond to individual crenulations on the keel.
In robust specimens, e.g. BMNH C79983, striae are accompanied by distant,
pronounced ribs, which are strongest at mid-flank; there is every intermediate
between.
Middle growth stages are also characterized by a wide range of variation
(Figs 4E-H, 8A-F). The holotype demonstrates the features of a strongly
ornamented individual. It is wholly septate (Fig. 8B) retaining traces of shell.
Coiling is very involute, with most of the inner whorls being covered. The tiny
umbilicus is deep, with a high, flat, subvertical wall, abruptly rounded shoulder,
and faint umbilical carina. The whorl section is compressed (breadth to height
ratio is 0,48) with faintly concave inner flanks, a weakly inflated mid-flank
region converging to distinct shoulders, clearly demarcated from a high,
septate ventral keel. In none of the medium-sized specimens does the keel bear
serrations. Ornament consists of fine, rectiradiate striae on the inner part of the
flank, effaced at a spiral mid-flank depression. The outer flank bears low, blunt,
concave ribs, declining at the ventrolateral shoulder, but projected forwards to
meet the keel.
The largest available specimens are up to 120 mm in diameter, and are still
wholly septate (Fig. 12A—C). In these, the whorls are quite robust, and ornament
consists of striae on the inner flank with blunt, concave ribs on the outer flank.
None of the present specimens has the aperture preserved. A few fragments show
103
CRETACEOUS FAUNAS FROM SOUTH AFRICA
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104
ANNALS OF THE SOUTH AFRICAN MUSEUM
F
Fig. 13. Sanmartinoceras (Sanmartinoceras) africanum sp. NOV., paratypes.
A-B, F. SAS Zo(i). C. SAS LJE 131a. D. SAS H54/41a. E. BMNH C79979.
A-D, E-F x1; C-D x2.
——
i re
105
CRETACEOUS FAUNAS FROM SOUTH AFRICA
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UOHI[OD J9d00D “YW “ds sosav0yvissiaog “4-C “Z11 ALT SVS edAjered “Aou ‘ds wnuvd1fo (spsavouljavUUDs) Sp4aI0UlJADULUDS *D-Y*pl “BLT
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106 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 15. A-E. Borissiakoceras sp., BMNH C80003. F—-J. Sanmartinoceras (Sanmartinoceras)
africanum sp. nov., paratypes. F is SAS LJE 112; G-J is BMNH C79985.
A-E, G-J x2; F x1.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 107
weak to strong strigations (Figs 4B—D, 13E). Where well preserved, this appears
to be a feature of the dorsal shell layer, and not of the original external shell
surface. That this occurs in some very large specimens suggests the species may
have reached diameters close to 200 mm.
The available specimens show the details of suture development to
advantage, as shown in Figure 16.
Mature sutures are deeply and intricately subdivided, as shown in Figure 17.
Discussion
The specimens show the ontogeny and range of intraspecific variation far
better than in any previously described Sanmartinoceras whilst the specimens
are the largest known for the genus. It is most unfortunate that none of the
specimens retains apertures, but the rather distinct spiral flank depression
present in the holotype suggests that it may be a microconch; the largest
individuals (Figs 1OA—B, 11A, 12A—D) are probably macroconchs.
Thomson (1974: 23) reviewed criteria used to differentiate species of
Sanmartinoceras as follows:
(i) strength and form of ribbing,
(ii) the way in which ribs appear (i.e. abruptly or increasing gradually in
strength),
(iii) the size of the individual at which ribs first appear.
In his discussion, however, he concluded that the form of the falcate ribbing
alone was sufficient to separate species. Figure 18 compares the line of ribs and
striae in the five described species; on this criterion alone it can be seen that the
straight ‘haft’ and form of ‘sickle’ of the present species are quite different from
the markedly biconcave rib of S. olenae, S. fontinale and S. patagonicum.
A straight ‘haft’ characterizes S. groenlandicum (Fig. 18), but here the ‘blade’
is much more deeply concave with the inner half almost straight. As noted
elsewhere, the authors do not regard the presence or absence of a lateral spiral
groove as of specific significance, whilst the present specimens vary greatly in
strength of ornament and growth stage at which it appears. The other feature
which separates S. africanum from other species is its great size, but this may
be no more than an artefact of preservation.
Occurrence
Upper Barremian of Zululand only.
Subgenus Theganeceras Whitehouse, 1927
Theganoceras nodosum sp. nov.
Figs 19A-C, 20
Holotype
SAM-PCZ5708 from the Makatini Formation, Aptian I, locality 170,
Mlambongwenya Spruit, northern Zululand.
108 ANNALS OF THE SOUTH AFRICAN MUSEUM
A jf
/
/
B
4
Vg
/
/
Py
Cc i
itn
“e$
Fig. 16. Suture development in Sanmartinoceras (Sanmartinoceras) africanum sp. nov.
A-C. BMNH C79979. D-E. SAS H54/41A. All x 12,5.
109
CRETACEOUS FAUNAS FROM SOUTH AFRICA
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‘LI ‘31g
110 ANNALS OF THE SOUTH AFRICAN MUSEUM
a ————
i
A B
Vite
Fig. 18. Rib patterns in Sanmartinoceras (Sanmartinoceras) compared.
A. S. (S.) olenae. B-C. S. (S.) fontinale. D. S. (S.) africanum. E. S. (S.) groenlandicum.
F-G. S. (S.) patagonicum.
Diagnosis
A large species of Theganeceras with broad, flat, falcoid ribs on the inner
whorls which bear umbilical bullae and incipient ventral clavi. Outer whorls
ornamented by bullae, flexuous growth lines, folds and constrictions.
Description
The unique holotype of this species comprises a crushed body chamber and
a nucleus. Growth of calcite between shell and sedimentary infilling has
produced an unduly thick ‘shell’ of partly diagenetic origin.
The inner whorl, at a diameter of approximately 38 mm, bears Santos to
sixteen closely spaced, low, broad, sickle-shaped ribs which are straight and
prorsiradiate on the inner flank, flexing backwards at mid-flank and becoming
markedly concave on the outer flank. The ribs are strong and well developed
down to the umbilicus where a distinct bulla is present, whilst on the ventro-
lateral shoulder they strengthen into an incipient clavus. The ribs themselves
and interspaces between bear dense striae, sometimes strengthened into riblets,
giving some ribs a bunched, fasciculate appearance. The venter is fastigiate, and
bears strongly projected striae.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 111
B Cc
Fig. 19. A-C. Sanmartinoceras (Theganeceras) nodosum sp. nov., holotype, SAM-—PCZ5708. x 1.
—E. S. (Sanmartinoceras) africanum sp. nov., paratype, BMNH specimen. x 1.
112 ANNALS OF THE SOUTH AFRICAN MUSEUM
The outer body whorl shows coiling to have been involute with a moderately
large, deep, crater-like umbilicus with a flat, outwards-sloping wall. The whorl
section is compressed, lanceolate, so that the overall shell form was an oxycone.
The whorl breadth to height ratio is estimated as approximately 0,38, the
greatest breadth being close to the angular umbilical shoulder. The preserved
fragment bears seven small, sharp, comma-shaped bullae at the shoulder.
These give rise to bundles of fine, dense, flexuous striae which, being crowded
close to the umbilicus, appear as ribs, which fade out at a short distance from
the bulla. The striae flex forwards to mid-flank, then backwards, to form the
very shallow blade of a sickle. They are strongly projected across the ventro-
lateral shoulder. Paralleling these striae are irregular, low folds and constrictions.
The imperfectly exposed suture is illustrated in Figure 20.
Discussion
The presence of well-developed ribs which extend to the umbilical shoulder
indicates this specimen to be a Theganeceras. It differs from all previously
described species in having broader, less markedly flexed ribs when young, as
well as possessing umbilical bullae, which feature is unique for the subgenus.
Only S. (Theganeceras) grande Thomson (1974, pl. 4 (fig. 9)) approaches this
specimen in size; the unique holotype of that species is quite distinct, however,
lacking bullae and having strongly flexed ribs and striae (Fig. 6G).
Equally, no S. (Sinzovia) bears bullae, and the closest species, S. (S.) stolleyi
Casey (Fig. 6A—D), has ribs which efface on the inner flank. The presence of
bullae also distinguishes our specimen from described S. (Sanmartinoceras) at
comparable dimensions.
Occurrence
Lower Aptian I of northern Zululand only.
Subgenus Sinzovia Sazonova, 1958
Sanmartinoceras (Sinzovia) trautscholdi (Sinzow)
Figs 4I-K, 6E-F
Ammonites bicurvatus Trautschold (non Michelin), 1865: 22, pl. 3 (figs 17a-c).
Oppellia Trautscholdi Sinzow, 1870: 118, pl. A (figs 1, la—b only). -
Sinzovia trautscholdi (Sinzow): Sazonova, 1958: 128, pl. 6 (fig. 2), pl. 8 (figs 1, 1a, 3-7),
pl. 10 (figs 4-5) (with synonymy). Druschchitz & Kudryavtseva, 1960: pl. 42 (figs 7a—b).
Collignon 1962: 31, pl. 229 (fig. 974).
? Sanmartinoceras (Sinzovia) sp. cf. trautscholdi (Sinzow): Casey 1961b: 136, pl. 26 (fig. 7).
Holotype
Trautschold’s original of his plate 3 (fig. 17a-c), from the Lower Aptian of
Simbirsk (now Polivna) in the Stalingrad area of Russia, on which Sinzow
(1870: 118) based his species. The original figures are reproduced here as
Figure 6E-F.
113
CRETACEOUS FAUNAS FROM SOUTH AFRICA
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114 ANNALS OF THE SOUTH AFRICAN MUSEUM
Material
SAM-PCZ5919 from the Makatini Formation, Aptian IV at locality 152,
Mkuze Game Reserve, northern Zululand.
Description
The specimen is a beautifully preserved disc of which the last half of a
whorl is body chamber. It retains extensive areas of replaced shell, in consequence
of which the sutures are not fully decipherable. The dimensions are as follows:
D Wb Wh Wb:Wh U
43,0 10,0(23,3) 22,0(51,2) 0,45 —-9,0(20,9)
Coiling is very involute, oxycone, with two-thirds of the previous whorl
being covered. The umbilicus is small and shallow (20° of diameter), with a
flat, subvertical wall. The umbilical shoulder is sharp and angular. The whorl
section is very compressed, with a breadth to height ratio of 0,45. The inner
flank is flattened to concave, with gently rounded, convergent outer flanks,
converging to an acute, delicately keeled venter.
The inner, concave part of the flank is virtually smooth where shell is
present, due to weathering. On the mould, however, there are delicate, concave,
prorsiradiate, bunched striae which sweep forwards to a distinct spiral ridge at
mid-flank and a parallel groove external to this. On the ridge and groove, the
striae flex backwards in a marked convex projection. On the outer flank, they
flex backwards to form a ‘blade’ corresponding to the bunches of ‘handles’ to
what is an overall sickle-like ornament. The blades are strong, concave, blunt
ribs, strong over most of the last whorl, although effacing on the venter, which
is smooth on either side of a narrow, sharp ventral keel.
The sutures cannot be seen.
Discussion
On the basis of weakness of inner flank ornament, and strength and style
of ribs on the outer flank, the authors would refer their single specimen to
Sinzow’s species. It is particularly close to the specimen from Ulyanovsk,
Russia, illustrated by Casey (19616, text-fig. 143 (1a-b)).
Sinzovia stolleyi Casey (19616, text-fig. 143(g)) is more distantly and
narrowly ribbed, as can be seen from Figure 6A—D herein.
Sinzovia aptianum (Sarasin) (1893: 155, pls 4-6 (fig. 12a—c); Casey 19615:
134, pl. 26 (fig. 6a—b)) has blunter, fewer ribs on the outer flank.
Sanmartinoceras ? (Sinzovia?) sp. nov. of Casey (19616: 136, text-fig. 43(f))
is a Lower Albian species of uncertain affinity.
Occurrence
Lower Aptian of the U.S.S.R. and southern England; Upper Aptian of
Madagascar and Zululand.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 115
Family Binneyitidae Reeside, 1927
The Binneyitidae are a family of micromorphs, characteristically platycone,
with much reduced sutures. Two genera, Borissiakoceras Arkhangelsky, 1916,
and Binneyites Reeside, 1927, were referred to the family by Wright (1957) who
suggested derivation from the Acanthocerataceae, but subsequent work has
revealed the presence of passage forms linking the family to the Aconeceratidae
via Falciferella Casey, 1954. This Middle to Upper Albian genus has a vestigal
keel when young and can be linked to the platycone but still feebly carinate
Aptian aconeceratids Doridiscus Casey, 1961, and Nothodiscus Casey, 1962
(in Collignon 1962). The evolutionary origins of the family thus involved a
change from oxycone to platycone with corresponding loss of keel, and sutural
simplification.
Genus Borissiakoceras Arkhangelsky, 1916
Type species
Borissiakoceras mirabile Arkhangelsky, 1916.
Diagnosis
Small, compressed, moderately involute to moderately evolute platycones.
Flanks typically smooth, sometimes bearing narrow, falcoid ribs. A few species
bear ventrolateral tubercles. Suture simple with narrow, bifid lobes and broader,
bifid or trifid saddles.
Discussion
Borissiakoceras is a distinctive genus, differing obviously from the later
(Coniacian) Binneyites Reeside, 1927, which has stronger lateral and ventro-
lateral ornament, sharp ventrolateral shoulders and more auxiliary elements in
the suture. Johnsonites Cobban, 1961, has a simpler suture and a flat or concave
venter.
Borissiakoceras is clearly descended from Falciferella, species of which
extend to the Upper Albian. It is the only binneyitid in which dimorphism has
been demonstrated (Kennedy & Cobban 1976: pl. 1 (figs 3-4)); males have
stronger, more markedly biconcave growth striae and ribs, reflecting a similar
aperture, with a short rostrum.
Occurrence
Borissiakoceras is best known from North America, ranging from Texas
(Stephenson 1952, 1955) to Kansas and Colorado (Morrow 1935). Other United
States occurrences are summarized by Cobban (1961). It also occurs in British
Columbia (Warren & Stelck 1958) and Alaska (Cobban & Gryc 1961). In these
regions it ranges from Middle Cenomanian to Lower Turonian. The type
species comes from Turkestan and is probably of late Cenomanian age;
116 ANNALS OF THE SOUTH AFRICAN MUSEUM
Bodylevsky & Shulinga (1958) record the genus from the Turonian—Coniacian
of the northern U.S.S.R., and Kennedy & Juignet (1973) record it from the
Middle Cenomanian of Normandy, whilst the present authors have seen a
fragment from the Lower Cenomanian of Sarthe (Sorbonne collections).
Wright (1963) records a doubtful species from the Middle Cenomanian of
northern Australia, and the Engonoceratidae gen. et sp. nov. of Henderson
(1973: 106, pl. 14 (fig. 8)) belongs here, as do the Middle Cenomanian specimens
from Zululand described below.
Borissiakoceras sp.
Figs 14D-F, 15A-E
Material
Two specimens, BMNH C80003, and no. 16 in M. R. Cooper’s collection,
University of Natal, Durban, both from the Middle Cenomanian (Cenomanian
III) Locality 62, the Skoenberg, Zululand.
Description
The specimens comprise a small phragmocone and a body chamber with a
maximum whorl height of 7,4 mm. Coiling is involute with a small, shallow
umbilicus comprising approximately 20 per cent of the diameter. The overall
form is platycone, the whorl breadth to height ratio being approximately 0,5,
with a low umbilical wall, flattened inner, and slightly convergent outer flanks,
abruptly and narrowly rounded ventrolateral shoulder, and somewhat flattened
venter. Both specimens are corroded, so that no trace of any original ornament
remains. The suture is poorly exposed, but much simplified, with such elements
as are visible resembling those of B. mirabile.
Discussion
Overall shell form and suture indicate these specimens to be Borissiakoceras,
the first representatives of the genus, and indeed the Binneyitidae, to be described
from Africa. Because of poor preservation it is not possible to identify them
fully. They most closely resemble feebly ornamented species such as B. mirabile
(see Kennedy & Juignet 1973) and B. orbiculatum (see Cobban, 1961: 750,
pl. 88 (figs 15-41), text-figs 5a-f).
Occurrence
Middle Cenomanian of Zululand.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 117
ANNOTATED LIST OF SPECIES REFERRED TO THE
ACONECERATIDAE AND BINNEYITIDAE
Family Aconeceratidae Spath, 1923
Genus Protaconeceras Casey, 1954
Type species: Oppelia patagoniensis Favre, 1908, by original designation.
Protaconeceras patagoniense (Favre), 1908: 634, pl. 34 (fig. 7), pl. 37 (figs 3-5), text-fig. 6.
Lower Hauterivian of Patagonia.
Protaconeceras spp. nov. Casey, 1954: 270, pl. 7 (fig. 7), text-fig. 2. Upper Hauterivian,
England.
Genus Aconeceras Hyatt, 1903
(= Adolphia Stolley, 1907; Adolphites Hennig, 1932)
Type species: Ammonites nisus d’Orbigny, 1841, by monotypy.
Aconeceras nisus (d’Orbigny), 1841: 184, pl. 55 (figs 7-9). Casey 19616: 128. Upper Aptian of
western Europe, notably Gargasian clays of the Vocontian Trough. Also recorded from
Madagascar and elsewhere, although many records are dubious.
Aconeceras haugi (Sarasin), 1893: 156, pls 4—6, text-fig. 1la-c. Lower Aptian of western
Europe, Nepal (Bordet ef a/. 1971), and Madagascar (Collignon 1962) where it is said to
be of Upper Aptian age.
Aconeceras neonisoides Casey, 19616: 129, pl. 26 (figs 1, 9-10), text-fig. 41d—e. Lower Albian
of southern England and north Africa (Sornay 1955; Dubourdieu 1956).
Aconeceras australonisoides Brunnschweiler, 1959: 11, pl. 1 (fig. la—b). Aptian of western
Australia.
Aconeceras whitehousei Brunnschweiler, 1959: 12, pl. 1 (fig. 2a—b). Aptian of western Australia.
Aconeceras walshense (Etheridge), 1892: 493, pl. 42 (figs 10-11). Whitehouse 1926: 203, pl. 34
(fig. 1), pl. 37 (fig. 3); 1927a@: 114, pl. 16 (figs 2-3), text-figs 1, 6-7. Aptian of Queensland.
Aconeceras nisoides (Sarasin), 1893: 155, pls 4-6 (fig. 10a—c), text-figs 3, 5. Lower Aptian of
western Europe.
Aconeceras luppovie (Sazonova), 1958: 130, pl. 8 (fig. 2). Aptian of the U.S.S.R.
Aconeceras saratoviensis (Sazonova), 1958: 130 (= Oppelia trautscholdi Sinzow, 1898, pl. A
(figs 4-5 only)). Lower Aptian of the U.S.S.R.
Genus Sanmartinoceras Bonarelli, 1921
Type species: Sanmartinoceras patagonicum Bonarelli, 1921, by monotypy.
Sanmartinoceras (Sanmartinoceras) patagonicum Bonarelli, 1921, in Bonarelli & Nagera 1921:
27, pl. 5 (figs 3-6). See also Howarth 1958: 5, pl. 1 (figs 6-10); Leanza 1970: 215, fig. 14;
Thomson 1974: 24, pl. 4b-f, text-fig. 7a. Aptian of Argentina and Antarctica. (See
Fig. 6A—-D herein.)
Sanmartinoceras (Sanmartinoceras) groenlandium Rosenkrantz, 1934, in Bogvad & Rosen-
a 1934: 20, pl. 4 (fig. 3), pl. 5 (figs 1-5). Aptian of east Greenland. (See Fig. 1A—C
erein.)
Sanmartinoceras (Sanmartinoceras) olenae (Tenison-Woods), 1883: 150, pl. 7 (fig. 8), pl. 8
(fig. 1). Whitehouse 1926: 205, pl. 41 (fig. 3); 1927a: 117, pl. 17 (fig. 6), text-figs 3, 4, 9.
Aptian of Australia. (See Fig. 3A—C herein.)
Sanmartinoceras (Sanmartinoceras) fontinale (Hudleston), 1890: 241, pl. 9 (fig. 1). Whitehouse,
1927a: 116, pl. 17 (figs 2-5). Aptian of Australia. (See Figs 3D-G, 7A-H herein.)
Sanmartinoceras (Sanmartinoceras) africanum Kennedy & Klinger, 1978 sp. nov. (See p. 96.)
Upper Barremian of Zululand.
Subgenus Sinzovia Sazonova, 1958
Type species: Ammonites trautscholdi Sinzow, 1870 (= Ammonites bicurvatus Trautschold,
1865 non Michelin, 1838) by original designation.
Sanmartinoceras (Sinzovia) trautscholdi (Sinzow), 1870: 118-119. See also Ammonites bicurvatus
Trautschold, 1865: 22, pl. 3 (fig. 17a—-c) (non Michelin); Casey, 19615: text-fig. 135a—c.
Lower Aptian of the U.S.S.R. and southern England; Upper? Aptian of Madagascar;
Upper Aptian of Zululand.
118 ANNALS OF THE SOUTH AFRICAN MUSEUM
Samartinoceras (Sinzovia) aptianum (Sarasin), 1893: 155, pls 4-6, fig. 12a—c. Casey 19616:
134, pl. 26 (fig. 6a—b), text-fig. 43d—e. Aptian of France, southern England and elsewhere
in Europe; Nepal (Bordet et a/. 1971). (See Fig. 6E—F herein.)
Sanmartinoceras (Sinzovia) stolleyi Casey, 19616: 133, 136; text-fig. 135g—h. Upper Aptian of
Germany. (See Fig. 6A—D herein.)
Sanmartinoceras ? (Sinzovia ?) sp. nov. Casey, 19616: 136, text-fig. 43f. Aptian of England.
vnon Sinzovia luppovie Sazonova, 1958: 130, pl. 8 (fig. 2) = S. luppovi Casey, 19616: 130.
Lower Aptian of the U.S.S.R. An Aconeceras according to Casey (19616: 133).
?non Sinzovia saratoviensis Sazonova, 1958 (= Oppelia trautscholdi Sinzow, 1898, pl. A
(figs 4-5a only). Lower Aptian of the U.S.S.R. Also an Aconeceras according to Casey
(19616: 133).
Subgenus Theganeceras Whitehouse, 1926
Type species: Oppelia scalata von Koenen, 1902, by original designation.
Sanmartinoceras (Theganeceras) grande Thomson, 1974: 25, pl. 4g. Lower Aptian of Alexander
Island. (See Fig. 6G herein.)
Sanmartinoceras (Theganeceras) scalatum (von Koenen), 1902: 54, pl. 45 (fig. 6). Lower Aptian
of north Germany. (See Fig. 6H herein.)
Sanmartinoceras (Theganeceras) falcatum (yon Koenen), 1902: 48, pl. 45 (figs 7-8). See also
Casey 19616: 132, pl. 26 (fig. 2). Lower Aptian of north Germany and England. (See
fig. 6I-J.)
Sanmartinoceras (Theganeceras) (?) sp. Thomson, 1974: 26, pl. 4k. Lower Aptian of Alexander
Island.
Sanmartinoceras (Theganeceras) nodosum Kennedy & Klinger sp. nov. (See p. 107.) Lower
Aptian of Zululand.
Genus Gyaloceras Whitehouse, 1927
Type species: Gyaloceras smithi Whitehouse, 1927 by original designation.
Gyaloceras smithi Whitehouse, 1927a: 115, pl. 17 (fig. 1), text-fig. 8. Aptian of Australia.
(See Fig. 2A.)
Gyaloceras ibo Reyment, 1955: 15, pl. 2 (figs 1-3), text-figs 3-4. Upper Albian of Nigeria.
Casey (19616: 139) has suggested that this species is ‘a completely new development whose
relationship to the Aconeceratidae is doubtful. Possibly it is congeneric with the
Aconeceras ? described by Haas (1942: 165) from the Upper Albian of Angola’.
? Genus Eofalciferella Brunnschweiler, 1959
Type species: Eofalciferella condoni Brunnschweiler, 1959, by original designation.
Eofalciferella condoni Brunnschweiler, 1959: 13, pl. 1 (figs. 3-4). Aptian of Australia. The only
illustrations of this species are pencil sketches of the unique holotype, a crushed specimen
from the Windalia Radiolarite. The species and genus are best treated as nomen dubia;
Casey (19615: 131) implies in his discussion of the genus that its affinities may be with
Sanmartinoceras.
Family Binneyitidae Reeside, 1927
Genus Falciferella Casey, 1954
Type species: Falciferella milbournei Casey, 1954, by original designation.
Falciferella milbourni Casey, 1954: 274, pl. 7 (figs 1-5), text-fig. 3. Middle Albian of southern
England.
Falciferella malandiandrensis Collignon, 1962: 32, pl. 229 (fig. 975). Upper Aptian of
Madagascar. This species is keeled to a diameter of approximately 45 mm. It may be an
Aconeceras, although Collignon (1962) indicates that it has a Falciferella-like suture.
Genus Doridiscus Casey, 1961
Type species: Doridiscus rotulus Casey, 1961, by original designation.
Doridiscus rotulus Casey, 1961b: 139, pl. 26 (fig. 8a—b), text-fig. 44c-e. Low Upper Aptian of
southern England.
Doridiscus sp. nov. indet? Casey, 19615: 140, text-fig. 44f. Upper Lower Aptian of southern
England.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 119
Genus Nothodiscus Casey in Collignon, 1962
Type species: Nothodiscus planus Casey, in Collignon, 1962, by original designation.
Nothodiscus planus Casey in Collignon, 1962: 32, pl. 229 (fig. 976). Upper Aptian of Madagascar.
Genus Borissiakoceras Arkhangelsky, 1916
Type species: Borissiakoceras mirabile Arkangelsky, 1916: 55, pl. 8 (figs 2-3), Lower Turonian
of Turkestan. Kennedy & Juignet, 1973: 900, text-figs 1-2, Middle Cenomanian of France.
Borissiakoceras compressum Cobban, 1961: 747, pl. 87 (figs 19-33); pl. 89 (figs 1-9), text-
fig. 4a—k. Middle Cenomanian of the United States Western Interior.
Borissiakoceras reesidei Morrow, 1935: 463, pl. 49 (fig. 7a—b), pl. 50 (fig. 5), text-fig. 8.
Cobban, 1961: 749, pl. 88 (figs 1-14), text-fig. 3h-k. Middle Cenomanian of the United
States Western Interior.
Borissiakoceras orbiculatum Stephenson, 1955: 64, pl. 6 (figs 1-4). Cobban, 1961: 750, pl. 88
(figs 15-41), text-fig. Sa-f. Middle/Upper Cenomanian of Texas and the United States
Western Interior.
Borissiakoceras cf. B. orbiculatum Stephenson, 1955; Cobban 1961: 753, pl. 89 (figs 10-14),
text-fig Sg, i. Uppermost Cenomanian of the Black Hills, United States Western Interior.
Borissiakoceras inconstans Cobban & Gryc, 1961: 187, pl. 38 (figs 30-37), text-fig. 2i-l.
Latest? Cenomanian of Alaska.
Borissiakoceras ashurkoffae Cobban & Gryc, 1961: 188, pl. 38 (figs 38-43), text-fig. 2j-k, m.
Lower Turonian of Alaska.
Borissiakoceras (?) sp. Wright, 1963: 602, pl. 89 (fig. 5). Middle Cenomanian of Bathurst
Island, Australia.
Borissiakoceras ? sp. Lower Cenomanian of Sarthe, France (Sorbonne collections).
Borissiakoceras sp. Kennedy & Klinger, 1978. (See p. 116.) Middle Cenomanian of Zululand.
Genus Johnsonites Cobban, 1961
Type species: Johnsonites sulcatus Cobban, 1961, by original designation.
Johnsonites sulcatus Cobban, 1961: 743, pl. 87 (figs 1-18), text-fig. 3a—g. Middle Cenomanian
of Wyoming and Colorado in the United States Western Interior.
? Johnsonites sp., the original of Stephenson’s (1952: 198, pl. 45 (figs 5—6)) Euhoplites ? sp.
from the Middle Cenomanian of Texas may also belong to this genus.
Genus Binneyites Reeside, 1927
Type species: Binneyites parkensis Reeside, 1927, by original designation.
Binneyites parkensis Reeside, 1927: 5, pl. 3 (figs 1-10). Cobban, 1961: 754, pl. 89 (figs 32-37),
text-fig. Ss, t. Coniacian of Wyoming and Utah in the United States Western Interior.
Binneyites carlilensis Cobban, 1961: 755, pl. 89 (figs 15-22), text-fig. Sh, j-m. Mid-Turonian
of South Dakota and Wyoming in the United States Western Interior.
Binneyites aplatus (Morrow), 1935: 465, pl. 49 (fig. 5), pl. 50 (fig. 6), text-fig. 7. Mid-Turonian
of Kansas in the United States Western Interior.
Binneyites rugosus Cobban, 1961: 756, pl. 89 (figs 26-31), text-fig. Sn—p.
The Engonoceratidae gen. et sp. nov. of Henderson, 1973: 106, fig. 14 (no. 8), text-fig. 15,
is either a Borissiakoceras or Binneyites of Turonian age.
ACKNOWLEDGEMENTS
We thank Dr H. W. Ball, Dr M. K. Howarth and Mr D. Phillips of the
British Museum (Natural History), Dr R. Casey and Mr C. J. Wood (Institute
of Geological Sciences), Mr C. W. Wright and Dr M. R. Cooper (Oxford), for
their advice and useful discussions. The assistance of the staff of the Geological
Collections at Oxford University Museum, and South African Museum,
Cape Town, is gratefully acknowledged as is the financial support of the Trustees
of the Sir Henry Strakosh Bequest, the Royal Society, and Natural Environment
Research Council.
120 ANNALS OF THE SOUTH AFRICAN MUSEUM
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(In Russian.)
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BorbDetT, P., COLCHEN, L., KRUMMENACHER, D., LE Fort, P., MOUTERDE, R., & Remy, M.
1971. Recherches Géologiques dans Himalaya du Népal, Région de la Thakkhola.
Paris: Centre National de la Recherche Scientifique.
BRUNNSCHWEILER, R. O. 1959. New Aconeceratinae (Ammonoidea) from the Albian and
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CALLOMON, J. H. 1963. Sexual dimorphism in Jurassic ammonites. Trans. Leicester lit. phil. Soc.
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Casey, R. 1954. Falciferella, a new genus of Gault ammonites, with a review of the family
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CosBan, W. A. 1961. The ammonite family Binneyitidae Reeside in the Western Interior of the
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CosBaNn, W. A. & Gryc, G. 1961. Ammonites from the Seabee Formation (Cretaceous) of
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COLLIGNON, M. 1962. Atlas des fossiles carctéristiques de Madagascar (Ammonites) LX (Aptien).
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DrusHcuitz, V. V. & KupryAvtTsevA, M. P. 1960. Atlas of Lower Cretaceous fauna of the
northern Caucasus and Crimea. Moscow: Gostoptekhizdat. (In Russian.)
DusourDieu, G. 1956. Etude géologique de la région de l’OQuenza (Confins Algéro-Tunisiens).
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Favre, F. 1908. Die Ammoniten der unteren Kreide Patagoniens. Neues Jb. Min. Geol.
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HENNIG, E. 1932. Wesen und Wege der Paldontologie. Berlin.
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HUuDLEsTON, W. H. 1890. Further notes on some Mollusca from South Australia. Geol.
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Palaeontology 6: 597-614.
6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
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Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-1SA
Nucula (Leda) bicuspidata og (ome: 37.
Leda plicifera A. Adams, 1856:
Laeda bicuspidata Hanley, 1859: “tis pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
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In describing new species, one specimen must be designated as the holotype; other speci-
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Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.
WILLIAM JAMES KENNEDY
&
HERBERT CHRISTIAN KLINGER
CRETACEOUS FAUNAS FROM ZULULAND
AND NATAL, SOUTH AFRICA —
THE AMMONITE SUPERFAMILY ~
HAPLOCERATACEAE ZITTEL, 1884
,
°- 1} Capele: Feet a
VOLUME 77 PART 7 FEBRUARY 1979 ISSN 0303-2515
MUS. COMP. ZOOLU
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scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part
number (only if independently paged) in parentheses, pagination (first and last pages of article).
Examples (note capitalization and punctuation)
BuLLouGn, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FiscHer, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FiscHer, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. 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.
Turere, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 77 Band
February 1979 February
Part 7 . Deel
A NEW GENUS AND SPECIES OF
THE PENAEOID FAMILY
SOLENOCERIDAE (CRUSTACEA, DECAPODA) FROM
SOUTH-EAST AFRICAN WATERS
By
ANTONIO J. DE FREITAS
Cape Town Kaapstad
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A NEW GENUS AND SPECIES OF THE PENAEOID FAMILY
SOLENOCERIDAE (CRUSTACEA, DECAPODA) FROM
SOUTH-EAST AFRICAN WATERS
By
ANTONIO J. DE FREITAS
Oceanographic Research Institute, Durban
(With | figure and 1 table)
[MS accepted 31 October 1978]
ABSTRACT
A new genus, Cryptopenaeus, is proposed for a new solenocerid species, Cryptopenaeus
catherinae, which is described and illustrated. The new genus is related to Hymenopenaeus and
Haliporoides formerly belonging to the genus Hymenopenaeus, sensu lato. C. catherinae has so
far been found only in three localities off southern Mozambique at depths of 310-500 metres.
CONTENTS
PAGE
Introduction ae ant. Pe
Cryptopenaeus gen. nov. . : : 2 2s
Cryptopenaeus catherinae sp. nov. . a0 125
Acknowledgements . . . ... 130
References..." eae ok! Bee 180
INTRODUCTION
In June 1969, while collecting penaeoid shrimps aboard a trawler operating
in deep water off southern Mozambique, one male specimen representing a new
genus and species was caught in 350 metres of water. Further male specimens
were subsequently found in the same area in June 1973, and the only female
available was caught in September 1976 at a depth of 500 metres.
All specimens were caught together with the commercially important pink
or knife prawn Haliporoides triarthrus, as well as the less common Aristeomorpha
foliacea and Penaeopsis balssi.
Genus Cryptopenaeus gen. nov.
Diagnosis
Body robust; carapace elongate; integument firm. Rostrum short, not
reaching distal margin of first antennular segment; ventral margin moderately
convex; armed only with dorsal teeth; epigastric tooth and first rostral separated
by interval equal to or only slightly greater than that between first and second
rostral teeth. Orbital, suprahepatic and branchiostegal spines absent; antennal,
postorbital, hepatic and pterygostomian spines present; cervical sulcus deep,
long but not reaching mid-dorsum of carapace; hepatic sulcus deep and long,
bending anteroventrally from horizontal posterior part and almost reaching
123
Ann. S. Afr. Mus. 77 (7), 1979: 123-131, 1 fig., 1 table.
124 ANNALS OF THE SOUTH AFRICAN MUSEUM
base of pterygostomian spine; orbito-antennal groove shallow and wide;
branchiocardiac carina distinct but not sharp; mid-dorsal abdominal carina
present on segments two to six. Telson with pair of short fixed spines ; no movable
marginal spines. Prosartema narrow, long, extending beyond end of eye.
Antennular flagella similar, subcylindrical and equal to, or slightly longer than,
carapace. Mandibular palp two jointed; articles subequal in length, distal one
narrower than basal and tapering to rounded apex. Exopodites on all maxillipeds
and pereiopods. Lateral ramus of uropod with very small, blunt distolateral
spine. Petasma with ventrolateral lobule entirely occupied by ventral costa and
distally free from dorsolateral lobule; both ventrolateral and dorsolateral
lobules heavily sclerotinized ; appendix masculina and appendix interna present.
Thelycum simple, of open type. Podobranch on maxilliped II only; epipodites
on maxillipeds II and III and on pereiopods I-IV.
Type species
Cryptopenaeus catherinae sp. nov.
Etymology
The generic name is derived from the prefix crypto, from the Greek kryptos
meaning hidden, in combination with the generic name Penaeus, denoting the
fact that this shrimp has been hidden from science until now; gender masculine.
Taxonomic status and comments
From the works of Bate (1881, 1888), Bouvier (1906), and Burkenroad
(1936), it is clear that the generic complex forming the then accepted subfamily
Solenocerinae presented many taxonomic difficulties. This subfamily consisted
of three genera: Solenocera, shrimps with concave antennular flagella (Lucas
1849; Wood-Mason & Alcock 1891; Barnard 1950); Haliporus, shrimps with
subcylindrical antennular flagella and movable lateral spines on the telson
anterior to fixed pair (Bate 1881; Kensley 1968); and Hymenopenaeus, shrimps
with subcylindrical antennular flagella and lacking lateral spines on the telson
(Smith 1882; Wood-Mason & Alcock 1891; Barnard 1950).
It is apparent from the literature, however, that Burkenroad (1936) was
somewhat unhappy with the taxonomic status of Hymenopenaeus and went to
the point of dividing the genus into four superspecies ‘. . . according to the
presence or absence of branchiostegal or pterygostomian spines and to the
nature of the postrostral armature’. Pérez Farfante (1977) revised the subfamily
and, besides proposing that the subfamilies hitherto accepted should be elevated
to the category of families of the superfamily Penaeoidea, divided Hymeno-
penaeus into five genera partly based on the superspecific groups elaborated by
Burkenroad (1936). In doing so Pérez Farfante takes into consideration the
‘... Shape of the antennular flagella and rostrum, proportions of the carapace,
number and comparative size of the articles of the mandibular palp, presence or
absence of certain carinae on the carapace, relative dimensions of the posterior
A NEW GENUS AND SPECIES OF THE PENAEOID FAMILY SOLENOCERIDAE 125
two pairs of pereiopods, location of the distolateral spine of the lateral ramus
of the uropod, structure of the petasma and degree of development of the
arthrobranchia on somite VII’, as well as those characteristics originally used
by Burkenroad.
The system presented is very comprehensive and covers the previously
known species extremely well. However, whereas the six specimens caught in
southern Mozambique belong clearly to the family Solenoceridae and would
have fitted into the genus Hymenopenaeus sensu lato, they do not belong to any
of the five genera established by Pérez Farfante (1977), and have therefore been
placed in a new genus, Cryptopenaeus.
Table 1 sets out the similarities and differences between Cryptopenaeus and
the other five genera, Hymenopenaeus, Haliporoides, Pleoticus, Hadropenaeus,
and Mesopenaeus. The new genus is closely allied to Hymenopenaeus and
Haliporoides but differs from the former in the arrangement of the rostral
teeth, the absence of branchiostegal spines, the presence of a mid-dorsal carina
on abdominal segments 2 and 3 and also by the short rostrum with a strongly
convex ventral margin. Cryptopenaeus differs from Haliporoides in the arrange-
ment of the rostral teeth, presence of a dorsal carina on the second abdominal
segment, by the absence of a suprahepatic spine and by the short rostrum with a
strongly convex ventral margin. The petasma of Cryptopenaeus differs from that
of the other five genera in having the ventrolateral lobule entirely occupied by
the ventral costa.
Cryptopenaeus catherinae sp. nov.
Fig. 1
Material
Holotype: SAM-A16148 in the South African Museum, Cape Town,
3, 46,7 mm carapace length, caught off Cape Santa Maria in southern Mozam-
bique (26°06’S 33°08’E) at a depth of 350 metres, on 16 December 1969.
Alloptype: SAM-A16149 in the South African Museum, Cape Town,
2, 63,2 mm carapace length, caught off Monte Bello in southern Mozambique
(25°00’S 35°21’E) at a depth of 500 metres, in September 1976.
Paratypes: 4 gg, 44,5-47,7 mm carapace length, caught off southern
Mozambique at a depth of 310 metres, in June 1973. One male paratype is in
the National Museum of Natural History in Washington, D.C., and the
remaining three will be sent to the British Museum (Natural History).
Description
Rostrum. Slightly downwardly directed, reaching to or just beyond end of
first antennular segment; ventral margin convex; rostral teeth 74°; epigastric
and three other teeth situated behind postorbital margin of carapace; adrostral
carina short, just reaching postorbital margin; postrostral carina very well
developed, long, almost reaching posterior margin of carapace and with
conspicuous notch behind epigastric tooth; median groove absent.
ANNALS OF THE SOUTH AFRICAN MUSEUM
126
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A NEW GENUS AND SPECIES OF THE PENAEOID FAMILY SOLENOCERIDAE 127
Carapace. Uniformly glabrous and lightly punctate; gastrofrontal and
postocular grooves and horizontal suture absent; no supra-orbital spine;
cervical sulcus and carina very well developed and long but not reaching dorsal
midline; cervical carina terminating anteroventrally in prominent hepatic spine;
gastro-orbital carina absent; postorbital spine prominent; antennal spine
present but relatively small; antennal carina absent; orbito-antennal groove
restricted to wide depression extending from below postorbital spine to below
hepatic spine; hepatic carina sharp, anteroventrally directed and situated just
anterior and below hepatic spine; hepatic sulcus wide and deep; extending
posteriorly below hepatic spine; branchiocardiac carina distinct but not very
sharp. Branchiostegal spine absent; pterygostomian spine prominent and sharp;
submarginal carina long and sharp; no vertical suture or carina.
Antennule. Flagella subequal in length; about 2,5 times length of antennular
peduncle; subcylindrical; mesial flagella twice as thick as lateral flagella;
prosartema flexible with pointed apex, reaching just beyond distal end of first
antennular peduncle, copiously provided with long setae; stylocerite sharply
pointed distally, straight and reaching just beyond end of eye or to end of first
antennular article; distolateral spine prominent and long; parapenaeid spine
absent.
Scaphocerite. Distolateral spine reaching just beyond distal end of anten-
nular peduncle; apex of lamella extending beyond distolateral spine; basicerite
with single broad blunt tooth distally.
Mandibular palp. Reaching to about basal one-third of carpocerite;
proximal article 1,8 times as long as wide; distal article subequal to proximal,
tapering to rounded apex.
Maxilliped III. Endopodite not exhibiting sexual dimorphism; reaching
distal end of scaphocerite; exopodite short, reaching less than half-way along
merus of endopodites; epipodite present.
Pereiopods. Exopodites present on all pereiopods, well developed, longest
on first periopod and shortest on fifth; epipodites present on pereiopods I-IV;
basipodites of first, second and third with prominent spines; ischial spine
present on first pereiopod only; distinct coxal spine on fifth pereiopod.
Pereiopod IV reaching to apex of mandibular palp; pereiopod V reaching to
distal end of antennular peduncle. Extended laterally lengths of pereiopods in
ascending order are: first, second and fourth, third and fifth.
Abdomen. Uniformly glabrous; mid-dorsal carina present from posterior
half of second segment to end of sixth where it terminates in short spine; short
vertical groove on pleura of first segment; lateral carinae absent.
Telson. Slightly longer than sixth segment; about as long as mesial ramus
of uropod; median groove deep, occupying only anterior half of telson; apical
spine somewhat blunt; pair of inconspicuous, very short, fixed subapical spines
present; movable marginal spines lacking.
Thelycum. Simple open structure; anterior portion formed by vertical
posterior face of sternite between fourth pereiopods; posterior face with short
128 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 1. Cryptopenaeus catherinae sp. noy. A. Lateral view, holotype ¢ 46,7 mm carapace
length. B. Right prosartema. C. Telson. D. Left mandibular palp, ventral view. E. Left
appendix masculina and interna, ventral view. F. Appendix masculina, dorsal view. G. Distal
subquadrate process of right half of petasma. H. Petasma, ventral view. I. Thelycum.
A NEW GENUS AND SPECIES OF THE PENAEOID FAMILY SOLENOCERIDAE 129
median groove and low lateral ridges; covered with short setae and obscured by
coxal protuberances of fourth pereiopods. Posterior portion (between fifth
pereiopods) consisting of elongate plate; broad central ridge occupying slightly
more than anterior two-thirds of plate; well-defined lateral ridges extending
from posterior sternal process to anterior margin of somite; two suboval,
setose, boss-like structures present between anterior third of lateral ridges and
median ridge; anterior margin of posterior sternal process interrupted by deep
median sulcus.
Petasma. Simple and very slightly involuted; dorsomedian lobule about
two-fifths of total length of petasma; entirely united along midline; ventro-
median lobule elongate and subtriangular; inner membranous section folded
slightly on itself; heavily sclerotinized central ridge running length of lobule
terminating distally in thick subquadrate process; distal margin smooth;
proximolateral angle of quadrate process beak-like with four or five blunt teeth;
ventral face of process concave; dorsal face convex; dorsolateral lobule elongate
and subtriangular; apex situated under proximolateral angle of distal ventro-
median process; lower inner angle forms small proximal process; ventral surface
sparsely covered with long setae; ventral costa (occupying entire ventrolateral
lobule) extending along lateral margin of petasma reaching half way into distal
ventromedian process; apex free and bi-lobed; dorsal lobe longer than ventral
lobe; ventral lobe with one to four minute teeth.
Appendix masculina. Dorsally convex, ventrally concave, roughly trapezoid
in shape; distal margin with row of short, stout setae; appendix interna subequal
in length to appendix masculina, half fitting into concave face of latter;
elongate with concave median surface; apex with short stout setae. Basal segment
of endopodite of pleopod II as wide as long, its distolateral portion concave,
subtriangular and produced into long, blunt spur.
Colour in life. Body generally red, carapace with broad white stripe running
from below hepatic sulcus to almost posterior margin of carapace on each side;
this stripe wider in posterior half of carapace; on abdominal segments brighter
red patch running anteroventrally on each pleuron; distinct white longitudinal
stripe on dorsal carina of abdominal segments 4-6; telson and uropods pinkish
white becoming red along posterior edges; lateral margins of scaphocerite red;
rostral crest and pereiopods pinkish white; basal segments of pleopods grey to
white; pleopodal endopodites greyish white becoming bright red distally;
marginal setae of pleura, pleopods and uropods orange while those of scapho-
cerite and antennules white.
Distribution
Known only from the type locality.
Ecological notes
All the specimens were collected from the same general area, namely the
Limpopo Bight in southern Mozambique. This is an important fishing ground
130 ANNALS OF THE SOUTH AFRICAN MUSEUM
where the main species of interest are the spiny lobster, Palinurus delagoae,
caught in about 280-350 metres, the langoustine, Nephrops andamanicus,
trawled in 320-380 metres, and the pink or knife prawn, Haliporoides triarthrus,
found in 300-500 metres.
In this area, the continental slope is relatively gentle down to 500 metres
after which it drops very steeply to about 3 000 metres. The substrate consists
of muddy sand to sandy mud and the temperature at 300-500 metres depth
recorded in April and September 1964 was 11-13°C (Instituto Hidrografico
Lisbon 1965, 1967).
Although Cryptopenaeus catherinae has so far been found only in the type
locality, it seems feasible to expect that, as its associate species, Haliporoides
triarthrus, Aristeomorpha foliacea and Penaeopsis balssi are found off the coast
of Natal, the distribution of C. catherinae could possibly extend southward as
well.
ACKNOWLEDGEMENTS
I wish to thank the Fisheries Development Corporation and the South
African Association for Marine Biological Research for their financial and
administrative assistance; Dr Isabel Pérez Farfante of the Systematics
Laboratory, National Marine Fisheries Service, Washington D.C., U.S.A., for
having checked the identity of this species and for the meticulous way in which
she read and criticized my manuscript; Professor A. E. F. Heydorn, Director of
the Oceanographic Research Institute, Durban, for his encouragement and for
his criticism of the manuscript; Dr P. F. Berry for reading and criticizing the
paper; and finally my wife, after whom this species is named, for her encourage-
ment, patience and understanding.
REFERENCES
BARNARD, K. H. 1950. Descriptive catalogue of South African decapod crustacea. Ann. S. Afr.
Mus. 38: 1-837.
Bate, C. S. 1881. On the Penaeidae. Ann. Mag. nat. Hist. (5) 8: 169-196.
Bate, C. S. 1888. Report on the Crustacea Macrura collected by H.M.S. Challenger during
the years 1873-1876. Rep. Sci. Results Voyage H.M.S. Challenger 1873-76, Zool. 24: 1-942.
Bouvier, E. L. 1906. Observations sur les Peneides du genre Haliporus Bate. Bull. Mus.
Oceanogr., Monaco 81: 1-10.
BURKENROAD, M. D. 1936. The Aristaeinae, Solerocerinae, and pelagic Penaeinae of the
Bingham Oceanographic Collection. Bull. Bingham Oceanogr. Coll. 5(2): 1-151.
INstiITUTO HiprRoGRAFICO LisBon. 1965. Resultados das observagées oceanograficas no Canal
de Mocambique, Cruzeiro AL 1/64: Abril-Maio 1964 1: 1-73.
INstrTuTO HipROGRAFICO LisBoN. 1967. Resultados das observagées oceanograficas no Canal
de Mocgambique, Cruzeiro AL 2/64: Setembro-Outubro 1964 3: 1-107.
KENSLEY, B. F. 1968. Deep sea decapod Crustacea from west of Cape Point, South Africa.
Ann. S. Afr. Mus. 50: 283-323.
Lucas, P. H. 1849. Genus Solenocera Lucas. Rev. Mag. Zool. (2) 1: 300.
PEREZ FARFANTE, I. 1977. American solenocerid shrimps of the genera Hymenopenaeus,
Haliporoides, Pleoticus, Hadropenaeus new genus, and Mesopenaeus new genus. Fish. Bull.
75: 261-346.
A NEW GENUS AND SPECIES OF THE PENAEOID FAMILY SOLENOCERIDAE
131
SmiTH, S. I. 1882. Report on the results of dredging, under the supervision of Alexander
Agassiz, on the East Coast of the United States, during the summer of 1880 by the
U.S. Coast Survey Steamer ‘Blake’. 17: Report on the Crustacea, 1: Decapoda. Bull. Mus.
comp. Zool., Harv. 10: 1-108.
Woop-Mason, J. & Atcock, A. 1891. Natural history notes from H.M. Indian Marine
Survey steamer ‘Investigator’, Commander R. F. Hoskyn, R.N., commanding. No 21.
Note on the results of the last season’s deep-sea dredging. Ann. Mag. nat. Hist. (6) 8:
268-286.
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2? Ath 4. Vi!
6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861:
Leda bicuspidata: Nicklés, 1950: is, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.
In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holotype
SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and date.
7. SPECIAL HOUSE RULES
Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. *... the Figure depicting C. namacolus ...’; *. . . in C. namacolus (Fig. 10)...’
(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene
(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian
Punctuation should be loose, omitting all not strictly necessary
Reference to the author should be expressed in the third person
Roman numerals should be converted to arabic, except when forming part of the title of a
book or article, such as
‘Revision of the Crustacea. Part VIII. The Amphipoda.’
Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.
Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.
ANTONIO J. DE FREITAS
A NEW GENUS AND SPECIES OF
THE PENAEOID FAMILY
SOLENOCERIDAE (CRUSTACEA, DECAPODA) FROM
SOUTH-EAST AFRICAN WATERS
Bf nkleirw
VOLUME 77 PART 8 FEBRUARY 1979 ISSN 0303-2515
[32 A
me fe "Ac Y
=
APR 1 3 1979
( OUTH AFRICAN
Sa a MUSEUM
JAPE TOWN
INSTRUCTIONS TO AUTHORS
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Examples (note capitalization and punctuation)
BuLLouGu, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FiscHer, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FiscHer, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann, Mag. nat. Hist. (13) 2: 309-320.
Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr..Coll. 17 (4): 1-51.
THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.
(continued inside back cover)
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2S 6 ES <= PS es = Ow
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 77 Band
February 1979 Februarie
Part 8 _ Deel
YPE SPECIMENS OF HYDROIDA (COELENTERATA)
IN THE SOUTH AFRICAN MUSEUM
By
N. A. H. MILLARD
Cape Town Kaapstad
The ANNALS OF THE SOUTH AFRICAN MUSEUM
are issued in parts at irregular intervals as material
becomes available
Obtainable from the South African Museum, P.O. Box 61, Cape Town
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TYPE SPECIMENS OF HYDROIDA (COELENTERATA) IN THE
SOUTH AFRICAN MUSEUM
By
N. A. H. MILLARD
South African Museum, Cape Town
[MS. accepted 31 October 1978]
ABSTRACT
A list is given of seventy-nine types of Hydroida housed in the South African Museum,
including spirit material, whole mounts and microscope sections.
CONTENTS
PAGE
Introduction : : : : He yile's
Species list . : Ge gt: : co! 134
References . : : : 2 . 149
INTRODUCTION
This paper lists the type holdings of Hydroida in the South African Museum,
and includes not only material preserved in alcohol, but also whole mounts and
sections prepared for microscopic examination.
The arrangement of families, genera and species is the same as that used by
Millard (1975).
Each specimen has a South African Museum registered number (prefixed
by the letters SAM-H), and in most cases a second number is quoted too, which
is that of the coilector’s catalogue. For the latter the following abbreviations are
used:
UCT University of Cape Town; material from the Ecological Survey of the
intertidal rocky shore, the estuaries and the benthic region of the South
African seas. It should be noted that most of the U.C.T. collection of
hydroids is now housed in the South African Museum.
PF Material collected by the old Government Survey vessel, the R.S. Pieter
Faure. The positions given in the Pieter Faure catalogue were not very
reliable, but have been converted here as accurately as possible into
latitude/longitude positions.
MD _ Duplicate material from the Museum National d’Histoire Naturelle,
Paris, collected during the cruise MD.03 of the Marion-Dufresne
(Millard 1977a).
MT Duplicate material from the collection of the Musée Royal de I’Afrique
Centrale, Tervuren, Belgium, collected by J. Bouillon (Millard &
Bouillon 1973, 1974, 1975).
133
Ann. S. Afr. Mus. 77 (8), 1979: 133-150.
134 ANNALS OF THE SOUTH AFRICAN MUSEUM
SM __— Material collected by the R.V. Meiring Naude off the east coast of South
Africa (Millard 19775).
TME Duplicate material from the Universitetets Zoologiske Museum, Copen-
hagen, collected during Th. Mortensen’s Java—South Africa Expedition,
1929-1930 (Millard 1968).
SPECIES LIST
Suborder ATHECATA
Family Myriothelidae
Monocoryne minor Millard, 1966: 435, fig. 1.
Holotype: SAM-—H417 (UCT SCD 215C). One fertile specimen detached
from substratum.
Locality: Agulhas Bank, 34°03’S 25°58’E; 78 m; 25 November 1960.
Collected by: University of Cape Town.
Mpriothela tentaculata Millard, 1966: 437, fig. 2.
Holotype: SAM-—H418 (UCT WCD 7Q). One male individual on encrusting
bryozoan.
Locality: Off west coast of Cape Peninsula, 34°09,3’S 18°17,5’E; 43 m;
24 March 1959.
Paratype: SAM-H2915 (UCT WCD 7Q). Microscope sections of male
individual; locality as holotype.
Collected by: University of Cape Town.
Family Corynidae
Bicorona elegans Millard, 1966: 441, fig. 3.
Holotype: SAM-H419 (UCT SB 161X). Rich male and female colonies;
plus one whole mount; plus microscope sections.
Locality: Saldanha Bay, 33°02,5’S 18°02’E; littoral; September 1957.
Collected by: University of Cape Town.
This is the type species of the genus Bicorona Millard, 1966.
Family Eudendriidae
Eudendrium deciduum Millard, 1957: 184, fig. 2.
Holotype: SAM-H98 (UCT FAL 52V). Male and female colonies; plus
one whole mount; plus microscope sections.
Locality: False Bay, from 34°09,3’S 18°49,6’E to 34°09’S 18°50,1’E; 18 m;
25 June 1952.
Collected by: University of Cape Town.
TYPE SPECIMENS OF HYDROIDA IN THE SOUTH AFRICAN MUSEUM 135
Eudendrium ritchiei Millard, 1975: 87, fig. 30.
Holotype: SAM-H1803. Abundant male and female colonies; plus two
whole mounts.
Locality: Saunders Rocks, Sea Point; littoral; 5 March 1973.
Collected by: N. A. H. Millard.
Family Bougainvilliidae
Clavopsella quadranularia Thiel, 1962: 227, figs 1-28.
= Clavopsella navis: Millard, 1975: 100, fig. 34A—D. (See also Rhizorhagium
navis.)
Schizoparatypes: SAM-H1774, H1775 (presented by Dr H. Thiel 1972).
Locality: Kiel Canal.
Rhizorhagium navis Millard, 1959b: 244, fig. 2.
= Clavopsella navis: Millard, 1975: 100, fig. 34A—D. (See also Clavopsella
quadranularia.)
Holotype: SAM-—H124 (UCT SH 429B). Male and female colonies on weed
and other hydroids; plus one whole mount.
Locality: Cape Town docks, on ship’s hull; 10 February 1958.
Collected by: N. A. H. Millard.
Silhouetta uvacarpa Millard & Bouillon, 1973: 25, fig. 3A—D, pls 2-3.
Schizoholotype: SAM-H2916 (MT 2664). Three whole mounts only,
bearing medusa buds.
Locality: Silhouette Island, Seychelles; 1966.
Collected by J. Bouillon.
This is the type species of the genus Silhouetta Millard & Bouillon, 1973.
Family Hydractiniidae
Clavactinia multitentaculata Millard, 1975: 106, fig. 35C-G.
As Hydractinia sp.: Millard, 1968: 255.
Holotype: SAM-—H389 (PF station 10724). Male colony on shell of gastro-
pod Melapium lineatum (Lamarck), occupied by hermit Dardanus arrosor
(Herbst); plus two whole mounts; plus microscope sections.
Locality: off Natal, approx. 29°54’S 31°11’E; 99 m; 14 December 1900.
Collected by: R.S. Pieter Faure.
136 ANNALS OF THE SOUTH AFRICAN MUSEUM
Hydractinia altispina Millard, 1955: 215, fig. 1.
Lectotype: SAM-—H87 (UCT F 274). Male colony on gastropod Thais
squamosa (Lamarck); plus microscope sections.
Locality: St James, False Bay; littoral; 1 August 1936.
Paralectotypes: SAM-—H88 (UCT CP 258). Infertile colonies on two
specimens of Thais squamosa; plus one whole mount; St James, False
Bay; littoral; April, 1944.
SAM-H89 (UCT B 92). Colonies with very young gonophores on two
specimens of Thais squamosa; Lambert’s Bay; littoral; 30 July 1938.
Collected by: University of Cape Town.
Hydractinia canalifera Millard, 1957: 179, fig. 1.
Holotype: SAM—H97 (UCT CP 332). Female colony on weed; plus one
whole mount; plus microscope sections.
Locality: Clovelly, False Bay; littoral; 23 August 1949.
Collected by: N. A. H. Millard.
Hydractinia diogenes Millard, 1959a: 305, fig. 2.
Holotype: SAM-H123 (UCT MOR SIH). Male and female colonies on
five gastropod shells occupied by hermits Diogenes costatus (Fabricius);
plus three whole mounts; plus microscope sections.
Locality: off mouth of Rio Coche, Mozambique; 3—5 m; 21 January 1954.
Collected by: University of Cape Town.
Hydractinia kaffraria Millard, 1955: 217, fig. 2.
Lectotype: SAM—H90 (UCT BRE 111A). Female colony on gastropod
Nassarius kraussianus (Dunker).
Locality: Breede River estuary; 3 February 1952.
Paralectotypes: SAM-—H92 (UCT SUN 3N). Colonies on five specimens of
Nassarius kraussianus; plus one whole mount; Sunday’s River estuary;
7 January 1950.
SAM-H91 (UCT HAM 3Q). Microscope sections of female colony;
Keiskama River estuary, The Haven; 9 January 1950.
Collected by: University of Cape Town.
Hydractinia marsupialia Millard, 1975: 113, fig. 38.
Holotype: SAM-H1854 (UCT TB 13). Female colony on gastropod
Nassarius speciosus Adams; plus microscope sections of male and female
colonies.
Locality: Table Bay; 9-18 m; 4 August 1946.
TYPE SPECIMENS OF HYDROIDA IN THE SOUTH AFRICAN MUSEUM 137
Paratypes: SAM-H1855 (UCT TB 14 plus 15). Nine colonies, male and
female, on Nassarius speciosus; plus one whole mount; Table Bay;
17-20 m; 25 October 1946 and 11 February 1947.
Collected by: University of Cape Town.
Family Cytaeidae
Podocoryne nassa Millard, 1959a: 307, fig. 3.
= Cytaeis nassa: Rees, 1962: 390, figs 8-9, pl. 11.
Holotype: SAM-H122 (UCT IN 112). Fertile colonies on two shells of
gastropod Nassarius fenestratus Marrat; plus one whole mount; plus
microscope sections.
Locality: Punta Torres, Inhaca Island, Mozambique; littoral; 18 September
1955.
Collected by: University of the Witwatersrand.
Suborder THECATA
Family Campanulinidae
Aequorea africana Millard, 1966: 461, fig. 8.
Holotype: SAM—H420 (UCT MB 70G). Fertile colony on empty gastropod
shell; plus two whole mounts.
Locality: Mossel Bay, 34°08,9’S 22°07,9’E; 18 m; 19 January 1956.
Collected by: University of Cape Town.
Egmundella amirantensis Millard & Bouillon, 1973: 40, fig. SA—D.
Schizoholotype: SAM-—H2917 (MT 2665). One whole mount only of fertile
material.
Locality: Amirante Island, Seychelles; 1966.
Collected by: J. Bouillon.
Lineolaria gravierae Millard, 1975: 134, fig. 43H.
As Lineolaria sp.: Gravier, 1970: 144, figs 11, 13A. Millard & Bouillon, 1974:
22, fig. 2D.
Schizoholotype: SAM-—H1955. One whole mount (infertile) only.
Locality: Barreira Vermelha, Inhaca, Mozambique, 26°02’S 32°54’E; 1969.
Collected by: J. Bouillon.
138 ANNALS OF THE SOUTH AFRICAN MUSEUM
Lovenella chiquitita Millard, 1957: 198, fig. 7.
Holotype: SAM-—H103 (UCT FAL 288J). One whole mount only of fertile
colony epizootic on Eudendrium deciduum Millard.
Locality: False Bay, 34°09,6’S 18°49,8’E; 18 m; 15 October 1897.
Collector unknown.
Paratypes: SAM-H104 (UCT FB 131K). Two whole mounts (infertile),
one epizootic on Sertularella polyzonias falsa Millard; False Bay,
34°09’S 18°26,7’E; 5-8 m; 13 December 1949.
SAM-H105 (UCT FAL 1080). One whole mount (infertile) epizootic on
Sertularella polyzonias falsa; False Bay, 34°09,3’S 18°51’E; 8-12 m;
23 January 1953.
Collected by: University of Cape Town.
Family Haleciidae
Halecium dufresneae Millard, 1977a: 8, fig. 2A—D.
Schizoholotype: SAM-—H2785 (MD station 26/63). Fragments of female
colony; plus one whole mount.
Locality: Crozet Island, Chenal des Orques, 46°21,5’S 51°55’E; 230 m;
20 April 1974.
Collected by: R.V. Marion Dufresne.
Halecium inhacae Millard, 1958: 168, fig. 1.
Holotype: SAM-—H114 (UCT IN 140H). Female colony on weed; plus two
whole mounts.
Locality: Inhaca Island, Mozambique, east shore rocks; 20 July 1956.
Collected by: University of the Witwatersrand.
Halecium parvulum, var. magnum Millard, 1957: 190, fig. 4B—O.
Since incorporated in Halecium delicatulum Coughtrey, 1876 (large form).
Holotype: SAM-—H99 (UCT FAL 274R). Male and female colonies; plus
three whole mounts.
Locality: False Bay, 34°10,9’S 18°27,2’E; 14-17 m; 21 September 1954.
Collected by: University of Cape Town. ,
Paratypes: SAM-H100 (UCT FAL 159L). Infertile colony; plus one whole
mount; False Bay, 34°12,5’S 18°28’E; 0-3 m; 21 April 1953.
Collected by: University of Cape Town.
SAM-HI11 (PF station 405). Male colony; plus two whole mounts;
off Buffels Bay, False Bay; depth unknown; 8 October 1898.
SAM-H30 (PF station 16287). Female colony; plus two whole mounts;
off Cape Point, approx. 34°20’S 18°32’E; 59 m; 9 December 1902.
Collected by: R.S. Pieter Faure.
TYPE SPECIMENS OF HYDROIDA IN THE SOUTH AFRICAN MUSEUM 139
Hydrodendron sympodiformis Millard & Bouillon, 1974: 25, fig. 5.
Schizoholotype: SAM-H1732 (MT station InPT 42). Part of a female
colony on weed; plus one whole mount.
Locality: Punta Torres, Inhaca, Mozambique; August 1969.
Collected by: J. Bouillon.
Family Lafoeidae
Hebella furax Millard, 1957: 200, fig. 8.
Holotype: SAM-H34 (PF station 18293). Infertile colony parasitic and
epizootic on Lytocarpus filamentosus (Lamarck); plus two whole mounts.
Locality: False Bay, approx. 34°05,5’S 18°39’E; 15— 18 m; 7 December 1903.
Collected by: R.S. Pieter Faure.
Paratype: SAM-H106 (UCT FAL 58Y). Infertile colony on Lytocarpus
filamentosus; one whole mount only; False Bay, 34°09,4’S 18°50,4’E;
12 m; 25 June 1952.
Collected by: University of Cape Town.
Hebella muscensis Millard & Bouillon, 1975: 10, fig. 3A—B.
Schizoholotype: SAM-—H2918 (MT station SEY 20K). Two whole mounts
only of fertile colony epizootic on Synthecium sp.
Locality: Anse la Mouche, Seychelles; 1972.
Collected by: J. Bouillon.
Hebella urceolata Millard, 1964: 11, fig. 2A.
Since incorporated in Hebella scandens (Bale, 1888)
Holotype: SAM-H410 (UCT SCD 154H). Infertile colony epizootic on
Halecium beanii (Johnston); plus one whole mount.
Locality: Agulhas Bank, 34°03’S 25°59’E; 84 m; 25 November 1960.
Collected by: University of Cape Town.
Scandia tubitheca Millard & Bouillon, 1975: 10.
As Scandia corrugata: Millard & Bouillon, 1973: 60 (pro parte), fig. 8D-F.
Schizoholotype: SAM-—H2919 (MT station Am IP). Male colony epizootic
on Synthecium dentigerum Jarvis; two whole mounts only.
Locality: Amirante Island, Seychelles; 30 August 1966.
Collected by: J. Bouillon.
Zygophylax cornucopia Millard, 1955: 219, fig. 3.
= Hydrodendron cornucopia: Millard, 1973: 33, fig. 6A—E (transferred to
Haleciidae).
Holotype: SAM—H93 (UCT FB 131B). Male and female colonies epizootic
on Antennella quadriaurita Ritchie; plus one whole mount.
140 ANNALS OF THE SOUTH AFRICAN MUSEUM
Locality: False Bay, 34°09’S 18°26,7’E; 5-8 m; 13 December 1949.
Paratypes: SAM-H94 (UCT TB 1B). Infertile colonies epizootic on
A. quadriaurita; plus one whole mount; Table Bay, 33°47,5’S 18°24,3’E;
19-20 m; 11 February 1947.
SAM-—H95 (UCT FAL 78S). Fertile colonies epizootic on A. quadriaurita:
plus one whole mount; False Bay, several positions; 4-17 m; 19 August
1952-23 September 1954.
SAM-H96 (UCT FAL 217N); one whole mount of infertile colony
epizootic on A. quadriaurita; False Bay, 34°07’S 18°32,5’E; 18 m;
9 September 1953.
Collected by: University of Cape Town.
Zygophylax crozetensis Millard, 1977a: 15, fig. 4.
Schizoholotype: SAM—H2779 (MD station 26/64). Part of a large fertile
colony; plus one whole mount.
Locality: Crozet Island, Chenal des Orques, 46°24’S 51°59’E; 180 m;
20 April 1974.
Collected by: R.V. Marion Dufresne.
Zygophylax cnigmatica Millard, 1964: 19, fig. SA—F.
= Hydrodendron gracilis (Fraser, 1914): Millard, 1973: 33, fig. 6F—G (trans-
ferred to Haleciidae).
Holotype: SAM-H411 (UCT WCD 12E); female colony epizootic on
Nemertesia ramosa Lamouroux; two whole mounts only.
Locality: off Cape Town, 34°09,4’S 18°16,5’E; 75 m; 24 March 1959.
Collected by: University of Cape Town.
Zygophylax geminocarpa Millard, 1958: 177, fig. 4D-G.
Holotype: SAM-HS59 (PF station 12308). About eight fertile stems; plus
two whole mounts.
Locality: off Natal, approx. 30°53’S 30°28’E; 66 m; 14 March 1901.
Collected by: R.S. Pieter Faure.
Zygophylax inconstans Millard, 1977b: 117, fig. 5.
Holotype: SAM-H1975 (SM station 23J). A fertile, tangled colony on a
bryozoan; plus one whole mount.
Locality: off Natal, 27°44,4’S 32°42,8’E; 400-450 m; 26 May 1975.
Collected by: R.V. Meiring Naude.
Zygophylax infundibulum Millard, 1958: 180, fig. 4B—C.
Holotype: SAM-—H36 (PF station 10781). Several infertile stems; plus two
whole mounts.
Locality: off Natal, approx. 29°53’S 31°11’E; 155 m; 17 December 1900.
Collected by: R.S. Pieter Faure.
TYPE SPECIMENS OF HYDROIDA IN THE SOUTH AFRICAN MUSEUM 141
Family Campanulariidae
Campanularia laminacarpa Millard, 1966: 472, fig. 12E-K.
Holotype: SAM-—H421 (UCT TRA 32A). A female colony epizootic on
Thyroscyphus aequalis Warren; plus two whole mounts.
Locality: Agulhas Bank, 34°49’S 20°21,5’E; approx. 91 m; 9 November
1947.
Collected by: commercial trawler.
Campanularia morgansi Millard, 1957: 195, fig. 6.
Holotype: SAM-H24 (PF station 15675). Female colonies epizootic on
Corhiza scotiae (Ritchie) and Sertularella arbuscula (Lamouroux); plus
one whole mount.
Locality: False Bay, approx. 34°28’S 18°32’E; 73 m; 9 October 1902.
Collected by: R.S. Pieter Faure.
Paratypes: SAM-H7 (PF station 351). Infertile colony epizootic on
Salacia articulata (Pallas); plus one whole mount; False Bay, approx.
34°19’S 18°31’E; 58-62 m; 28 September 1898.
SAM-H32 (PF station 18232). Fertile colony epizootic on Eudendrium
?deciduum Millard; False Bay, approx. 34°27’S 18°45’E; 110 m;
11 November 1903.
Collected by: R.S. Pieter Faure.
SAM-H101 (UCT FB 119L plus FAL 289B). Male and infertile colonies
epizootic on Salacia articulata (Pallas); plus two whole mounts; False
Bay, 34°08,5’S 18°34,5’E; 27 m; 21 April 1947; and another locality in
False Bay, exact location and collector unknown; 40 m; September 1897.
Collected by: University of Cape Town.
SAM-H102 (UCT FAL 26L). Infertile colony on empty tubes; one whole
mount only; False Bay, 34°13’S 18°29’E; 15-21 m; 5 March 1952.
Collected by: J. C. Morgans.
Campanularia pecten Gow & Millard, 1975: 1, fig. 1.
Holotype: SAM-H1659. Male colony growing on sea-grass Caulerpa
filiformis (Suhr); plus four whole mounts.
Locality: St James, False Bay; littoral; 29 March 1968.
Collected by: C. Gow.
Campanularia roberti Gow & Millard, 1975: 3, fig. 2.
Holotype: SAM-H1660. Female colony growing on weed Sargassum
longifolium (Turner) attached to the kelp Ecklonia maxima (Osbeck);
plus four whole mounts.
Locality: Partridge Point, False Bay; 24 March 1968.
Collected by: R. W. Day.
142 ANNALS OF THE SOUTH AFRICAN MUSEUM
Campanularia subantarctica Millard, 1971: 403, fig. 5.
Holotype: SAM—H2920. Two whole mounts only of fertile colony epizootic
on Symplectoscyphus marionensis Millard.
Locality: Near mouth of Soft Plume River, Marion Island; littoral;
7 February 1965.
Collected by: N. R. Fuller.
Clytia latitheca Millard & Bouillon, 1973: 55, fig. 7H-L.
Schizoholotype: SAM—H2921 (MT 2666). Two whole mounts only of fertile
colony.
Locality: Praslin Island, Seychelles; 1966.
Collected by: J. Bouillon.
Clytia serrata Millard, 1958: 173, fig. 3C, H.
= Clytia gravieri (Billard): Millard & Bouillon, 1973: 51, fig. 7E-G.
Holotype: SAM-H115 (UCT MOR 216C). One whole mount only of
infertile colony epizootic on Eudendrium carneum Clarke.
Locality: On wreck at Linga Linga, Morrumbene estuary, Mozambique;
15 July 1954.
Collected by: University of Cape Town.
Family Syntheciidae
Hincksella corrugata Millard, 1958: 181, fig. 5.
Holotype: SAM-H85 (PF station 12456). Infertile colony on weed; plus
one whole mount.
Locality: off Natal, approx. 30°32’S 30°38,5’E; 46 m; 22 March 1901.
Collected by: R.S. Pieter Faure.
Hincksella indiana Millard, 1967: 178, fig. 3D-G.
Holotype: SAM-H1646 (UCT AFR 1235A). One infertile stem in three
parts; plus one whole mount of hydrocladia.
Locality: south of Madagascar, 27°48’S 47°19’E; 875 m; 22 June 1961.
Collected by: R.V. Africana II.
Synthecium hians Millard, 1957: 204, fig. 9A-C.
Holotype: SAM-—H107 (UCT FAL 214G). Several infertile stems; plus two
whole mounts.
Locality: False Bay, 34°12,4’S 18°43,5’E; 42 m; 10 September 1953.
Collected by: University of Cape Town.
TYPE SPECIMENS OF HYDROIDA IN THE SOUTH AFRICAN MUSEUM 143
Family Sertulariidae
Salacia disjuncta Millard, 1964: 31, fig. 1OA-F.
Holotype: SAM-H412 (UCT SCD 37K). One whole mount only (infertile).
Locality: Agulhas Bank, 32°15,2’S 28°57,7’E; 50 m; 19 May 1958.
Paratype: SAM-H1782 (UCT SCD 296H). One whole mount only
(infertile); Agulhas Bank, 33°09’S 28°02’E; 84 m; 6 February 1962.
Collected by: University of Cape Town.
Sertularella agulhensis Millard, 1964: 35, fig. 12A.
Holotype: SAM-—H413 (UCT TRA 151F). Several infertile stems; plus one
whole mount.
Locality: Agulhas Bank, 34°51’S 19°55’E; 22 m; 6 March 1958.
Collected by: commercial trawler.
Sertularella capensis Millard, 1957: 210, fig. 10H.
Holotype: SAM-H108 (UCT FB 114A). A rich infertile colony; plus one
whole mount.
Locality: False Bay, 34°07,5’S 18°31’E; 27-28 m; 22 February 1947.
Paratypes: SAM-H109 (UCT FB 115D). A fairly rich infertile colony;
False Bay, 34°08’S 18°31,5’E; 27-28 m; 22 February 1947.
SAM-H110 (UCT FAL 64L). A fairly rich infertile colony; plus one
whole mount; False Bay, 34°17,3’S 18°48,7’E; 37-38 m; 29 July 1952.
Collected by: University of Cape Town.
Sertularella capensis delicata Millard, 1964: 38, fig. 12B—D.
Since incorporated in Sertularella annulaventricosa Mulder & Trebilcock, 1915.
Holotype: SAM-H414 (UCT NAD 22F). Several infertile stems; plus one
whole mount.
Locality: off Natal, 29°58’S 31°02’E; 49 m; 12 August 1958.
Collected by: University of Cape Town.
Sertularella congregata Millard, 1964: 39, fig. 13A—D.
Holotype: SAM-H415 (UCT SCD 254Q). Fertile colony; plus one whole
mount.
Locality: Agulhas Bank, 33°07,3’S 28°01’E; 88 m; 16 July 1961.
Collected by: University of Cape Town.
Paratype: SAM-H18S. Infertile colony; plus one whole mount; Agulhas
Bank, 33°09’S 28°03’E; 86 m; 28 December 1898.
Collector unknown.
144 ANNALS OF THE SOUTH AFRICAN MUSEUM
Sertularella dubia, var. magna Millard, 1958: 189, fig. 7A.
= Sertularella dubia magna Millard.
Holotype: SAM-HS54 (PF station 12028). Several infertile stems; plus one
whole mount.
Locality: off Natal, approx. 28°41’S 32°22’E; 62 m; 27 February 1901.
Collected by: R.S. Pieter Faure.
Sertularella falsa Millard, 1957: 211, figs 1OF, 11D.
= Sertularella polyzonias falsa Millard.
Holotype: SAM-H111 (UCT FB 119C). A fairly rich fertile colony; plus
one whole mount.
Locality: False Bay, 34°08,5’S 18°34,5’E; 27 m; 21 April 1947.
Paratypes: SAM-—H112 (UCT FB 131H). Rich fertile colonies from several
localities in False Bay and on various dates; plus one whole mount.
SAM-H113 (UCT CP 333B). Fertile colony; plus one whole mount;
Dalebrook, False Bay; littoral; 25 August 1949.
Collected by: University of Cape Town.
Sertularella gilchristi Millard, 1964: 44, fig. 12E, G-H.
Holotype: SAM-H416 (UCT SCD 85J). A fairly rich male colony; plus
one whole mount.
Locality: Agulhas Bank, 33°03’S 27°55’E; 27 m; 17 July 1959.
Collected by: University of Cape Town.
Sertularella mediterranea var. asymmetrica Millard, 1958: 191, fig. 7B.
= Sertularella mediterranea asymmetrica Millard.
Holotype: SAM-H116 (UCT IN 49K). Several female stems; plus two
whole mounts.
Locality: Inhaca Island, Mozambique; littoral; 1954.
Collected by: University of the Witwatersrand.
Sertularella natalensis Millard, 1968: 271, fig. 4E-G.
Schizoholotype: SAM—H2922 (TME 23M). One whole mount only.
Locality: off Natal, 29°47,5’S 31°11,8’E; 64 m; 22 August 1929.
Collected by: Th. Mortensen’s Java—South Africa Expedition.
Sertularia linealis var. longa Millard, 1958: 197, fig. 8E.
= Sertularia longa: Millard & Bouillon, 1974: 33, fig. 7D, J.
Holotype: SAM-H117 (UCT IN 140E). Infertile colony on weed; plus one
whole mount.
Locality: East Shore Rocks, Inhaca Island, Mozambique; 20 July 1956.
Collected by: University of the Witwatersrand.
TYPE SPECIMENS OF HYDROIDA IN THE SOUTH AFRICAN MUSEUM 145
Symplectoscyphus marionensis Millard, 1971: 405, fig. 7.
Holotype: SAM—H2923. One whole mount only of fertile colony.
Locality: near Soft Plume River, Marion Island; littoral; 1965.
Collected by: N. R. Fuller.
Uniscyphus fragilis Millard, 19776: 122, fig. 7E-G.
Holotype: SAM-H1982 (SM station 43). Two hydrothecae; plus one whole
mount of three hydrothecae.
Locality: off Natal, 28°45,5’S 32°24,5’E; 360-420 m; 29 May 1975.
Collected by: R.V. Meiring Naude.
This is the type species of the genus Uniscyphus Millard, 1977.
Family Plumulariidae
Subfamily Halopterinae
Corhiza bellicosa Millard, 1962: 275, fig. 2A—E.
Holotype: SAM-—H364 (UCT SCD 84S). Several infertile stems; plus one
whole mount.
Locality: Agulhas Bank, 33°03’S 27°55’E; 27 m; 17 July 1959.
Paratype: SAM-H1947 (UCT SCD 153J). One infertile stem; plus one
whole mount; Agulhas Bank, 34°03’S 25°59’E; 84 m; 25 November 1960.
Collected by: University of Cape Town.
Corhiza mortenseni Millard, 1968: 274, fig. SA—D.
Schizoholotype: SAM-—H2914 (TME 24). Two whole mounts only.
Locality: off Natal, 29°48,5’S 31°18’E; approx. 219 m; 22 August 1929.
Collected by: Th. Mortensen’s Java—South Africa Expedition.
Corhiza pannosa Millard, 1962: 278, fig. 3A—B, D-G.
Holotype: SAM-—H362 (UCT SCD 5H). One infertile colony; plus one
whole mount.
Locality: Agulhas Bank, 34°15’S 25°05’E; 11 m; 19 April 1958.
Collected by: University of Cape Town.
Gattya tropicalis Millard & Bouillon, 1973: 79, fig. 1OA—D.
Schizoholotype: SAM—H2924 (MT 2667). One whole mount only.
Locality: Beau Vallon, Mahé Island, Seychelles; 1966.
Collected by: J. Bouillon.
146 ANNALS OF THE SOUTH AFRICAN MUSEUM
Halopteris gemellipara Millard, 1962: 282, fig. 4A—F.
Holotype: SAM-—H308 (PF station 13810). An infertile colony; plus one
whole mount. ;
Locality: Agulhas Bank, approx. 33°54’S 26°51’E; 120 m; 10 September
1901.
Collected by: R.S. Pieter Faure.
Halopteris pseudoconstricta Millard, 1975: 355, fig. 114D-G.
As Halopteris constricta: Millard, 1957: 227, fig. 14A.
Holotype: SAM—H542 (UCT CP 740D). Numerous pinnate stems bearing
female gonophores and a few infertile simple stems; plus one whole
mount.
Locality: Melkbosstrand, Table Bay; littoral; 17 November 1967.
Collected by: University of Cape Town.
Halopteris rostrata Millard, 1975: 357, fig. 114A-C.
Holotype: SAM-—H543 (UCT NAD 89H). Eleven infertile stems and several
damaged ones growing on skeleton of antipatharian; plus two whole
mounts.
Locality: off Natal, 29°11’S 32°02’E; 70 m; 30 July 1964.
Collected by: University of Cape Town.
Monostaechas faurei Millard, 1958: 204, fig. 11.
Holotype: SAM-H58 (PF station 12028). Six male stems; plus two whole
mounts.
Locality: off Natal, approx. 28°41’S 32°22’E; 62 m; 27 February 1901.
Collected by: R.S. Pieter Faure.
Monostaechas natalensis Millard, 1958: 206, fig. 12.
Holotype: SAM-H79 (PF station 12456). Many stems, male and female;
plus two whole mounts.
Locality: off Natal, approx. 30°32’S 30°38,5’E; 46 m; 22 March 1901.
Paratypes: SAM-H48 (PF station 11803). One infertile stem; plus one
whole mount; off Natal, approx. 29°00’S 31°49’E; 24 m; 8 February 1901.
SAM-H76 (PF station 12392). Several infertile stems; plus one whole
mount; off Natal, approx. 31°02,5’S 30°18’E; 46 m; 14 March 1901.
Collected by: R.S. Pieter Faure.
TYPE SPECIMENS OF HYDROIDA IN THE SOUTH AFRICAN MUSEUM 147
Subfamily Plumulariinae
Plumularia antonbruuni Millard, 1967: 185, fig. 5.
Holotype: SAM-H1647 (UCT ABD 14C). Three male stems; plus one
whole mount.
Locality: off Natal, 29°45’S 31°40’E; 440 m; 8 September 1964.
Collected by: R.V. Anton Bruun.
Plumularia irregularis Millard, 1958: 210, fig. 13A—C.
= Kirchenpaueria irregularis: Millard, 1975: 370, fig. 118D-G (transferred to
subfamily Kirchenpaueriinae).
Holotype: SAM-H119 (UCT DBN 70Q). Three infertile stems; plus one
whole mount.
Locality: Salisbury Island, Durban Bay; littoral; 22 July 1950.
Collected by: University of Cape Town.
Plumularia mossambicae Millard, 1975: 393, fig. 123E-G.
Holotype: SAM-H1866 (UCT PED 4V). One whole mount only (infertile
stem).
Locality: off Mozambique, 24°46’S 35°18’E; 110 m; 18 August 1964.
Collected by: University of Cape Town.
Plumularia pennycuikae Millard & Bouillon, 1973: 85, fig. 1ON-P.
As Plumularia sp.: Pennycuik, 1959: 183, pl. 3 (fig. 7).
Schizoholotype: SAM-—H2925 (MT 2668). One whole mount only.
Locality: Silhouette Island, Seychelles; 1966.
Collected by: J. Bouillon.
Subfamily Aglaopheniinae
Cladocarpus crepidatus Millard, 1975: 417, fig. 130A-C.
Holotype: SAM-—H537 (UCT SST 6GG). One female stem; plus one whole
mount.
Locality: Agulhas Bank, 35°22’S 22°31’E; 200 m; 20 June 1972.
Collected by: University of Cape Town.
Cladocarpus natalensis Millard, 1977b: 127, fig. 10.
Holotype: SAM-—H2861 (SM station 86). Five fertile stems; plus one whole
mount.
Locality: off Natal, 27°59,5’S 32°40,8’E; 550 m; 22 May 1976.
Collected by: R.V. Meiring Naude.
148 ANNALS OF THE SOUTH AFRICAN MUSEUM
Cladocarpus paries Millard, 1975: 483, fig. 143.
Holotype: SAM-—H1914. One infertile stem; plus two whole mounts.
Locality: Agulhas Bank, 33°25’S 27°29’E; 70 m; 23 January 1975.
Collected by: South African Museum.
Cladocarpus unicornus Millard, 1975: 429, fig. 133E-F.
Holotype: SAM—H538 (UCT PED 4S). One infertile stem; plus one whole
mount.
Locality: off Mozambique, 24°46’S 35°18’E; 110 m; 18 August 1964.
Collected by: University of Cape Town.
Halicornaria africana Millard, 1958: 215, fig. 1SA—C.
= Gymnangium africanum (Millard).
Holotype: SAM-H120 (UCT AFR 1028B). One infertile colony; plus one
whole mount.
Locality: off Natal, 28°28’S 32°25,8’E; 27 m; 15 May 1948.
Collected by: R.V. Africana IT.
Halicornaria arcuata var. epizootica Millard, 1958: 218, fig. 15F.
= Gymnangium arcuatum (Lamouroux, 1816), epizootic form.
Holotype: SAM-H73 (PF station 12392). One whole mount only, infertile,
epizootic on Thecocarpus formosus (Busk).
Locality: off Natal, approx. 31°02,5’S 30°18’E; 46 m; 14 March 1901.
Collected by: R.S. Pieter Faure.
Halicornaria exserta Millard, 1962: 309, fig. 11A—E.
= Gymnangium exsertum (Millard).
Holotype: SAM-H169 (PF station 596). One fertile stem; plus one whole
mount.
Locality: Agulhas Bank, approx. 33°49’S 25°56’E; depth unknown;
11 November 1898.
Collected by: R.S. Pieter Faure.
Halicornaria exserta epizootica Millard, 1962: 309, fig. 11F—H.
= Gymnangium exsertum (Millard), epizootic form.
Holotype: SAM-—H320 (PF station 13915). Colony epizootic on Thecocarpus
flexuosus umbellatus Millard; plus one whole mount.
Locality: Agulhas Bank, approx. 34°05’S 26°34’E; 115 m; 23 September
1901.
Collected by: R.S. Pieter Faure.
TYPE SPECIMENS OF HYDROIDA IN THE SOUTH AFRICAN MUSEUM 149
Thecocarpus flexuosus umbellatus Millard, 1962: 316, fig. 12B, M.
Holotype: SAM—H363 (UCT SCD 36M). Large infertile colony; plus one
whole mount.
Locality: Agulhas Bank, 32°15,2’S 28°57,7’E; 49,5 m; 19 May 1958.
Collected by: University of Cape Town.
Thecocarpus giardi var. solidus Millard, 1958: 222, fig. 16B—C.
= Thecocarpus flexuosus solidus Millard.
Holotype: SAM-H121 (UCT AFR 1028A). Large fertile colony; plus two
whole mounts.
Locality: off Natal, 28°28’S 32°25,8’E; 27 m; 15 May 1948.
Collected by: R.V. Africana II.
REFERENCES
BaLe, W. M. 1888. On some new and rare Hydroida in the Australian Museum collection.
Proc. Linn. Soc. N.S.W. (2) 3: 745-799.
CouGuTrey, M. 1876. Critical notes on the New Zealand Hydroida. Trans. Proc. N.Z. Inst. 8:
298-302.
Fraser, C. M. 1914. Some hydroids of the Vancouver Island region. Trans. R. Soc. Can. (3) 8:
99-216.
Gow, C. & MILLARD, N. A. H. 1975. Two new species of campanularian hydroids from
South Africa. Ann. S. Afr. Mus. 67: 1-6.
GrAVIER, N. 1970. Etude des Hydraires epiphytes des Phanérogames marines de la région de
Tulear (sud-oest de Madagascar). Rec! Trav. Stn mar. Endoume-Marseille 10: 111-161.
Lamouroux, J. V. F. 1816. Histoire des polypiers coralligénes flexibles, vulgairement nommés
zoophytes. Caen: Poisson.
MILLARD, N. A. H. 1955. New species of Hydrozoa from South Africa. Ann. S. Afr. Mus. 41:
215-222.
MILLARD, N. A. H. 1957. The Hydrozoa of False Bay, South Africa. Ann. S. Afr. Mus. 43:
173-243.
MILLARD, N. A. H. 1958. Hydrozoa from the coasts of Natal and Portuguese East Africa.
Part I. Calyptoblastea. Ann. S. Afr. Mus. 44: 165-226.
MILLARD, N. A. H. 1959a. Hydrozoa from the coasts of Natal and Portuguese East Africa.
Part II. Gymnoblastea. Ann. S. Afr. Mus. 44: 297-313.
MILLARD, N. A. H. 19596. Hydrozoa from ships’ hulls and experimental plates in Cape Town
docks. Ann. S. Afr. Mus. 45: 239-256.
MILLARD, N. A. H. 1962. The Hydrozoa of the south and west coasts of South Africa. Part I.
The Plumulariidae. Ann. S. Afr. Mus. 46: 261-319.
MILLARD, N. A. H. 1964. The Hydrozoa of the south and west coasts of South Africa. Part II.
The Lafoeidae, Syntheciidae and Sertulariidae. Ann. S. Afr. Mus. 48: 1-56.
MILLARD, N. A. H. 1966. The Hydrozoa of the south and west coasts of South Africa. Part III.
The Gymnoblastea and small families of Calyptoblastea. Ann. S. Afr. Mus. 48: 427-487.
MILLARD, N. A. H. 1967. Hydroids from the south-west Indian Ocean. Ann. S. Afr. Mus. 50:
169-194,
MILLarD, N. A. H. 1968. South African hydroids from Dr Th. Mortensen’s Java—South Africa
Expedition, 1929-1930. Vidensk. Meddr dansk naturh. Foren. 131: 251-288.
MILLARD, N. A. H. 1971. Hydrozoa. In: ZINDEREN BAKKER, E. M. VAN, WINTERBOTTOM, J. M.
& Dyer, R. A., eds. Marion and Prince Edward Islands: 396-408. Cape Town:
A. A. Balkema.
MILLARD, N. A. H. 1973. Auto-epizoism in South African hydroids. Proc. Second Internat.
Symp. Cnidaria; Publs Seto mar. biol. Lab. 20: 23-34.
150 ANNALS OF THE SOUTH AFRICAN MUSEUM
MILLARD, N. A. H. 1975. Monograph on the Hydroida of southern Africa. Ann. S. Afr. Mus.
68: 1-513.
MILLARD, N. A. H. 1977a. Hydroids from the Kerguelen and Crozet shelves, collected by the
cruise MD.03 of the Marion-Dufresne. Ann. S. Afr. Mus. 73: 1-47.
MILLARD, N. A. H. 19776. The South African Museum’s Meiring Naude cruises. Part 3.
Hydroida. Ann. S. Afr. Mus. 73: 105-131.
Miiiarpb, N. A. H. & BourLton, J. 1973. Hydroids from the Seychelles (Coelenterata).
Annls Mus. r. Afr, cent. Sér. 8 vo (Sci. zool.) 206: 1-106.
MILLARD, N. A. H. & BourILLon, J. 1974. A collection of hydroids from Mocgambique, East
Africa. Ann. S. Afr. Mus. 65: 1-40.
MILLARD, N. A. H. & BoumLton, J. 1975. Additional hydroids from the Seychelles. Ann. S. Afr.
Mus. 69: 1-15.
Mu.per, J. E. & Trepitcock, R. E. 1915. Victorian Hydroida. With description of new
species. Part V. Geelong Nat. 1915: 51-S9.
PENNYCUIK, P. R. 1959. Faunistic records from Queensland. Part VY. Marine and brackish
water hydroids. Pap. Dep. Zool. Univ. Qd 1: 141-210.
Rees, W. J. 1962. Hydroids of the family Cytaeidae L. Agassiz, 1862. Bull. Br. Mus. nat. Hist.
(Zool.) 8: 381-400.
TureL, H. 1962. Clavopsella quadranularia nov. spec. (Clavopsellidae nov. fam.), ein neuer
Hydroidpolyp aus der Ostsee und seine phylogenetische Bedeutung. Z. Morph. Okol.
Tiere 51: 227-260.
6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.
An author's name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.
In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and date.
7. SPECIAL HOUSE RULES
Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. *... the Figure depicting C. namacolus ...?; *. . . in C. namacolus (Fig. 10). .
(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. DuToit but A.L.du Toit; Von Huene but F. von Huene
(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian
Punctuation should be loose, omitting all not strictly necessary
Reference to the author should be expressed in the third person
Roman numerals should be converted to arabic, except when forming part of the title of a
book or article, such as
‘Revision of the Crustacea. Part VIII. The Amphipoda.’
Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.
Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.
N. A. H. MILLARD
TYPE SPECIMENS OF HYDROIDA (COELENTERATA)
IN THE SOUTH AFRICAN MUSEUM
j - a 52 .
A
)
MUS. COMP. ZOOL.
LIBRARY
JUL 2 3 407
HARVARD
OF THE SOUTH vi. RN
APE TOWN
INSTRUCTIONS TO AUTHORS
1. MATERIAL should be original and not published elsewhere, in whole or in part.
2. LAYOUT should be as follows:
(a) Centred masthead to consist of
Title: informative but concise, without abbreviations and not including the names of new genera or species
Author’s(s’) name(s)
Address(es) of author(s) (institution where work was carried out)
Number of illustrations (figures, enumerated maps and tables, in this order)
(b) Abstract of not more than 200 words, intelligible to the reader without reference to the text
(c) Table of contents giving hierarchy of headings and subheadings
(d) Introduction
(e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents
(f) Summary, if paper is lengthy
(g) Acknowledgements
) References
(i) Abbreviations, where these are numerous
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All illustrations, whether line drawings or photographs, should be termed figures (plates
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The number of the figure should be lightly marked in pencil on the back of each illustration.
5. REFERENCES cited in text and synonymies should all be included in the list at the end of
the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable):
(a) Author’s name and year of publication given in text, e.g.:
“Smith (1969) describes...’
‘Smith (1969: 36, fig. 16) describes...’
‘As described (Smith 1969a, 19696; Jones 1971)’
“As described (Haughton & Broom 1927)...’
‘As described (Haughton ef a/. 1927)...’
Note: no comma separating name and year
Pagination indicated by colon, not p.
names of joint authors connected by ampersand re
et al. in text for more than two joint authors, but names of all authors given in list of references.
(b) Full references at the end of the paper, arranged alphabetically by names, chronologically
within each name, with suffixes a, 6, etc. to the year for more than one paper by the same
author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a).
For books give title in italics, edition, volume number, place of publication, publisher.
For journal article give title of article, title of journal in italics (abbreviated according to the World list o,
scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part
number (only if independently paged) in parentheses, pagination (first and last pages of article).
Examples (note capitalization and punctuation)
BuLLouGuH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FIscHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
Fiscuer, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.
Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann, Mag. nat. Hist. (13) 2: 309-320.
Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
TurELe, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 77 Band
May 1979 Mei
Part 9 Deel
THE UNIONACEA
(MOLLUSCA, LAMELLIBRANCHIATA)
OF SOUTH-CENTRAL AFRICA
By
C.<. APPLETON
Cape Town Kaapstad
The ANNALS OF THE SOUTH AFRICAN MUSEUM
are issued in parts at irregular intervals as material
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap
THE UNIONACEA (MOLLUSCA, LAMELLIBRANCHIATA)
OF SOUTH-CENTRAL AFRICA
By
C. C. APPLETON*
Bilharzia Field Research Unit, South African Medical Research Council,
Nelspruit
(With 8 figures and | table)
[MS. accepted 11 December 1978]
ABSTRACT
Twelve species of the Unionacea (Mollusca, Lamellibranchiata), five Unionidae and six
Mutelidae and one Etheriidae, are recorded from the south-central region of Africa. Their
distribution within this region, which is taken to embrace the catchments of the Kunene,
Cuvelai, Okavango, Linyanti and Zambezi River systems, is updated. Attention is drawn to
the paucity of ecological data relevant to the freshwater bivalves of Africa and the detailed
locality notes made by several collectors are therefore included.
CONTENTS
PAGE
Introdiction a=” 2] 2 Se os: 151
Systematic discussion ae eee aso
General discussion . . . .-. 168
Acknowledgements . . . . . 172
Refererices 9% | ae seep eee Bc 2
INTRODUCTION
Knowledge of the Unionacea of the southern central African region
embracing the Kunene, Cuvelai, Okavango and Zambezi River systems (i.e.
between latitudes 9° and 21°S) has hitherto been limited, particularly with
regard to the Kunene and Okavango. During the past decade, however, a
considerable amount of material has been collected from these drainages and
through the co-operation of a number of individuals and institutions the
author has been able to examine examples of eleven of the twelve recorded
unionacean species, five Unionidae, five Mutelidae and one Etheriidae, per-
mitting an updating of knowledge on the composition and distribution of this
bivalve fauna. Unfortunately no material is available from the middle and lower
Zambezi, east of Lake Kariba. However, the bivalve fauna of Lake Macllwaine
on the Hunyani River, a tributary to the middle Zambezi, has been discussed
by Marshall (1975) and the malacofauna of Lake Malawi has been dealt with
by Crowley et a/. (1964) and Mandahl-Barth (1972). In addition, the bivalves
of the adjacent Bangweulu—Luapala basin, the Congo River system, and Mozam-
bique have been variously discussed by Mandahl-Barth (1968), Pilsbry &
* Present address:
School of Environmental & Life Sciences, Murdoch University, Murdoch, Western
Australia 6153.
151
Ann. S. Afr. Mus. 77 (9), 1979: 151-174, 8 figs, 1 table.
152 ANNALS OF THE SOUTH AFRICAN MUSEUM
Bequaert (1927), Azevedo et al. (1961), and Braga (1952). In the present contri-
bution the systematic arrangement of Connolly (1939) has been followed,
though changes in nomenclature made by Haas (1969), Mandahl-Barth (1968,
1972) and Heard & Vail (1976) have been incorporated.
Figure 1 shows the watercourses referred to in the text. These comprise
both large, perennial rivers such as the Kunene and Okavango with volumes
that may exceed 6 000 m/s during high flood which can drop to less than 15 m3/s
in dry years (Stengel 1963), as well as largely seasonal floodplain or deltaic
environments such as the Okavango delta, Linyanti swamp, eastern Caprivi
floodplain and the Etosha Pan.
Where measurements of shell height (altitude) are given, maximum height
is used (except where indicated) in preference to umbonal height because in
many cases the umbones were eroded to such an extent that considerable
inaccuracies would have been introduced.
The following abbreviations are used:
SAM — South African Museum, Cape Town
NM Natal Museum, Pietermaritzburg
™ Transvaal Museum, Pretoria
MMK McGregor Memorial Museum, Kimberley
SMN — State Museum, Windhoek
Where no abbreviations are given, the material has been lodged in the
collection of the South African Museum, Cape Town.
Lake
Malawi
2
vate FR. <ambezi R a
Cuvelai
R
Qkavang,
P~
2 Ly
Ekuma R \ JAR
Sf R
D 3
Bie nar Okavango delta y KS
Fig. 1. Map of south-central Africa (between latitudes 9° and 21°S) showing the river systems
referred to in the text.
21
THE UNIONACEA OF SOUTH-CENTRAL AFRICA 153
SYSTEMATIC DISCUSSION
Superfamily UNIONACEA
Family Unionidae
Although many species of Unionidae have been described from southern
Africa (south of but including the investigated area) only four are recognized
here. These are Unio caffer, Caelatura kunenensis, C. mossambicensis and
C. framesi and may even be too many. All except C. framesi, a southerly species,
occur in the area under discussion.
Unio Retzius, 1788
Unio caffer Krauss, 1848
Fig. 2A
Material examined
Zambezi River system:
Kafue River south of Itezhitezhi, Zambia. R. M. Derricourt leg. 1975,
2 specimens + | valve.
Matopos, Rhodesia, M. N. Bruton leg. 1973, 3 specimens.
Nyamandlelovu (on Gwaai River), Rhodesia. R. H. R. Stevenson leg.
26 May 1924, 1 specimen, NM-A5364.
Victoria Falls, collector unknown, 1 valve, MMK MOL-328.
Upper Zambezi River, G. C. Shortridge leg., 1 valve. MMK MOL-327.
Hunyani River, Darwendale, B. E. Marshall leg. May 1975, 2 specimens
+ 1 valve.
Gwenoro dam, Gwelo, Rhodesia. B. E. Marshall leg., 1 specimen.
Heard & Vail (1976) have discussed in detail the anatomy and systematic
position of this species and have illustrated its hooked glochidium. These
glochidia are considerably smaller than those of Caelatura kunenensis found
by the present author. Unio caffer, which attains a far larger size than any other
southern African unionid, is characterized by its heavy shell, strong cardinal
and lateral teeth and downward-sloping hinge line. The posterodorsal ridge is
usually pronounced though not sharp. Some zigzag sculpture is present on
juvenile shells up to about 21 mm in length; on older shells this is generally
worn away and only growth striae remain. Only the outer demibranchs are
modified to form marsupia.
The records from the Kafue and upper Zambezi rivers are the most
northerly for this southern African species. Marshall (1975) has reported it
from Lake Macllwaine (Rhodesia). It is, however, uncommon in the area
under discussion and has not been recorded from the Okavango, Cuvelai or
Kunene River systems, i.e. west of longitude 23°E.
A record of U. caffer from Amaramba in the Niassa district of Mozam-
bique (Azevedo et al. 1961), consisting of a single juvenile approximately
154 ANNALS OF THE SOUTH AFRICAN MUSEUM
6,1 x 3,6 X 2,2 mm, has been suggested by Oberholzer & Van Eeden (1967)
to refer to a higher-shelled species, viz. Caelatura mossambicensis. This is
probably correct since the Amaramba specimen has a L/H ratio of 1,70 whereas
a juvenile U. caffer of similar size from the eastern Transvaal escarpment has
a ratio of 1,95. No juvenile C. mossambicensis are available but comparable
C. kunenensis from the Kafue River (NM-—A5366) have L/H ratios of 1,58—1,62.
Furthermore, the illustration by Azevedo ef al. (1961) of sculpture round only
the umbone is in agreement with the general paucity of sculpture found in
C. mossambicensis and is in marked contrast to juvenile U. caffer where a
sculpture pattern of two to three broken chevrons crosses the entire shell.
Unio is a palearctic genus whose distribution in Africa is, as Brown (1978)
has pointed out, restricted to the cooier regions of the continent. There is thus
a wide gap between the ranges of U. caffer in South Africa, South West Africa
and Rhodesia and that of the nearest related species in North Africa.
Caelatura Conrad, 1853
Caelatura kunenensis (Mousson, 1887)
Fig. 2B-C
Material examined
Kunene River system:
Pump station, Huila (district), Angola, M. J. Penrith & J. Batista leg.
29 September 1972, 2 specimens, SMN-75540.
Pool below Ruacana Falls, B. F. Kensley leg. 1972, 50 specimens + 11
valves.
Ruacana Falls (17°23’S, 14°13’E), M.-L. Penrith leg. 1976, 10 specimens,
SMN-75598.
Great (Kambele) Falls (according to M. J. Penrith this is probably an old
name for the minor cataracts about 10 km above Ruacana), K. H.
Barnard leg. 1923, 2 valves, SAM-—A6640 and A6647.
44 km west of Ondarusu Falls, M. J. Penrith leg. 23-25 October 1971,
1 specimen, SMN-75494.
72 km west of Ondarusu Falls, M. J. Penrith leg. 28 October 1971, 1 speci-
men, SMN-75494.
Kunene River at Otjinungua, W. D. Haacke leg. May 1966, 2 specimens,
TM-9334 and 9335.
Cuvelai/Etosha system:
Ekuma floodplain, Etosha Pan National Park, K. L. Tinley leg. 27 Decem-
ber 1965, 1 specimen, NM-A5369.
Okavango River system:
Okavango River, G. C. Shortridge leg., 5 specimens, NM-3999; 3 speci-
mens, TM-38541.
THE UNIONACEA OF SOUTH-CENTRAL AFRICA 155
20mm
Fig. 2. Left valves of ten species of Unionacea from south-central Africa.
A. Unio caffer Krauss, 1848, Kafue River south of Itezhitezhi, Zambia. 62,5 <x 31,9 mm.
B. Caelatura kunenensis (Mousson, 1887), ‘typical form’, Thamalakane River in Okavango
delta, Botswana. 50,2 x 33,2 mm. C. Caelatura kunenensis (Mousson, 1887), ‘Zambezi form’,
Kalala Island in Kafue River, Zambia. 33,8 x 23,4 mm. D. Caelatura mossambicensis
(Martens, 1859), eastern end of Lake Kariba, Rhodesia. 50,8 x 27,0 mm. E. Caelatura
nyassaensis (Lea, 1964), off Nkopola, Lake Malawi, Malawi. 37,6 x 31,0 mm. F. Caelatura
hypsiprymna (von Martens, 1897), off Nkopola, Lake Malawi, Malawi. 37,3 x 28,3 mm.
G. Mutela dubia (Gmelin, 1793). Thamalakane River in Okavango delta, Botswana. 101,3 x
36,1 mm. H. Mutela alata (Lea, 1864). Lake Malawi, 87,7 x 41,0 mm. I. Aspatharia (Aspa-
tharia) pfeifferiana (Bernardi, 1860), upper Zambezi River, Zambia. 78,3 x 38,0 mm. J. Aspa-
tharia (Spathopsis) wahlbergi (Krauss, 1848), eastern end of Lake Kariba, Rhodesia. 102,5 x
49,2 mm. K. Aspatharia (Spathopsis) nyassaensis (Lea, 1864), Lake Malawi, Malawi.
91,5 x 56,4 mm.
156 ANNALS OF THE SOUTH AFRICAN MUSEUM
Omuramba Omataka, G. C. Shortridge leg., 22 specimens, MMK
MOL-327. Label reads: ‘everything collected in dry annually flooded
bed of Omuramba Omataka at or within 15 miles (23 km) of junction
with Okavango’. (The Omuramba Omataka is the largest, albeit
seasonal, watercourse flowing from South West Africa to the Okavango
River.)
Thamalakane River at Maun, Okavango delta, collector unknown, 20 May
1930, 181 specimens + 14 valves, TM-5565; same locality, P. A.
Reavell leg. 1973; 147 specimens.
Rakops on the Botletle River between the Okavango delta and the Makari-
kari Pan, collector unknown, 3 specimens.
Chobe/Linyanti River system:
Luapala Island in Linyanti swamp, 17°22’S 23°42’E ‘from shallow sand-
bank, water level dropped’. B. van der Waal leg. 1975, 1 specimen
+ 2 valves.
Zambezi River system:
‘Zambezi’, | specimen + 1 valve, NM-3371.
‘Zambezi’, B. E. Marshall don., 4 specimens.
Zambezi at Mongu Lealui, Barotse Valley, Zambia, 5 specimens + 1 valve,
NM-4000. Type locality of Nodularia croninae, synonym of
C. kunenensis.
Upper Zambezi River, G. C. Shortridge leg., 2 specimens -+ 21 valves,
MMK MOL-327.
Zambezi at Katima Mulilo, eastern Caprivi, W. D. Haacke leg. May 1966,
TM-11009; same locality, B. van der Waal leg. 1975, 3 specimens;
R. J. Pitchford leg. 1976, 2 valves.
Victoria Falls, K. H. Barnard leg. 1946, 2 specimens + 1 valve.
Kafue River, Kalala Island Rest Camp, Kafue National Park, Zambia,
A. C. van Bruggen leg. 2-3 November 1966, 3 specimens + 8 valves,
‘ex flood debris of Kafue River’, NM—A5366.
Kalala Island in Kafue River, Zambia, R. M. Derricourt leg. 1975, 26 speci-
mens + 8 valves.
Itezhitezhi area of Kafue River, Zambia, R. M. Derricourt leg. 1975,
10 specimens + 5 valves.
Although Haas (1936, 1969) included Unio zambeziensis Preston and Nodu-
laria croninae Walker from the Zambezi River system in the synonomy of this
species, two distinct forms are nevertheless recognizable, at least in adult shells.
A generally small, robust form, dark brown in colour and usually with zigzag
sculpture over most of the shell, corresponding to Unio zambeziensis (hereafter
referred to as the Zambezi form) from the Victoria Falls is also known from
the northern tributaries of the Zambezi as well as the Kunene River. A larger
though more compressed form, with a lower length/height ratio, light brown
THE UNIONACEA OF SOUTH-CENTRAL AFRICA 157
in the adult stage and with sculpture over much of the shell, occurs in the
Okavango River and its delta, and in the Linyanti swamp. This latter form,
according to Mandahl-Barth (in /itt. 13 October 1976), corresponds to typical
C. kunenensis. An apparently intermediate form occurs in the Zambezi River
at Katima Mulilo, the Okavango River and Ekuma floodplain. Shells referable
to Nodularia croninae are known from the upper reaches of the Zambezi in
eastern Zambia. Figure 3 shows a significant difference (p < 0,01) between the
shapes (as indicated by L/H v. length) of the Zambezi form and typical
C. kunenensis, particularly amongst larger specimens. There is no meaningful
difference in shell width relative to length and these dimensions are not shown.
The measurements given by Dartevelle (1939) for five specimens of C. kunenensis
from the upper Kunene (no precise locality given) indicate that these should
belong to the typical form.
Juveniles up to approximately 34 x 23 mm may be sculptured over the whole
shell surface. In larger shells the sculpturing persists posteriorly and anteriorly,
although specimens corresponding to N. croninae are generally smoother than
the more southerly forms. Juveniles of the Zambezi form are usually green-
yellow. The very small valves from the Kafue River (NM-—A5366) are dominated
by rugose and sometimes lamellate zigzag sculpture over the whole shell surface.
Nothing is known of the nature of the localities where shells corresponding
to N. croninae were found, but the Zambezi form and typical C. kunenensis
may represent adaptations to fundamentally different environmental conditions.
The present data indicate that the former occurs in lotic environments (perennial,
flowing rivers) whereas the latter is found in lentic and often seasonally
inundated areas. G. Mandahl-Barth has suggested (in /itt. 13 October 1976)
that the Zambezi form may, in fact, be a subspecies of C. kunenensis. Caelatura
. ¥=19346+0:0045X *
20 25 30 35 40 45 50 55 60
Length (mm)
Fig. 3. Caelatura kunenensis: relationship between L/H and length of river (Ruacana Falls,
Kunene River, n = 71) and swamp (Thamalakane River, Okavango delta, n = 145) forms as
shown by linear regression lines fitted by the method of least squares. Solid circles = Kunene
River; open circles = Okavango delta.
158 ANNALS OF THE SOUTH AFRICAN MUSEUM
kunenensis is thus a western species confined to the Kunene and Okavango
River systems as well as the upper Zambezi and its tributaries (Kafue, Luapala
and Linyanti) as far north as approximately 11°S latitude. The ‘typical’ form
seems allied to Caelatura choziensis (Preston) from Lake Bangweulu and the
Chambeshi River as illustrated by Haas (1936) and Mandahl-Barth (1972).
Although the gonads were not examined histologically, dissection of a
series of specimens from below the Ruacana Falls in the Kunene River showed
that 58 per cent (25/43) were functional females with all four demibranchs
modified to marsupia. The remaining 42 per cent (18/43) bore no sign of mar-
supia and were considered males. The smallest female measured 25,6 x 17,8 x
12,0 mm. Although most marsupia were charged with embryos some contained
sub-oval glochidia. The uniformly punctate valves of these glochidia were
virtually equal in length and height (Fig. 4); the mean dimensions of six examples
were 0,263 « 0,265 mm. No hooks were seen, although the glochidia may not
have been fully developed.
adductor
mantle
foot
Fig. 4. Camera lucida drawing of a glochidium
from an outer demibranch of Caelatura
kunenensis (38,6 < 23,2 x 17,6 mm) from
Ruacana Falls, Kunene River.
Small leeches, Batracobdella tricarinata (Blanchard) were found in the
pallial cavities of 16,3 per cent (7/43) of Caelatura kunenensis from Ruacana
on the Kunene River and a single emphemeropteran nymph (Baetidae: Cen-
troptiloides ? sp.) was present in the pallial cavity of a specimen from the Zambezi
at Katima Mulilo. Although this nymph is unlikely to be a ‘pallial cavity’
dweller, Centroptiloides nymphs are unusual among mayflies in being pre-
daceous (F. M. Chutter, in /itt. 31 January 1977). The presence of insect larvae
inside bivalves was also noted by Beedham (1971) who found chironomid larvae
between the mantle and shell of Anodonta cygnea (Linnaeus) (Unionidae) in
England. Woodward (1974) recorded numerous examples of a hydracarine
(though not a parasitic stage) within the pallial cavity of Caelatura cf. aegyptica
(Cailliaud) from Lake Chad.
THE UNIONACEA OF SOUTH-CENTRAL AFRICA 159
Caelatura mossambicensis (von Martens, 1860)
Fig. 2D
Material examined
Zambezi River system:
Eastern end of Lake Kariba in Nyanyana River estuary, Sanyati basin,
5-7 m in soft sand/silt, D. H. S. Kenmuir leg. 17 December 1976,
49 specimens.
Lake Macllwaine (on Hunyani River, tributary to Zambezi), B. E. Marshall
leg., 14 specimens + 14 valves.
Hunyani River at Darwendale, Rhodesia (below Lake Mcllwaine), B. E.
Marshall leg., 3 specimens.
This species may be distinguished from Caelatura kunenensis by being
relatively longer and more compressed, and by having a thinner shell with
poorly developed sculpture. In order to demonstrate the differences between
the shell width v. length ratio of C. mossambicensis and C. kunenensis, the
Lake Kariba mossambicensis were compared statistically to kunenensis from
three other localities, the Kafue River, Kunene River and Okavango delta.
Linear regression lines were fitted to each of the four sets of data by the method
of least squares and their slopes (not shown) compared in a pairwise manner
(Table 1). This showed that the shell width v. length ratio of all the C. kunenensis
populations differed significantly from C. mossambicensis, whereas there was
no significant difference between any of the individual C. kunenensis populations.
The three sets of data for C. kunenensis were then pooled and compared with
C. mossambicensis from Lake Kariba (Fig. 5). This showed a significant differ-
ence (p < 0,001) between the shell width v. length ratio for the two species.
TABLE 1
Pairwise comparisons between (A) Caelatura mossambicensis from Lake Kariba and
(B) C. kunenensis from the Kafue and Kunene rivers and the Okavango delta, using
the slopes of individual regression lines for shell width v. length.
A. Comparison between C. mossambicensis and C. kunenensis
degrees of
C. mossambicensis C. kunenensis t freedom significance
Lake Kariba Kafue River 2,1400 68 .
Lake Kariba Kunene River 3,1318 108 se
Lake Kariba Okavango delta 3,7227 191 shi
Lake Kariba three above
localities
combined 19,5344 279 wa
B. Comparison between three geographically different populations of C. kunenensis
degrees of
Localities t freedom significance
Kafue River Kunene River 1,0691 84 ns
Kafue River Okavango delta 1,0069 167 ns
Kunene River Okavango delta 0,1677 207 ns
* — significant at 5°% level
** — significant at 1% level
*** — significant at 0,1 % level
ns = not significant
160 ANNALS OF THE SOUTH AFRICAN MUSEUM
30
25
£0
He
3
=
15
10
25 30 35 40 45 50 55 60
Length (mm)
Fig. 5. Diagram relating shell width to shell length for Caelatura kunenensis (open circles)
and Caelatura mossanbicensis (solid circles) by means of linear regression lines fitted by the
method of least squares.
Juvenile C. mossambicensis show zigzag sculpture anterior and posterior
to the umbones, but in adults only a little remains visible. The colour of the
periostracum varies from green-brown to uniform dark brown while juveniles
may be light brown to greenish. Teeth are much weaker and less projecting than
in C. kunenensis.
The type locality of C. mossambicensis is Tete on the lower Zambezi.
It has also been recorded near Chemba (Germain 1935) and Mazarra (Azevedo
et al. 1961; Connolly 1939) and in Lake Macllwaine on the Hunyani River,
Rhodesia (Marshall 1975). In addition, it has been collected from Lake Malawi,
its tributary the South Rukuru River, and the Shire River which connects the
lake to the Zambezi (Mandahl-Barth 1972). Caelatura mossambicensis is closely
allied to Caelatura framesi (Connolly 19256) from further south (Brown 1967;
Appleton 1977), and Mandahl-Barth (in litt. 14 December 1976) is of the
opinion that C. framesi is merely a southerly form of C. mossambicensis. The
distribution of these two ‘species’ abut one another in central Mozambique.
Caelatura mossambicensis thus appears to be an eastern species confined to the
coastal peneplain of east and south-east Africa from the Kingani River near
Bagamoyo in Tanzania (about 6°25’S) (Haas 1962) to the Pongola River
floodplain (about 27°00’S) in north-eastern Natal (Appleton 1977; Brown 1967;
Pretorius ef al. 1975), and penetrating the Zambezi River system as far as
Lakes Kariba and Malawi.
THE UNIONACEA OF SOUTH-CENTRAL AFRICA 161
Caelatura nyassaensis (Lea, 1864)
Fig. 2E
Material examined
Monkey Bay, Lake Malawi, M. N. Bruton leg. March 1973, 1 specimen;
alive in sandy mud at about 3 m, C. C. Appleton & D. H. Eccles leg.
September 1976, 4 specimens.
Off Nkopola, Lake Malawi, alive in sand about 3 m, C. C. Appleton leg.
September 1976, 3 specimens + 1 valve.
A small, squat shell with very little or no sculpture, ventral margin curved
downward posteriorly and sharp umbonal ridge. The L/H ratio of the specimens
examined ranged from 1,24 to 1,40 (mean 1,32).
Caelatura hypsiprymna (von Martens, 1897)
Fig. 2F
Material examined
Monkey Bay, Lake Malawi, B. Boswell leg., 3 specimens + 1 valve,
TM-11239; alive in sandy mud at about 3 m, C. C. Appleton &
D. H. Eccles leg. September 1976, 4 specimens.
Off Nkopola, Lake Malawi, alive in sand about 3 m, C. C. Appleton leg.
September 1976, 8 specimens + 2 valves.
Another small, squat species with more chevron sculpture than C. nyassa-
ensis. This sculpture sometimes extends over most of the shell and is particularly
evident in smaller individuals which, as Mandahl-Barth (1972) has pointed out,
bear a close resemblance to the Zambezi form of C. kunenensis. The umbonal
ridge of C. hypsiprymna is pronounced though not sharp and the shell tends
to be rather attenuated posteroventrally. The L/H ratios of those examined
range from 1,13 to 1,53 (mean 1,34).
These two species appear to be endemic to Lake Malawi and have been
recorded from many localities in the lake and Shire River at Mangoche to
depths of 12,2 m (C. nyassaensis) and 79,3 m (C. hypsiprymna) (Crowley et al.
1964; Mandahl-Barth 1972). Mandahl-Barth (1972) gives a good description
of the differences between these species. To this it may be added that their
L/H ratios are lower than those of other Caelaturae of south-central Africa.
Family Mutelidae
Mutela Scopoli, 1777
Mutela dubia (Gmelin, 1793)
Fig. 2G
Material examined
Kunene River system:
Pool below Ruacana Falls, B. F. Kensley leg. 1973, 4 specimens.
162 ANNALS OF THE SOUTH AFRICAN MUSEUM
Okavango River system:
Okavango River, S.W.A., G. C. Shortridge leg. 5 specimens, NM-3912;
3 specimens + 1 valve, MMK MOL-336.
Thamalakane River at Maun, Okavango delta, donor unknown, 2 speci-
mens, TM-1075; P. A. Reavell leg. 1973, 42 specimens.
Chobe/Linyanti River system:
Lupala Island in Linyanti swamp, 17°22’S 23°42’E, ‘from shallow sand-
bank, water level dropped’. B. van der Waal leg. 1973, 7 specimens.
Mutualutze Lusese, a large permanent channel or ‘mulapo’ in eastern
Caprivi floodplain, 17°45’S 24°44’E. B. van der Waal leg. 1973,
1 specimen.
Zambezi River system:
Zambezi at Mongu Lealui, Barotse Valley, Zambia, Mrs. T. H. Howard
leg., 1 specimen, NM-3914.
Upper Zambezi, G. C. Shortridge leg., 1 specimen, MMK MOL-336.
Zambezi at Katimo Mulilo, B. van der Waal leg. 1975, 1 specimen.
Eastern end of Lake Kariba, Nyanana estuary in Sanyati basin, 5-7 m in
soft sand/silt, D. H. S. Kenmuir leg. 17 December 1976, 8 specimens.
Connolly (1939) identified Mutela mabilli (Rochebrune) from the Oka-
vango River but Haas (1936), after examining the same material, recorded it
as M. nilotica mabilli, a southern race of the north African M. nilotica. Later,
however, Haas (1969) placed M. mabilli into synonomy with the west African
M. dubia. Although the present material varies considerably in shape from
almost triangular to lanceolate, the majority of shells do, in fact, correspond
well to M. dubia as illustrated by Pilsbry & Bequaert (1927).
Both thin and thick (robust) shelled forms may occur in the same locality,
but the former appears to be the most common. The thick-shelled form from
the Okavango delta is on average 4,9 g heavier than the thin-shelled form of
similar length, and has a rather distinct shape (Fig. 6).
Mutela dubia is a west African species which reaches the southernmost
limits of its distribution in the Kunene, Okavango and Zambezi systems. The
only other record of M. dubia from the Kunene is a brief reference to this
species from the upper Kunene by Dartevelle (1939). The accuracy of the
record of Mutela rostrata (Rang) (now included in the synonomy of M. dubia
by Haas (1936, 1969)) from Lourengo Marques (Maputo) was questioned by
Connolly (1925a, 1939) and to the present author’s knowledge it remains the
only record of Mutela from Mozambique south of the Zambezi River. Small
leeches, Batracobdella tricarinata (Blanchard) (Branchiobdellidae), were found
in the mantle cavities, notably around the palps, of two Ruacana (Kunene)
specimens. The decimation of the littoral M. dubia population of Lake
Mcllwaine (Rhodesia) after a drop in water level (Marshall, 1975, 1978) may
have been due in part at least to the natural gape of the valves of this species
reducing the species’ resistance to dessication.
THE UNIONACEA OF SOUTH-CENTRAL AFRICA 163
60 70 80 90 100 no
Length (mm)
Fig. 6. Mutela dubia: scatter diagram relating L/H jx to valve length. The dotted line indi-
cates the robust form which constituted 22,3 per cent of M. dubia from the Okavango delta.
Solid triangles = Zambezi River; open triangles = eastern Caprivi/Linyanti floodplain;
solid circles = Okavango River system; open circles = Kunene River.
Mutela alata (Lea, 1864)
Fig. 2H
Material examined
Monkey Bay, Lake Malawi, M. N. Bruton leg. March 1973, 3 specimens;
C. C. Appleton leg. September 1976, 1 specimen.
Known from a variety of localities in Lake Malawi and the Shire River
to a depth of 27,5 m (Crowley et al. 1964; Mandahl-Barth 1972). Both thin-
and thick-shelled forms occur amongst the material available. Mutela alata
is characterized, as its name implies, by its high (alate) posterodorsal ‘wing’,
though this is not as high as that of Mutela hargeri (Smith) from Lake Mweru
(Zambia). The L/H ratio (2,06-2,36) tends to be lower than for M. dubia
(2,26-2,82) (Fig. 6).
This species is apparently endemic to Lake Malawi and the Shire River
although Mandahl-Barth (1972) considered that it is related to M. hargeri and
(in litt. 2 February 1977) to the widely distributed dubia—nilotica—mabilli group.
Aspatharia Bourguignat, 1885
Representatives of two subgenera, Aspatharia s.s. and Spathopsis, occur
in southern central Africa and may be distinguished by the character of the
inhalent opening into the branchial chamber. In Aspatharia s.s. there is a
distinct inhalent aperture formed by a fusion of the mantle edges, while in
Aspatharia (Spathopsis) there is no such fusion and the mantle margins thus
164 ANNALS OF THE SOUTH AFRICAN MUSEUM
remain free below the exhalent aperture (Fig. 7). In this respect Aspatharia s.s.
is unique among the southern and central African Unionacea.
A
Fig. 7. Digrammatic illustration comparing the inhalent apertures of Aspatharia (Aspatharia)
and Aspatharia (Spathopsis). A. Aspatharia s.s. lacks a discrete aperture and the inhalent
current is simply drawn between the mantle edges into the branchial chamber. B. Aspatharia
(Spathopsis) possesses a distinct inhalent aperture through which the current enters the
branchial chamber.
Aspatharia (Aspatharia) pfeifferiana (Bernardi, 1860)
Fig. 21
Material examined
Kunene River system:
Pool below Ruacana Falls, B. F. Kensley leg. 1973, 1 specimen.
Ontoruthu Falls (= Ondorusu Falls, 17°24’S 13°57’E), B. F. Kensley leg.
1972, 4 specimens ++ 2 valves.
Otjinungua, Kaokoveld, W. D. Haacke leg. May 1966, 1 specimen,
TM-9335.
Okavango River system:
Okavango River, South West Africa, G. C. Shortridge leg., 6 specimens,
NM-3913.
Omuramba Omataka, G. C. Shortridge leg., 7 specimens, MMK MOL-334.
Boro River, Okavango delta, P. A. Reavell leg. 1973, 3 specimens + 5
valves.
Zambezi River system:
Upper Zambezi Valley, G. C. Shortridge leg., 7 specimens, MMK
MOL-334.
Sandbank in Zambezi at Katima Mulilo, B. van der Waal leg. 1974,
1 specimen.
Itezhitezhi area of Kafue River, Zambia, R. M. Derricourt leg. 1975,
1 specimen + 3 valves.
THE UNIONACEA OF SOUTH-CENTRAL AFRICA 165
Records of this smallish species from the Kunene River at Fort Rocados
in southern Angola, the Okavango River in Owambo, and the Hunyani River
(a tributary of the Zambezi) in Rhodesia were cited by Haas (1936).
Adpatharia pfeifferiana is a predominantly west African species reaching the
southern limits of its range in this area. In several localities it was found
together with A. (Spathopsis) wahlbergi from which it may be distinguished by
its concave ventral margin and much smaller pedal protractor muscle scar.
Empty shells of A. pfeifferiana can usually be distinguished from those of
A. wahlbergi as shown in Figure 8. This relates the L/H (ag umbone) Tatio to
ad umbone
ae
30 40 50 60 70 80 90 100 110 120
valve length (mm)
Fig. 8. Scatter diagrams relating the L/H ag umbone ratio to valve length for Aspatharia (A.)
Pfeifferiana (solid symbols) and Aspatharia (S.) wahlbergi (open symbols). Circles = Kunene
River; triangles = Kafue River; inverted triangles = Zambezi River system;
squares = Okavango River system.
valve length; umbonal height is used despite shell erosion in order to take into
account the marked concavity of the ventral margin (below the umbonal area)
typical of A. pfeifferiana.
The only other member of Aspatharia s.s. recorded from south-central
Africa is A. (A.) subreniformis (Sowerby) from Lake Malawi and the Shire
River. Mandahl-Barth (1972) doubts that this represents a valid species endemic
to the lake and has suggested that it may be closely related to A. (A.) pfeifferiana
(Bernardi) (= A. rugifera (Dunker) fide Haas 1969).
166 ANNALS OF THE SOUTH AFRICAN MUSEUM
Aspatharia (Spathopsis) wahlbergi (Krauss, 1848)
Fig. 2J
Material examined
Kunene River system:
Ruacana Falls, M. L. Penrith leg. 1973, 6 specimens, SMN-75597.
Pool below Ruacana Falls, B. F. Kensley leg. 1973, 4 specimens + 1 valve.
Okavango River system:
Nata River valley, between Maitengwe and Nata Police Station, Central
district Botswana, donor unknown, 3. specimens + 4_ valves,
TM-11237.
Zambezi River system:
Eastern end of Lake Kariba, Nyanana estuary in Sanyati basin, 5-7 m in
soft sand/silt, D. H. S. Kenmuir leg. 17 December 1976, 13 specimens.
Battlefields (on Sanyati River), Rhodesia, F. B. Parkinson leg., 1 specimen,
NM-2703; | specimen, NYVK MOL-335.
Kalala Rest Camp, Kafue National Park, Zambia, ‘ex flood debris of
Kafue River’, A. C. v. Bruggen leg. 2-3 November 1963, 1 valve,
NM-AS5365.
Itezhitezhi area of Kafue River, Zambia, R. M. Derricourt leg. 1975,
1 specimen + 3 valves.
Monkey Bay, Lake Malawi, B. Boswell leg. July 1968, 2 specimens,
TM-11238.
Juveniles may display up to thirteen irregular riblets on the posterodorsal
surface of the shell. They generally have a light-brown periostracum, but this
darkens with age and adult specimens are dark brown to black. A. wahlbergi
is also known from Mupa on the Cuvelai River, southern Angola (Haas 1936).
This large species is widely distributed in central and east Africa from the
Kunene River in the west to Mozambique in the east and thence southwards to
the Transvaal and Zululand. Except for a single high-shelled juvenile from the
confluence of Omuramba Omataka and Okavango rivers (Connolly 1931), and
several specimens from the Nata River which flows into the Makarikari pan,
A. wahlbergi has not been collected from the Okavango River system. Connolly’s
(1931) description of the singleton from South West Africa (L/H 1,71) is
reminiscent of the Kunene River specimens whose L/H ratios (1,75-1,86,
mean 1,82) are low for the species and which probably correspond to the
variety moutai described by Dartevelle (1939) from the upper Kunene. The
Nata River shells are much larger (to 120,9 x 62,1 mm, mean L/H 1,91),
resembling those from the Limpopo River system whose drainage abuts on that
of the Nata River. A. wahlbergi was not recorded from Lake Malawi itself by
either Crowley et al. (1964) or Mandahl-Barth (1972) and its occurrence here is
regarded as doubtful. Commenting on this, the latter author noted the species’
presence in a roadside pool 5 km south of Monkey Bay. The shells (empty)
THE UNIONACEA OF SOUTH-CENTRAL AFRICA 167
collected by Boswell (TM-11238) may have originated from a similar water
body in the vicinity of Monkey Bay.
Braga (1952) illustrated a rather attenuated shell from Lagoa Bamba, Chibuto,
Mozambique (L/H 2,36) which, although he gave no width measurement,
appears otherwise comparable with the unusually obese specimens collected
from Lagoa Pavi near Inhambane (MMK MOL-335) by H. A. Junod (L/H
2,13-2,21, mean 2,19) (not the same as those, also collected by Junod from
Lagoa Pavi, recorded by Connolly 1925a). These shells may represent a lacus-
trine form occurring in the freshwater coastal lakes of the Mozambique
peneplain.
The leech Batracobdella tricarinata (Blanchard) (Branchiobdellidae) was
found in the mantle cavities of two specimens from Ruacana in the Kunene
River.
Aspatharia (Spathopsis) nyassaensis (Lea, 1864)
Fig. 2K
Material examined
‘Lake Malawi’, donor unknown, | specimen NM-A5367.
Mangoche (Fort Johnston) on Shire River between Lakes Malawi and
Malombe, K. Fuller leg. 19 November 1971, | specimen.
Monkey Bay, Lake Malawi, M. N. Bruton leg. March 1973, | specimen;
C. C. Appleton & D. H. Eccles leg. September 1976, live in sandy mud
at about 3 m, | specimen.
An unmistakable large, thick-shelled species which is endemic to Lake
Malawi and the Shire River (Mandahl-Barth 1972). It is much higher posteriorly
than the other large Aspatharia, A. wahlbergi, and the specimens to hand have
a lower L/H ag umbone Fatio of 1,70—1,96.
Aspatharia (Spathopsis) petersi (von Martens, 1859)
Although no specimens of this small, slender species are available from the
area under discussion, Haas (1936) collected it from the Hunyani River 20 km
south of Salisbury, Rhodesia. Connolly (1939) cites a record from Tete (the
type locality) which lies on the lower Zambezi. A. petersi may be distinguished
from A. wahibergi by its much longer and narrower shell and hence its high
L/H ratio of between 2,2 and 2,6 as opposed to 1,8 and 2,1 of the latter.
The distribution of this species is not well documented but it appears to
be confined to the lower Zambezi as well as the eastwards flowing rivers of
south-eastern Africa to about latitude 27°21'S. Specimens collected by B. Davies
(Rhodes University) from Lake Tchal on the Incomati floodplain in Mozam-
bique are, like A. wahlibergi from lakes in this region, longer in relation to
height (L/H 2,30-2,57, mean 2,46) and more compressed than their counterparts
from riverine environments. Data to hand suggest that the L/H ratios of shells
from rivers range from 2,19 to 2,41 (mean 2,28).
168 ANNALS OF THE SOUTH AFRICAN MUSEUM
Family Etheriidae
Etheria Lamarck, 1807
Etheria elliptica Lamarck, 1807
Material examined
Kunene River:
Kunene River at Otjinungua, Kaokoveld, W. D. Haacke leg. May 1966,
1 specimen + 2 valves, TM-—93332.
Ontoruthu Falls (= Ondorusu Falls, 17°24’S 13°57’E), B. Kensley leg.
1973, 9 valves.
Yonge (1962) has discussed in detail the anatomy, evolution and life
history of Etheria elliptica and concurs (Yonge 1962, 1978) with Haas (1969)
and Thiele (1935) that E. elliptica is closely related to two other monotypic
genera, the Colombian Acostaea and the Indian Pseudomulleria, and that these
three comprise the family Etheriidae. The largest lower valve encountered
measured 99,2 x 49,0 mm and the largest upper valve, 119,2 « 82,4 mm.
One of the twelve upper valves available has the tube-like outgrowths (costae)
typical of E. elliptica f. tubifera Sowerby. The only other record of E. elliptica
from the Kunene River is by Germain in 1925 (cited by Haas 1936).
Only empty valves are available but M. J. Penrith (State Museum, Wind-
hoek) has seen live specimens. He notes (in /itt. 11 October 1976) that E. elliptica
is very common in the Kunene from below the Ruacana Falls to the Foz du
Cunene, where the last outcrops of rock occur (7 km from the mouth), and that
it is usually found on the large boulders which in places form either banks or
rapids in the river. However, it does not occur where flow is very fast, i.e. the
shallower and narrower rapids. At the Ontorusu Falls the highest living animals
were only a few centimetres below normal low water. He doubts whether the
species occurs upstream of Ruacana and, if not, it would possess a distribution
pattern which parallels that of some fish species. Dartevelle (1939) did not
record E. elliptica from the upper Kunene.
Yonge (1962) noted that E. e/liptica could be cemented to the substrate by
either valve and that this occurred relatively late in its growth. The equivalve
post-larva probably became attached (in the vertical position) initially by a
byssus enabling it to withstand the high current velocities in which the adults
are usually found. The later adoption of a horizontal posture was followed by
cementation and growth as an inequivalve.
E. elliptica is widely distributed in tropical Africa and the Kunene shells
represent its southernmost known locality.
DISCUSSION
Virtually nothing is known of the ecology of the freshwater bivalves of
south-central Africa nor, indeed, of Africa generally. This is despite the fact
that, although poor in numbers of species, the Unionacea of the river systems
THE UNIONACEA OF SOUTH-CENTRAL AFRICA 169
under discussion are among the largest members of the benthos. Indeed, they
may attain high population densities and may therefore constitute an important
proportion of the benthic biomass. Relevant ecological studies include those of
Pilsbry & Bequaert (1927) on the general malacology of the Congo River
system, Fryer (1959, 1961) on the life-cycle of Mutela bourguignati (Ancey) in
Lake Victoria, Marshall (1975) on the composition and distribution of the
Unionacea of Lake Macllwaine in Rhodesia, and Yonge (1962, 1978) on the
settlement and cementation of Etheria elliptica. It is thus useful here to quote
in full the locality notes accompanying specimens from the Kunene River and
the Okavango delta (identifications inserted by the author).
M. J. Penrith (in /Jitt. 11 May 1976) described the Kunene as a rather
silt-laden river whos bed alternates between sand and rock. In sandy localities
the channel is edged with Phragmites. Where rock is present it is either bare
where the river is wide and slow-flowing (favourable Etheria habitats), or
carpeted with Hydrostachis where it is fast flowing. There are fishes below the
Ruacana Falls having affinities with rivers such as the Curoca and Quanza in
Angola to the north.
At Ruacana B. F. Kensley collected four bivalve species from a single pool
below the falls. His notes read: ‘Hippo pool—small, muddy pool on Angolan
side of river, bounded by sandy bar on the flowing river side and by reeds on
the landward side. Maximum depth 0,9 m. Charophyte covering most of the
bottom which consisted of a 2,5 cm layer of ooze, coarse sand below being
black and anaerobic. Caelatura kunenensis most abundant, 6—10/m?, in the
ooze layer, Mutela dubia also lies (horizontally) in the first 2,5 cm of ooze.
Aspatharia pfeifferiana and A. wahlbergi lay (vertically) about 7,6 cm below
sand surface. Mutela and Aspatharia spp. were less common than C. kunenensis,
seldom more than |/m?*. Substrate surface scattered with dead shells, especially
C. kunenensis. Openbill storks (Anastomus lamelligerus Temminck) observed
feeding on bivalves. No water movement through pool and temperature higher
than in flowing river.’
P. A. Reavell, who made a collection of bivalves near Maun in the Okavango
delta, wrote: ‘All specimens taken from the seasonally inundated area below
perennial swamps. Caelatura kunenensis (typical form) prefers soft clay bottoms
of streams in seasonally inundated area. Very common along Thamalakane
River where in the dry season water may become stagnant with very low
dissolved oxygen content. Mutela dubia usually together with C. kunenensis
but more sporadic in distribution and of lower population density. Aspatharia
pfeifferiana found only on clean, sandy bottoms where water was always flowing
and dissolved oxygen content always high.’ It is perhaps noteworthy that in
many large C. kunenensis and M. dubia shells from the Thamalakane River
erosion of the prismatic layer round the umbones and dorsal areas is unusually
severe. This may be due to the action of aggressive carbon dioxide in the stagnant
conditions which Reavell noted developed here.
These notes suggest that, while the bivalve species concerned are tolerant
170 ANNALS OF THE SOUTH AFRICAN MUSEUM
of a wide range of dissolved oxygen concentration, C. kunenensis and M. dubia
may prefer fine substrates such as clay and ooze whereas Aspatharia spp. may
prefer coarser conditions.
Marshall (1975) found that in Lake Macllwaine, a man-made impoundment
on the Hunyani River in Rhodesia, Mutela mabilli (? = M. dubia) and Caelatura
mossambicensis attained their greatest population densities (31/m? and 3/m?
respectively) and largest sizes in the riverine part of the lake. This was tentatively
taken as evidence that these bivalves remain essentially riverine animals after
impoundment, an interpretation in agreement with Fryer (1961) who found that
the fish Barbus altianalis radcliffi was more heavily parasitized by larval Mutela
bourguignati in the Victoria Nile than in Lake Victoria itself. This may have
been due to the fish being more abundant in the riverine environment and
therefore easier for the larvae to locate. If so, it suggests an important role for
host fish in determining the distribution and population density of at least some
Unionacea in lacustrine environments. This aspect is also discussed by Jubb
(1976) who adds that U. caffer glochidia probably parasitize both truly fresh-
water fish species and those which are capable of moving from fresh to saline
water. Thus U. caffer could, when parasitic on fish of the latter type, i.e. the
widespread Sarotherodon mossambicus (Peters), be carried from one river system
to another via the sea, suggesting that spread of species from one drainage to
another might be effected by processes other than geomorphic ones (river
capture). He notes, too, that adult U. caffer are tolerant of salinities up to
3°/o0 but not 5°/oo.
The large perennial Kunene, Okavango and Zambezi rivers rise on the
southerly part of the central Angolan highlands at about | 700 m altitude; the
smaller Cuvelai River, more seasonal in character, rises on the southernmost
spur of this plateau, the Serra Encoco. Over much of the drainage of these
rivers the topography is very flat, with gradients of 1:30 000 or less. While the
Kunene flows westwards to the Atlantic Ocean and the Zambezi eastwards to
the Indian Ocean, the intervening area includes three endoreic or inland drainage
systems, the Cuvelai/Ekuma/Etosha, the Cubango/Okavango/Ngami/Makari-
kari and the Chobe/Liambezi/Linyanti. Where their beds are of sand or clay,
these rivers broaden to create extensive floodplains where, in years of high
rainfall, numerous low gradient valleys and channels (known in different areas
as oshanas, omurambas or mulapos) become flooded and inundate vast areas of
low-lying land. Water velocities in these seasonal watercourses seldom exceed
0,5 m/sec. (Stengel 1963: 375).
As far as a zoogeographical consideration of this malacofauna is possible,
it is evident that the Kunene, Okavango and upper Zambezi River systems not
only have several bivalve species in common but also constitute the southerly
limits of the ranges of three west African Mutelidae, Mutela dubia, Aspatharia
Pfeifferiana and Etheria elliptica. Of the twelve unionacean species recorded
from these systems the three Mutelidae listed above are predominantly west
African species. Dartevelle (1939) attributed the similarity between the species
THE UNIONACEA OF SOUTH-CENTRAL AFRICA 171
occurring in the upper Kunene and the Zambezi to capture of part of the
drainage of the latter by the Kunene. Caelatura kunenensis is endemic to the
western half of south-central Africa while C. mossambicensis and Aspatharia
petersi are endemic to the eastern half. Unio caffer is a southern African endemic
and Aspatharia wahlbergi enjoys a wide distribution in southern and east Africa.
Caelatura nyassaensis, C. hypsiprymna, Mutela alata and Aspatharia nyassaensis
are endemic to Lake Malawi.
Several examples exist of morphologically separable riverine and lacustrine
forms of a species, e.g. Caelatura kunenensis, Aspatharia wahlbergi and A. petersi.
Enough data are available to show (Fig. 3) that for C. kunenensis these differ-
ences are statistically significant (p < 0,01). This is probably a widespread
phenomenon among freshwater Mollusca. Sutcliffe & Durrant (1977) have
shown highly significant differences (p < 0,001) between the shell shapes of
riverine and lacustrine populations of the limpet Ancylus fluviatilis (Muller) in
England.
Similarities among the unionacean fauna of the river systems included in
this study are probably due to connections between them during the Tertiary
era (Gaigher & Mc. C. Pott 1973; Netterberg 1973) and to river capture.
Connections still occur between the Okavango and upper Zambezi systems
after exceptionally rainy seasons, when the waters of the Okavango flow via
the Chobe/Linyanti rivers and the eastern Caprivi floodplain to the Zambezi
at Kazungula, some 100 km upstream from the Victoria Falls. This floodplain
is extremely flat, with a maximum height difference of only 20 m, so that in
high flood the Zambezi may overflow in the reverse direction (Van der Waal
1974). Floods may occur more than once in each direction in a single rainy
season.
Although the Kunene does not today overflow into Owambo even at
high flood (Stengel 1963) there was before the breakthrough at the Ruacana
Falls a previous connection with the Cuvelai/Etosha Pan system. Unfortunately,
little bivalve material is available from this latter system. The upper tributaries
to the Kunene, Okavango and Zambezi rivers not only lie close together, but
their watersheds are ill-defined so that, in the dry seasons, it is difficult to tell
in which direction they would flow (Wright 1963). Since these drainages were
probably frequently connected during the pluvial periods of Pleistocene times,
the spread of freshwater bivalves between them could readily occur. Fish might
convey the parasitic stages of C. kunenensis and the Mutelidae from one drainage
to another. Floods, too, can transport live bivalves. The author has found
numerous large live Unio caffer scoured from the mud substratum of pools in a
stream bed and carried by floods for considerable distances both downstream
and away from its banks.
The apparent limits in the upstream distribution of E. elliptica in the
Kunene River at the Ruacana Falls and of C. mossambicensis in the Zambezi
at the Victoria Falls suggests that either large waterfalls or, since Ruacana is
the most westerly of a series of falls, several smaller ones, may play important
172 ANNALS OF THE SOUTH AFRICAN MUSEUM
roles in limiting bivalve distribution. This may be due to changes in substrate
or to the barriers imposed by waterfalls on host fish distribution.
Except for Caelatura mossambicensis, and perhaps the little known Aspa-
tharia (A.) subreniformis as well, the unionacean fauna of Lake Malawi consists
of endemic forms with noticeably heavy shells (Mandahl-Barth 1972). This
fauna seems to have affinities with the river systems to the west and south rather
than with the other African Great Lakes to the north. Little is known of the
Unionacea of the Shire River nor of how far south the Lake Malawi endemics
are distributed within it. However, most occur in Lake Malombe as well and
it is possible that the Murchison Falls (Shire River) act as a barrier here.
ACKNOWLEDGEMENTS
The assistance of the directors of the following institutions who made
their collections available for study is gratefully acknowledged: South African
Museum, Cape Town; Natal Museum, Pietermaritzburg; Transvaal Museum,
Pretoria; McGregor Memorial Museum, Kimberley; and State Museum,
Windhoek. The following collectors have also made material available: Mr P. A.
Reavell (University of Zululand), Mr B. van der Waal (Dept. of Agriculture and
Works, eastern Caprivi), Dr R. M. Derricourt (National Monuments Commis-
sion of Zambia), Dr M. N. Bruton (Rhodes University), Mr D. H. S. Kenmuir
(Lake Kariba Fisheries Research Institute), and Mr B. E. Marshall (Dept. of
National Parks and Wildlife Management, Rhodesia). Dr F. M. Chutter
(National Institute for Water Research) and Dr J. H. Oosthuizen (University
of Pretoria) identified ?Centroptiloides sp. and Batracobdella tricarinata respec-
tively. Thanks are also due to Mr M. J. Andrew (Institute of Biostatistics) for
analysing the data presented in Figures 3 and 5, to Dr G. Mandahl-Barth
(Danish Bilharziasis Laboratory) for his ready advice on various taxonomic
issues, and to the director of the McGregor Memorial Museum for permission
to publish Figure 2]. Dr R. C. Lethbridge (Murdoch University) kindly read
th manuscript which is published with the permission of the South African
Medical Research Council.
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I1I—Moluscos de Mocgambique. Estudos Ensaios Docum. Jta Invest. Ultramar, Lisboa
88: 1-394.
BEEDHAM, G. E. 1971. The extrapallial cavity in Anodonta cygnea (L.) inhabited by an insect
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BERNARDI, A. 1860. J. Conch., Paris 4: 331.
BourGuiGnatT, J. R. 1885. Espéces nouvelles et generes nouveaux découverts par les Rév. Peres
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THE UNIONACEA OF SOUTH-CENTRAL AFRICA 173
BraGA, J. M. 1952. Materias para o estudo da fauna malacologica de Mogambique. An. Jta
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Brown, D. S. 1967. A review of the freshwater Mollusca of Natal and their distribution.
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261-267.
CoNNOLLY, M. 1931. Contribution to a knowledge of the fauna of South West Africa IX. The
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CONNOLLY, M. 1939. A monographic survey of the South African non-marine Mollusca.
Ann. S. Afr. Mus. 33: 1-660.
CROWLEY, T. E., PAIN, T. & WOODWARD, F. R. 1964. A monographic review of the Mollusca
of Lake Nyasa. Annis. Mus. Afr. cent. Sér. 8vo. Zoologie 131: 1-58.
DaRTEVELLE, E. 1939. Quelques mollusques fiuviatiles du Cunene. J. Conch., Paris 83: 327-331.
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Fryer, G. 1961. The developmental history of Mutela bourguignati (Ancey) Bourguignat
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GAIGHER, I. G. and Mc. C. Pott, R. 1973. Distribution of fishes in southern Africa. S. Afr.
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of Lake McIlwaine, Rhodesia. Arnoldia 7: 1-15.
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OBERHOLZER, G. & VAN EEDEN, J. A. 1967. The freshwater molluscs of the Kruger National
Park. Koedoe 10: 1-42.
174 ANNALS OF THE SOUTH AFRICAN MUSEUM
Pirspry, H. A. & BEQUAERT, J. 1927. The aquatic mollusks of the Belgian Congo, with a
geographical and ecological account of Congo malacology. Bull. Amer. Mus. nat. Hist. 53:
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freshwater Mollusca of the Pongola River floodplain pans. S. Afr. J. Sci. 71: 208-212.
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and the development of fishery. 2nd report, Oct. 1973-Aug. 1974. Katima Mulilo: Dept.
Agriculture & Works, Eastern Caprivi Government Service.
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Notes on the anatomy of Caelatura sp. from Lake Chad, Northern Nigeria. Revue Zool.
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(Zoology) 10: 449-528.
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tion of monomyarianism, in the family Etheriidae (Bivalvia: Unionacea). Phil. Trans.
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YONGE, C. M. 1978. On the monomyarian, Acostaea rivoli and evolution in the family
Etheriidae (Bivalvia: Unionacea). J. Zool., Lond. 184: 429-448.
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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.
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An author’s name when cited must follow the name of the taxon without intervening
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species or subspecies is transferred from its original genus. The name of a subsequent user of
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Synonymy arrangement should be according to chronology of names, i.e. all published
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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:
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Holotype
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Biological Abstracts.
C. C. APPLETON
THE UNIONACEA
(MOLLUSCA, LAMELLIBRANCHIATA)
OF SOUTH-CENTRAL AFRICA
S- Hy - ajcetow ste
VOLUME 77 PART 10 JUNE 1979 ISSN 0303-2515
MUS. COMP. ZOOL,
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BuLLOuGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FiscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gen. 74: 627-634.
Kon, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.
Konn, A. J. 19606. Spawning behaviour, i sa aes and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4):
Turee, J. 1910. Mollusca: B. Seiatemainon: Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 77 Band
June 1979 Junie
Part 10 £Deel
UPPERMOST ALBIAN (STOLICZKAIA DISPAR ZONE)
AMMONITES FROM THE ANGOLAN LITTORAL
By
M. R. COOPER
&
W. J. KENNEDY
Cape Town Kaapstad
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UPPERMOST ALBIAN (STOLICZKAIA DISPAR ZONE)
AMMONITES FROM THE ANGOLAN LITTORAL
By
M. R. CooPER
Queen Victoria Museum, Salisbury
&
W. J. KENNEDY
Geological Collections, University Museum, Oxford
(With 82 figures)
LMS. accepted 11 December 1978]
ABSTRACT
Rich, well-preserved collections of late Albian (Stoliczkaia dispar Zone) ammonites from
the Angolan littoral provide the basis for discussion of the ontogenetic and intraspecific
variation within S. (Stoliczkaia) tenuis Renz, Tetragonites jurinianus (Pictet), Eogaudryceras
italicum Wiedmann & Dieni, Desmoceras latidorsatum perinflatum subsp. nov., Phylloceras
(Hypophylloceras) seresitense Pervinquiére, Anisoceras haasi sp. nov., and Mortoniceras
(Durnovarites) collignoni sp. noy. Other important elements present include Jdiohamites
dorsetensis Spath, I. pygmaeus sp. nov., I. cf. elegantulus Spath, Anisoceras armatum
(J. Sowerby), A. perarmatum Pictet & Campiche, A. phillipsi sp. nov., Mortoniceras (Durno-
varites) perinflatum (Spath), M. (D.) subquadratum Spath, M. (Angolaites) simplex (Choffat),
M. (A.) gregoryi (Spath), Cantabrigites? curvatum Renz, Hysteroceras? cf. ootaturense
(Stoliczka) and Borissiakoceras sp. nov. ? aff. reymenti (Brunnschweiler).
The faunas form the basis for discussion of the subdivisions of the Stoliczkaia dispar Zone,
and a return to the simple division into subzones of S. (Faraudiella) blancheti below and
M. (Durnovarites) perinflatum above is proposed.
CONTENTS
PAGE
Introduction . : : : 2 : : ; «. Lib
Systematic palaeontology . : P : : ee i 8
Family Phylloceratidae p , : : Re ae) WS i
Genus Phylloceras . : F : ‘ eo TT
Family Tetragonitidae . : , : : St
Genus Tetragonites : : : : . 181
Family Gaudryceratidae . : ; ‘ . 489
Genus Eogaudryceras . ‘ : : ; 189
Family Ancyloceratidae : ‘ A : Oo
Genus Anisoceras . : A , ; ‘ 196
Genus Idiohamites . : ‘ ; ; ee:
Family Hamitidae . : 4 . 5 : at 26
Genus Hamites . P : , ‘ » tees
Family Turrilitidae : : : : ; < eee
Genus Mariella. ; ; : ; . 228
Family Desmoceratidae . ‘ ‘ : a 28
Genus Puzosia ; : : , ; ee
Genus Desmoceras ~ : : : 7) ma
175
Ann. S. Afr. Mus. 77 (10), 1979: 175-308, 82 figs.
176 ANNALS OF THE SOUTH AFRICAN MUSEUM
Systematic palaeontology (cont.) PAGE
Family Lyelliceratidae . : : = out el
Genus Stoliczkaia . . ; : : . 243
Family Brancoceratidae ‘ : : ; ~ 265
Genus Hysteroceras? . : : : » 265
Genus Mortoniceras . ‘ j : . 269
Subgenus M. (Durnovarites) : F . 269
Subgenus M. (Angolaites) . : : - 285
Genus Cantabrigites? . : : : . 290
Genus Drakeoceras : ‘ ; : e293
Family Binneyitidae . : ‘ ‘ : . 296
Genus Borissiakoceras . : : : » . 296
Age of the fauna . ; P : : , : z - 298
Summary : : F ; : : : , = » S01
Acknowledgements : : : : ; : . 303
References. : : ‘ : : : : = 303
INTRODUCTION
Although Albian ammonites have long been known from Angola
(Szajnocha 1884; Meunier 1887; Choffat in Choffat & De Loriol 1888), Paul
Choffat was the first to record uppermost Albian ammonites when he described
Ammonites dispar Choffat (non d’Orbigny) (= Stoliczkaia tenuis Renz) from the
environs of Catumbela. Subsequently, many authors have reported on the
presence of strata with Ammonites dispar (Mouta & Borges 1926, 1928; Mouta
1937, 1954, 1956; Mouta & O’Donnell 1933) and the presence of uppermost
Albian strata in Angola is now well established. In contrast, the faunal
constituents of these rocks are poorly known.
The collections which form the basis of this paper are from Porto Amboim,
Cabo Ledo and Praia-Egito. The material from Porto Amboim comes from the
lime kiln at the south side of the boat landing, and was collected mostly by
C. W. Washburn on 22 July 1914, although some material from this locality, in
the South African Museum, was described by Haughton (1924, 1925). Elements
of this fauna were also described subsequently by Haas (1942), whose localities
R28, R30 and 3041 have yielded an identical fauna in like preservation, with
recrystallized test preserved. The fauna from Egito was collected by the senior
author and comes from the coastal exposures at Praia-Egito where thinly-bedded
limestone-shale alternations have yielded a rich fauna, mostly preserved as
composite internal moulds. Faunas collected by the senior author from the
Quissama Ridge, the promontory on which stands the Farol de Cabo Ledo,
were obtained from road gravels and may represent more than one faunal
horizon.
These faunas include a number of new species, and also populations, of
S. (Stoliczkaia), Mortoniceras (Durnovarites), E. (Eogaudryceras), Phylloceras
(Hypophylloceras), D. (Desmoceras), Anisoceras and Tetragonites which provide
unique information on the intraspecific variation, development and morphology
of these ammonites. Furthermore, the composition of the faunas provides an
opportunity for discussion of the stratigraphic and geographic distribution of
the late Albian Stoliczkaia dispar Zone fauna.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 177
SYSTEMATIC PALAEONTOLOGY
The following abbreviations are used to indicate the source of material:
BM _ British Museum (Natural History), London
SAM_ South African Museum, Cape Town
USNMNH USS. National Museum of Natural History, Washington D.C.
Measurement abbreviations are as follows: D, diameter; H, whorl! height;
W, whorl width (intercostal unless otherwise specified); U, diameter of umbilicus
between umbilical seams; “/H, whorl width/height ratio. All measurements are
given in millimetres and dimensions, as a percentage of the diameter, follow in
parentheses.
Class CEPHALOPODA Cuvier, 1797
Subclass AMMONOIDEA Zittel, 1884
Order PHYLLOCERATIDA Arkell, 1950
Superfamily PHYLLOCERATACEAE Zittel, 1884
Family Phylloceratidae Zittel, 1884
Subfamily Phylloceratinae Zittel, 1884
Genus Phylloceras Suess, 1865
Subgenus Hypophylloceras Salfeld, 1924
Type species Phylloceras onoense Stanton, 1895
Phylloceras (Hypophylloceras) seresitense Pervinquiére, 1907
Figs 1-2, 31
Ammonites velledae Pictet & Campiche (non Michelin), 1860: 268, pl. 36 (fig. 8). Stoliczka,
1865: 116, pl. 59 (figs 1-3).
Phylloceras velledae (Michelin) Kossmat 1895: 12, pl. 1 (fig. 3). Boule, Lemoine & Thévenin,
1906: 7, pl. 1 (figs 6, 11), fig. 2. Crick, 1907: 166, pl. 10 (figs 10-11). Bése, 1923: 119,
pl. 7 (figs 15-17). Spath, 1925b: 180. Besairie, 1936: 164, pl. 16 (fig. 1). Venzo, 1936: 66,
pl. 5 (fig. 4). Haas, 1942: 146, pl. 27 (fig. 1), pl. 44 (fig. 1), fig. 19. Matsumoto, 1942: 676.
Almela & Revilla, 1957: 17, pl. 3 (fig. 2). Anderson, 1958: 180, pl. 16 (fig. 4).
Phylloceras velledae var. seresitense Pervinquiére, 1907: 52; 1910: 9, pl. 1 (figs 1-3), fig. 2.
Phylloceras tanit Pervinquiére, 1907: 53, pl. 3 (figs 3-9), fig. 5.
Phylloceras angolaense Haughton, 1924: 85, pl. 1 (figs 1-2); 1925: 267, pl. 12 (figs 1-2).
Phylloceras seresitense Pervinquiére, Spath, 1923: 18, pl. 1 (fig. 2), pl. 2 (fig. 1). Collignon
(in Besairie) 1936: 190, pl. 21 (figs 1-2). Fabre, 1940: 211, pl. 5 (fig. 1).
Phylloceras boulei Collignon, 1928: 144, pl. 15 (fig. 5), fig. 1.
Hyporbulites seresitensis (Pervinquiére) Breistroffer, 1947: 82.
Phylloceras (Hyporbulites) seresitense Pervinquiére, Collignon 1950: 66; 1963: 4, pl. 241
(fig. 1038), pl. 242 (fig. 1041).
178 ANNALS OF THE SOUTH AFRICAN MUSEUM
Hyporbulites seresitensis var. raynaudiensis Collignon, 1956: 16, pl. 4 (fig. 1).
Neophylloceras seresitense (Pervinquiére) Matsumoto, 1959: 55, pl. 12 (figs 4-5), fig. 3.
? Phylloceras (Euphylloceras) vohipalense Collignon, 1962: 1, pl. 215 (fig. 940).
? Phylloceras sp. (Ph. velledae ?) da Silva, 1962: 26, pl. 15 (figs 1-3).
Hypophylloceras seresitense seresitense (Pervinquiére) Wiedmann, 1962a: 142, pl. 8 (figs 1-2)
fig. 8; 19625: 249, pl. 16 (fig. 1).
Hypophylloceras seresitense tanit (Pervinquiére) Wiedmann, 1962a: 142; 1962b: 250, fig. 2.
Phylloceras (Hypophylloceras) seresitense seresitense Pervinquiére, Wiedmann, 1964: 221,
pl. 15 (fig. 4), pl. 21 (fig. 1), fig. 52. Wiedmann & Dieni, 1968: 26. Kennedy & Klinger,
1977a: 364, pl. 4 (fig. 6), pl. 6 (fig. 4), pl. 7 (fig. 4), pl. 9. Renz, 1968: 17, pl. 1 (fig. 1).
Phylloceras (Hypophylloceras) seresitense tanit Pervinqui¢re, Wiedmann, 1964: 226, pl. 21
(figs 2-3), fig. 54. Wiedmann & Dieni, 1968: 26, pl. 1 (fig. 6), pl. 3 (figs 1-2).
Phylloceras (Hypophylloceras) seresitense boulei Collignon, Wiedmann, 1964: 224, pl. 20
(figs 2-3), fig. 53. Renz, 1968: 18, pl. 1 (fig. 2), figs 6a, 7c. Forster, 1975: 140, pl. 1 (fig. 1),
fig. 27.
? Phylloceras (Hypophylloceras) seresitense vohipalense Collignon, Forster, 1975: 139, fig. 26.
? Phylloceras serum var. perlobata Zwierzycki (non Sayn), 1913: 323, figs a-c.
? Phylloceras ex aff. ramosi Meek, Collignon, 1928: 1, pl. 1 (figs 2-4).
? Phylloceras aff. tanit Pervinquiére, Matsumoto, 1942: 674, fig. 2.
Phylloceras cf. seresitense Pervinquiére, Wright & Wright, 1951: 12.
? Phylloceras cfr. semistriatum Choffat (non d’Orbigny), 1903: 17, pl. 1 (fig. 10).
Material
Ten specimens, SAM-6527, USNMNH 236897-236904, 237013, all with
recrystallized shell preserved, and all from Porto Amboim. ~
Description
The coiling is very involute, with a narrow, crater-like umbilicus
(6-7% of the diameter). The whorl section is strongly compressed, elliptical
(W/~ = 0,55-0,67). The flanks are slightly convex, with maximum width just
below midflank, converging to a narrow, evenly rounded venter. Ornament
comprises fine, flexuous lirae, very faint or completely effaced on the inner half
of the flanks (faint growth striae suggest they were prorsiradiate there),
recurving at about midflank so as to pass straight or slightly backwards across
the venter. On USNMNH 236897 there are about 128 lirae per half-whorl.
None of the present specimens shows the sutures.
Measurements
No. D H W W/H U
USNMNH 236897 37 21(57) 13(35) 0,62 2,6(7)
» 26,5 15,5(58) +8,5(32) 0,55
USNMNH 236898 a2 30,5(59) 17,5(34) 0,57 3(6)
» 35 19(54) 11(31) 0,58 it
USNMNH 236900 15 8(53) 5(33) 0,63 1,1(7)
» 11 5,5(50) 3,7(34) 0,67 ?
USNMNH 236901 | OF, 10(59) 6,5(38) 0,65 1,0(6)
USNMNH 236902 || 12(57) TMD 0,64 ?
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 179
Fig. 1. Phylloceras (Hypophylloceras) seresitense Pervinquitre. A-B. USNMNH 236899.
C-E. USNMNH 236904. F-H. USNMNH 236897. I-K. USNMNH 236903. L-N. USNMNH
236898. O-P. USNMNH 236902. Q-S. USNMNH 236901. T-V. USNMNH 236900. x1.
180 ANNALS OF THE SOUTH AFRICAN MUSEUM
Intraspecific variation
Wiedmann (1964: 221) recognized three subspecies within P. (H.) seresitense:
P. (H.) seresitense seresitense—whorl section moderately compressed,
whorl width to height ratio up to 0,65. Umbilicus very narrow.
P. (A.) seresitense boulei Collignon—whorl section very compressed
(W/H = 0,50-0,57), with very narrow umbilicus.
P. (H.) seresitense tanit—whorl section compressed, but with moderately
open umbilicus.
The most noteworthy variation in the population before the writers is in
the degree of inflation of the whorls. As can be seen from Figure 2 the variation
in this respect (¥/H = 0,55-0,67) includes individuals of all of Wiedmann’s
(1964) subspecies and, since they are all broadly contemporaneous (ranging
from Aptian to Cenomanian), appear to have little biological significance. The
authors find, therefore, the application of the subspecies boulei and tanit
inappropriate to the present collection.
Discussion
The holotype of Phylloceras angolaense Haughton is in the South African
Museum, SAM-6527. The specimen is entirely septate and is replaced by
crystalline calcite.
The shell is strongly compressed and very involute, with a very narrow,
deep umbilicus. The flanks are slightly convex, almost flat, and converge
towards the narrow, evenly rounded venter (Fig. 31). Maximum width is about
one-quarter of the way up the flanks. Ribbing is very faint on the inner half of
the flanks, although growth striae suggest it was prorsiradiate here. On the
outer half of the flanks, the ribs recurve so as to become almost radial. The
ribbing is very fine, with fourteen ribs in a distance of 10 mm along the venter
on the adoral quarter of the outer whorl.
There are no features by which P. angolaense may be satisfactorily dis-
tinguished from P. seresitense and consequently the authors follow Wiedmann
(1962a, 19625) in regarding it as a junior subjective synonym of Pervinquiére’s
species.
Phylloceras velledae (Michelin) (Wiedmann 1964: 209, pl. 11 (fig. 1),
pl. 13 (fig. 4), pl. 21 (fig. 4), fig. 49) typically differs from the present species in
having more convex flanks, with much coarser, more flexuous lirae, and a
suture which shows diphyllic saddles in immaturity rather than tetraphyllic
saddles as in P. (H.) seresitense.
Hypophylloceras yeharai Nakai & Matsumoto (1968: 4, pl. 1 (figs 1-3),
pl. 3 (fig. 1)) is based upon crushed material said to differ from P. (H.) velledae
in its more compressed whorls (? enhanced by crushing) and less numerous and
broader lirae. Kennedy & Klinger (1977a) have recently demonstrated the wide
range of variation within contemporaneous populations of P. (H.) velledae and
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 181
WIDTH
YAEIGHT oO
0 30 60 90 120
SAE) —————
Fig. 2. Plot of inflation (whorl width/height ratio) versus diameter for Phylloceras seresitense
Pervinquiére, showing the unsatisfactory separation of subspecies. Circles = present Angolan
material; dots = P. seresitense tanit Pervinquiére; square = holotype of P. angolaense
Haughton; open triangles = P. seresitense seresitense Pervinquiére; black triangles =
P. seresitense boulei Collignon. (Data after Stoliczka 1865, Spath 1923, Wiedmann 1962,
Collignon 1963, Wiedmann & Dieni 1968, Renz 1968, Forster 1975, and Kennedy & Klinger
1977.)
figured examples (1977a: pls 10-11) as coarsely ribbed as the Japanese material,
Suggesting that P. (H.) yeharai may be within the limits of variation of
P. (A..) velledae.
Occurrence
Phylloceras (Hypophylloceras) seresitense ranges from Upper Aptian to
Middle Cenomanian, and is known from southern France, Switzerland,
southern England, Spain, Tunisia, Algeria, Sardinia, the Balearics, southern
India, Japan, Alaska, California, Mexico, Angola, Zululand, possibly Tanzania,
and Madagascar.
Order LYTOCERATIDA Hyatt, 1889
Superfamily LYTOCERATACEAE Neumayr, 1875
Family Tetragonitidae Hyatt, 1900
Genus Tetragonites Kossmat, 1895
Types species Ammonites timotheanus Pictet, 1848
Discussion
Wiedmann (1973) has provided the most recent discussion of the Albian to
Cenomanian members of this genus, recognizing three species groups. The
group of 7. rectangularis possesses persistent constrictions and a straight
umbilical suture, and includes 7. rectangularis Wiedmann, T. subtimotheanus
182 ANNALS OF THE SOUTH AFRICAN MUSEUM
Wiedmann, 7. kitchini (Krenkel), T. subbeticus Wiedmann, T. marrei Thomel
and 7. blaisoni Collignon. In the group of 7. timotheanus (Pictet), constrictions
are present only at an early growth stage, whilst the umbilical suture is retracted.
To this group Wiedmann (1973) assigned 7. timotheanus (Pictet), T. balmensis
Breistroffer, and 7. nautiloides (Pictet). Forms lacking constrictions at all
growth stages are referred to the group of 7. jurinianus (Pictet).
Tetragonites (Tetragonites) collignoni Breistroffer, 1940
Ammonites timotheanus Stoliczka (non Mayor), 1865: 146, pl. 73 (figs 3-4, 6).
Tetragonites collignoni Breistroffer, 1940: 110. Murphy, 1967a: 66, pl. 5 (figs 2-5), fig. 36.
Forster, 1975: 147, pl. 1 (fig. 5) (with synonymy).
Tetragonites subtimotheanus Wiedmann, 1962a: 131; 1973: 592, pl. 1 (fig. 5), pl. 2 (fig. 2),
pl. 3 (figs 1-5), pl. 7 (fig. 8), fig. 2 (with synonymy).
Tetragonites blaisoni Collignon, 1964: 31, pl. 324 (fig. 1448). Wiedmann, 1973: 601, pl. 1
(fig. 4), pl. 6 (figs 5-7), fig. 7 (with synonymy).
Material
One specimen, USNMNH 236916, preserved as an internal mould, from
Porto Amboim.
Description
Shell involute, somewhat inflated, with a depressed, trapezoidal whorl
section (W/H = 1,21). Umbilicus narrow (22% of the diameter), deep, with
steep, almost vertical umbilical walls. Umbilical shoulder evenly rounded, with
flattish flanks converging towards a broad, slightly convex venter which rounds
somewhat in maturity. There are two distinct constrictions in a distance equal
to the whorl height on the last portion of the outer whorl (representing part of
the body chamber), although the number of constrictions per whorl is unknown.
The constrictions are markedly prorsiradiate across the inner flanks, recurving
strongly in the region of the ventrolateral angulations to cross the venter with a
prominent concave sinus.
Measurements
No. D H W W/H U
USNMNH 236916 44 21,5(49) +26(59) 1,21 ++9,5(22)
Discussion
Problems concerning the specific identification of constricted tetragonitids
become very clear from the recent literature concerning this group (Wiedmann
1962a, 1962b, 1973; Collignon 1963, 1964; Murphy 1967a, 1967b; Wiedmann &
Dieni 1968; McLearn 1972; Forster 1975; Kennedy & Klinger 19776). Forster
(1975) has suggested that 7. subtimotheanus and T. blaisoni are junior subjective
synonyms of T. collignoni. Wiedmann (1973) rejected the latter name because it
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 183
was based upon a pyritic nucleus only 14 mm in diameter (although he retains
T. kitchini and T. subbeticus as valid species even though the types are only
18 and 16 mm in diameter respectively), considering it a nomen dubium. Because
the authors are at present unhappy with Wiedmann’s (1973) fine delimitation of
T. subtimotheanus Wiedmann, T. rectangularis Wiedmann, T. blaisoni Collignon,
and 7. kitchini (Krenkel), all of which are broadly contemporaneous, they have
some sympathy with Forster’s (1975) view in assigning the present material to
T. collignoni.
Pot)
Fig. 3. A-D. Mortoniceras (Angolaites) simplex (Choffat). A. SAM-PCA4613. B. SAM-
PCA4774. C. SAM-PCA4590. D. SAM-PCA4609. E-F. Drakeoceras cf. dellense Young.
E. SAM-PCA4662. F. SAM-—PCA4800. G. Mortoniceras (Durnovarites) perinflatum (Spath),
SAM-PCA4587. H. Mariella (Mariella) cf. oehlerti (Pervinquiére), SAM-—PCA4798.
I. Phylloceras (Hypophylloceras) seresitense (Pervinquiére), the holotype of P. angolaense
Haughton, SAM-6527. x1.
184 ANNALS OF THE SOUTH AFRICAN MUSEUM
Occurrence
Tetragonites collignoni ranges from the Lower Albian to the Middle
Cenomanian, and is known from Madagascar, southern India, Zululand,
Alaska, British Columbia, and Oregon.
Tetragonites (Tetragonites) kitchini (Krenkel, 1910)
Fig. 4
? Ammonites timotheanus Whiteaves (non Mayor), 1876: 41, pl. 3 (fig. 2).
Desmoceras (Puzosia; Latidorsella ?) kitchini Krenkel, 1910: 226, pl. 22 (fig. 8).
Tetragonites kitchini (Krenkel), Wiedmann 1962a: 171. Murphy, 1967a: 33, pl. 2 (figs 11-14),
figs 15-16. Wiedmann, 1973: 599, pl. 1 (fig. 7), pl. 6 (figs 1, ?2-4), figs 5, 26.
Tetragonites hulenensis Murphy, 1967a: 54, pl. 6 (figs 16-19), pl. 7 (figs 3, 6-8, 10), figs 28-30.
Murphy, 19675: pl. 4 (figs 8-9).
? Tetragonites aff. kitchini (Krenkel), McLearn 1972: 26, pl. 4 (figs 4-5).
Material
A single specimen, SAM-PCA3125, from low in the coastal cliffs
immediately north of the estuary at Praia-Egito (low dispar Zone).
Description
The shell is moderately involute (umbilicus 31,5°% of the diameter) and
partially retains recrystallized shell. The whorl section is subtrapezoidal, almost
subquadrate, as wide as high (W/n = 1,00), with slightly converging flanks and
a flattish, slightly convex venter. The umbilicus is rather narrow and deep, with
steep umbilical walls and subrounded umbilical shoulders. The ventrolateral
shoulders are evenly rounded. Maximum width is at the umbilical shoulder.
Fig. 4. Tetragonites kitchini (Krenkel). Lateral and ventral views of
SAM-PCA3125, partially retaining recrystallized test. x1.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 185
There would appear to have been about seven slightly flexuous (prorsi-
radiate concave in the terminology of Murphy (1967a)), strongly prorsiradiate
constrictions on the outer whorl, which cross the venter with a concave-adoral
sinus. The suture line was not observed.
Measurements
No. D H W W/H U
SAM-PCA3125 54 25(46) 25(46) 1,00 17(31,5 %)
Discussion
Tetragonites kitchini differs from all other contemporaneous species of
constricted Tetragonites in having less strongly inflated whorls with an almost
quadrate whorl section. However, there appear to be intermediate forms to
T. subtimotheanus Wiedmann and T. rectangularis Wiedmann, and the population
structures of these species require investigation to fully validate their specific
separation.
Occurrence
Tetragonites kitchini is known from the upper Lower Albian of Tanzania,
California, Alaska, and perhaps British Columbia, and from the Middle or
Upper Albian of Madagascar, and the uppermost Albian of Angola.
Tetragonites (Tetragonites) jurinianus (Pictet, 1847)
Figs 5-6
Ammonites jurinianus Pictet (in Pictet & Roux), 1847: 297, pl. 3 (fig. 3). D’Orbigny, 1850: 124.
Ammonites timotheanus Pictet & Campiche (zon Mayor), 1860: 289 (pars).
Lytoceras (Tetragonites) epigonum Boule, Lemoine & Thévenin (non Kossmat), 1906: 186,
pl. 3 (fig. 1).
? Lytoceras (Tetragonites) timotheanum Pervinquiére (non Mayor), 1907: 74 (pars.), pl. 3
(fig. 24 only).
Lytoceras (Tetragonites) jurinianum (Pictet) Jacob, 1908: 19, pl. 1 (fig. 12).
Latidorsella latidorsata Jacob (non Michelin), 1908: 35, pl. 5 (fig. 1 only).
Lytoceras (Tetragonites) zacatecanus Bose, 1923: 127, pl. 9 (figs 11-17).
Tetragonites jurinianus (Pictet) Spath, 1923: 26. Roman, 1938: 43. Breistroffer, 1940: 112.
Wiedmann, 1962a: 176, pl. 14 (fig. 2), figs 37-38. Almela & Revilla, 1957: 20, pl. 4 (fig. 3).
Murphy, 1967a: 23, pl. 2 (figs 1-4); figs 10-11. Renz, 1968: 19, pl. 1 (figs 6-7), figs 6d,
7Ja—b. Wiedmann & Dieni, 1968: 48, pl. 4 (fig. 4), pl. 5 (fig. 4). Wiedmann, 1973: 608,
pl. 8 (figs 3-4).
? Tetragonites brazoensis Bose, 1927: 203, pl. 1 (figs 2-7).
Tetragonites jurinianus var. angolana Haas, 1942: 170, pl. 44 (fig. 3), pl. 45 (fig. 1).
Tetragonites timotheanus Matsumoto (non Mayor), 1959: 78, fig. 16.
Tetragonites jurinianus angolanus Haas, Wiedmann, 1973: 609.
Material
Ten specimens, USNMNH 236915, 236917-21, 236977-79, all with
recrystallized shell preserved, and all from Porto Amboim.
186 ANNALS OF THE SOUTH AFRICAN MUSEUM
Description
Shell inflated, involute, with a slightly depressed, trapezoidal whorl section
in juveniles (“/q = 1,00-1,29), the venter becoming rounded in maturity.
Umbilicus narrow (16-30% of the diameter), deep, with almost vertical walls
and evenly rounded umbilical shoulders. The flanks are flattened, with maximum
width close to the umbilical shoulder, and converge towards the venter. Up to
25 mm diameter, the venter is flattened, very slightly convex, whereafter it
becomes rounded and the ventrolateral angulations disappear. At this stage the
whorl section is broadly ovate. Constrictions are lacking at all observed growth
stages.
Measurements
No. D H WwW W/H U
USNMNH 236915 32:5 25,5(49) +28(53) 1,10 10(19)
USNMNH 236917 38,5 20(52) +22(57) 1,10 +9(23)
Pe 30 14,5(48) 16(53) 1,10 9(30)
USNMNH 236918 43 19(44) 22,3(52) 1,17 9,8(23)
USNMNH 236919 16 7,5(47) 8,5(53) 1,13 4(25)
USNMNH 236920 18 8(44) 10(55) 125 +4(22)
USNMNH 236921 21 10(48) +10(48) 1,00 5(24)
USNMNH 236977 17,5 6,8(39) 8,5(49) 1,25 4,5(26)
USNMNH 236978 iW 7(41) 9(53) 1,29 4,7(28)
T. jurinianus
angolanus
(holotype) 105 56(53) 51(48) 0,91 17(16)
ES 63 32(51) 34(54) 1,06 ?
Intraspecific variation
The intraspecific variation seen in Tetragonites jurinianus shows features
which have an important bearing on the classification of tetragonitids in
general. The wide range of inflation within juveniles far exceeds the range of
variation admitted by Wiedmann (1973) within the constricted tetragonitids as
a whole, and population studies may show his subdivisions of the group to be
utilitarian rather than biologically significant.
As can be seen from Figure 6, there is not only a distinct decrease in the
umbilical ratio with growth, but there is also an ontogenetic increase in the
height of the whorls, which become progressively less inflated.
Wiedmann (1973) maintained 7. jurinianus angolanus as a separate sub-
species by virtue of its narrower umbilicus (16% versus 23% of the diameter in
T. jurinianus jurinianus), high-oval whorl section and large size. It appears,
however, that these differences result from a comparison of different onto-
genetic stages, and the authors include 7. jurinianus angolanus in the synonymy
of T. jurinianus sensu stricto.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 187
Fig. 5. Tetragonites (Tetragonites) jurinianus (Pictet). A-B. USNMNH 236918. C-D, G.
USNMNH 236921. E-F. USNMNH 236978. H-J. USNMNH 236977. K-M. USNMNH
236917. N-P. USNMNH 236919. Q-S. USNMNH 236920. x1.
188 ANNALS OF THE SOUTH AFRICAN MUSEUM
Discussion
As diagnosed by Wiedmann (1973: 606), Tetragonites nautiloides (Pictet)
does not have constrictions beyond 10 mm diameter and thus closely approaches
T. jurinianus. Indeed, the present material all exceeds this diameter and could,
therefore, equally well be assigned to 7. nautiloides. However, Murphy (1967a)
has noted that constrictions are present to 27 mm diameter in topotype material
of T. nautiloides when there is considerable difficulty in distinguishing this
1,50
Wi
0,70
0 30 60 90
DIAMETER (mm) ————>
UMBILICUS(%)
0 40 70 110
DIAMETER (mm) —————
Fig. 6. Plot of inflation (whorl width/height ratio) and umbilical ratio against diameter for
examples of Tetragonites jurinianus (Pictet). Dots = present Angolan material; squares =
the holotype of 7. jurinianus angolanus Haas; circles = material described by Wiedmann
(1962, 1973), Murphy (1967a), Wiedmann & Dieni (1968).
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 189
species from some constricted tetragonitids, e.g. 7. rectangularis Wiedmann.
Wiedmann’s (1973) diagnosis of T. nautiloides is difficult to support, since some
of his figured specimens (pl. 8 (figs 2, 5)) clearly show weak constrictions at
22 and 27 mm diameter respectively, suggesting that this species may better be
included in the T. rectangularis group.
Occurrence
Tetragonites jurinianus ranges from Middle Albian to Lower Cenomanian,
and is recorded from Switzerland, France, Sardinia, Mallorca, Madagascar,
Angola, Mexico, and California.
Family Gaudryceratidae Spath, 1927
Subfamily Gaudryceratinae Spath, 1927
Genus Eogaudryceras Spath, 1927
Type species Ammonites numidus Coquand, 1880
Discussion
Eotetragonites was originally separated (Breistroffer 1947) from Eogaudry-
ceras by the possession of strong constrictions throughout ontogeny, and a
suture with irregularly bifid saddles. However, Wiedmann (19625: 35) has noted
the occurrence of species which show combinations of these characters, conse-
quently treating Eotetragonites as a subgenus of Eogaudryceras. As defined by
him (19625), E. (Eogaudryceras) has initially trapezoidal whorls which become
rounded in maturity and are ornamented by fine, flexuous lirae, whereas
E. (Eotetragonites) has quadrate juvenile whorls and lacks liration.
Eogaudryceras (Eogaudryceras) italicum Wiedmann & Dieni, 1968
Figs 7-11
Eogaudryceras (Eogaudryceras) italicum Wiedmann & Dieni, 1968: 34, pl. 1 (fig. 8), fig. 6.
Material
Ten specimens, USNMNH 236905-236914, all with recrystallized shell
preserved, and all from Porto Amboim.
Description
Up to 23 mm diameter: shell evolute, compressed, with a wide, shallow
umbilicus (30-41 °% of the diameter) and steep umbilical walls which merge
imperceptibly with the strongly convex flanks. The whorl section is oval,
depressed (W/H = 1,00-1,36), with a broadly rounded venter. The outer whorl
conceals about 55 per cent of the preceding whorl. The earliest whorls are very
finely lirate, almost smooth to the naked eye, with sporadic, rather distinct,
prorsiradiate collars.
190 ANNALS OF THE SOUTH AFRICAN MUSEUM
24-45 mm diameter: the shell form is much as at the earlier growth stages,
except that the flanks flatten slightly and the whorl section changes from slightly
depressed to almost quadrate. At this stage the lirae are visible to the naked eye;
they arise at the umbilical seam and pass forwards (prorsiradiate) on the lower
third of the flanks. Thereafter they recurve slightly, only to flex forwards again
before crossing the venter. At intervals one or two adjacent lirae are strengthened,
presumably corresponding to the collars of the earliest whorls. The lirae become
flat-topped, band-like, much broader than the interspaces, and of variable
thickness at this stage.
Greater than 46 mm diameter: in maturity the whorls become flat-sided and
compressed (W/H = 0,66—-0,80), with an elliptical whorl section and narrowly
rounded venter. The umbilicus is moderately wide (26-33% of the diameter),
shallow, with steep umbilical walls and evenly rounded umbilical shoulders.
Maximum width is at about mid-flank. At this growth stage the irregular band-
like ribs are split by fine, threadlike grooves. In the largest specimen (still septate
at 82 mm diameter) there are still occasional strengthened ribs, followed by a
slightly deeper intercostal groove. These may correspond to the collars of the
earliest whorls.
Fig. 7. Eogaudryceras (Eogaudryceras) italicum Wiedmann & Dieni. Reconstruction of
juvenile and adult shells. x1.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 191
C
Fig. 8. Eogaudryceras (Eogaudryceras) italicum Wiedmann & Dieni. A~-B. USNMNH 236905.
C-E. USNM 236908. x1.
192 ANNALS OF THE SOUTH AFRICAN MUSEUM
Measurements
No. D H W Wily U
USNMNH 236905 81 37,5(46) 25(31) 0,66 21(26)
5s 56 25(45) 19,5(35) 0,78 17,561)
USNMNH 236906 34 13(38) 15,5(45) 1,19 +10(29)
USNMNH 236907 +47 +21(45) +18(38) 0,86 IS,363%
USNMNH 236908 v1 33,5(47) +27(38) 0,80 19(27)
USNMNH 236909 26 11(42) 11(42) 1,00 8(31)
fe 19,5 8(41) 8(41) 1,00 +5,5(28)
USNMNH 236910 24 9(38) 8(38) 1,00 8,5(35)
. 18 6,5(36) 7(39) 1,08 7(39)
USNMNH 236911 15 5,5(37) 6(40) 1,09 5(30)
USNMNH 236912 22 8(36) 9(41) 1,13 8(36)
USNMNH 236913 17 5,5(32) +7,5(44) 1,36 7(41)
USNMNH 236914 25 9(36) +11(44) 122 +8,5(34)
E. (E. italicum* +50 25(50) 19(38) 0,76 +14(28)
E. (E.)aenigmum* 58,3 28,3(48,5) 24(41,5) 0,86 14(24)
E. (E.)b. bourritianum* 33 14(42) 20(60) 1,43 10(30)
E. (E.)b. hispanicum* 47 20(42) 22(47) 1,10 13(28)
Gaudryceras aff.
madraspatanum
Spath (non Stoliczka) 28 11,8(42) 11,8(42) 1,00 10(36)
* Asterisks mark holotypes or neotypes.
Intraspecific variation
The most noteworthy feature of the present material is the great change in
whorl section with growth. The earliest whorls show a wide range in inflation
(W/a = 1,00-1,36) but, with continued growth, the shell becomes increasingly
high-whorled and the whorl section becomes strongly compressed (Fig. 11).
At the same time there is a slight, but distinct, tendency for the umbilical ratio
to decrease (Fig. 11).
Discussion
E. (Eogaudryceras) italicum Wiedmann & Dieni is based upon a smooth,
fragmentary internal mould from the Upper Albian of Sardinia. It agrees
precisely with the present material in whorl section and relative proportions;
lack of liration may simply be a reflection of different preservation.
Gaudryceras aff. madraspatanum (Stoliczka) (Spath 1923: 22, pl. 1 (fig. 4))
is based upon a smooth phosphatic internal mould from the Cambridge
Greensand of southern England. In relative proportions it falls well within the
range of variation of the present Angolan material, and the writers would
tentatively refer it, therefore, to E. italicum.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 193
K
Fig. 9. Eogaudryceras (Eogaudryceras) italicum Wiedmann & Dieni. A-C. USNMNH 236905.
D-F. USNMNH 236906. G, K-L. USNMNH 236910. H-J. USNMNH 236909.
M-O. USNMNH 236912. x1.
194 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 10. Eogaudryceras (Eogaudryceras) italicum Wiedmann & Dieni.
A-B. USNMNH 236907. C-E. 236911. A-D x1, E x2.
The present sample is the largest single population of an E. (Eogaudryceras)
species yet described, and shows a wide range of ontogenetic and intraspecific
variation. In view of this wide range of variation, it seems probable that
population studies will reduce in number the twelve species currently assigned
to this subgenus, viz. E. (E.) numidum (Coquand), E. (E.) intermedium (Fallot),
E. (E.) vocontianum (Fallot), E. (E.) elegans Basse, E. (E.) shimizui Breistroffer,
E. (E.) llosetae Breistroffer, E. (E.) turgidum Breistroffer, E. (E.) skoenbergense
Collignon, E. (£.) bourritianum (Pictet), E. (E£.) aenigmum (Haas) and
E. (£.) muntaneri Wiedmann.
Of other Upper Albian species of Eogaudryceras (Eogaudryceras),
Wiedmann (1962a: 154) selected a neotype for E. (£.) bourritianum (Pictet)
(in Pictet & Roux 1848: 298, pl. 4 (fig. 1)) and divided it into two chronological
subspecies: E. (Eogaudryceras) bourritianum bourritianum, from the uppermost
Albian (dispar zone) of south-west France is characterized by its strongly
depressed whorl section (W/H = 1,43), involute form (U = 30% of the diameter)
and smooth whorls (the neotype is preserved as an internal mould and hence the
lack of ornament may not be a diagnostic character). It differs from the Angolan
material in its much more depressed whorl section and narrower umbilicus;
E. (Eogaudryceras) bourritianum hispanicum Wiedmann (1962a: 155, pl. 12
(fig. 6), fig. 15) is a low Upper Albian form (associated with Hysteroceras and
Mortoniceras) which was said to differ from the typical form in its less depressed,
trapezoidal whorl section (W/H = 1,10), flattened venter, and greater involution
(U = 28% of the diameter). It differs from E. (E.) italicum in being more
involute, having a slightly more depressed whorl section and a flattened venter.
E. (Eogaudryceras) aenigmum (Haas) (1942: 167, pl. 42 (fig. 3), pl. 44 (fig. 2),
fig. 24) is from the Upper Albian of Angola. Crushed material assigned to this
species, in the British Museum (Natural History), comes from Praia do Jombe
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 195
1,40
we eG —$_—_—____—_
0,50
0 30 60 90
DIAMETER (mm) =>
UMBILICUS ( %) —————»>
0 100
DIAMETER (mm) ———
Fig. 11. Plot of inflation and umbilical ratio against diameter for Eogaudryceras italicum
Wiedmann & Dieni. Dots = present Angolan material; square = holotype (after Wiedmann
& Dieni 1968).
196 ANNALS OF THE SOUTH AFRICAN MUSEUM
where it is associated with a low Upper Albian fauna which includes Prohystero-
ceras wordiei Spath, Beudanticeras beudanti (Brongniart) and Mortoniceras cf.
inflatum (J. Sowerby). It is thus a contemporary of E. (E.) bourritianum
hispanicum. The style of ornament is difficult to judge from Haas’s (1942)
illustration, although the specimen is clearly lirate (it has shell preserved).
Although its whorl width/height ratio is closely comparable to E. italicum
(? due to crushing), it is much more involute (U = 24% of the diameter).
Haas’s (1942) whorl section of the species shows inner whorls with a strongly
fastigiate to keeled venter which suggest his material to be crushed. If true,
this species may prove to have priority over E. (E.) bourritianum hispanicum.
Occurrence
E. (Eogaudryceras) italicum is currently known from the uppermost Albian
of Sardinia, Angola, and possibly southern England.
Suborder ANCYLOCERATINA Wiedmann, 1966
Superfamily ANCYLOCERATACEAE Meek, 1896
Family Ancyloceratidae Meek, 1876
Subfamily Anisoceratinae Hyatt, 1900
Genus Anisoceras Pictet, 1854
Type species Hamites saussureanus Pictet, 1847
Anisoceras (Anisoceras) perarmatum Pictet & Campiche, 1861
Figs 12A—H, 13C—D, 14A-C, 15C-—F, 16B
Hamites armatus J. de C. Sowerby (non J. Sowerby), 1850: pl. 29 (fig. 13).
Anisoceras perarmatum Pictet & Campiche, 1861: 65, pl. 48 (figs 7-8), pl. 49 (figs 1-3, 5-7).
Pictet, 1861: 21. Ooster, 1863: 19. Pictet & Renevier, 1866: 103. Jukes-Browne, 1875:
288. Barrois, 1878: 271. Renevier, 1890: 340. Boule, Lemoine & Thévenin, 1907: 35.
Ganz, 1912: 121. Spath, 1921: 289. Bése, 1923: 144. Diener, 1925: 73. Spath, 19255: 191.
Barbu, 1932: 16. Roman, 1938: 52. Spath, 1939: 548, pl. 59 (figs 1-3), pl. 61 (figs 3-7),
fig. 192. Breistroffer, 1947: 62. Reyment, 1955: 12, pl. 1 (fig. 1). Collignon, 1963: 50,
pl. 259 (figs 1126-1127). Swensen, 1963: 67, pl. 4 (figs 1, 3, 7). Dieni & Massari, 1963: 798.
Clark, 1965: 25, pl. 6 (figs 1, 3, 7). Wiedmann & Dieni, 1968: 65, pl. 6 (fig. 4), pl. 7 (fig. 9),
figs 38-39. Renz, 1968: 74, pl. 13 (fig. Sa—c), pl. 14 (figs 1, 3, 5), figs 27a, 28g.
Hamites (Anisoceras) perarmatus (Pictet & Campiche) von Hauer, 1861: 644, pl. 2 (figs 2-4).
Pervinquiére, 1907: 85.
Hamites (Anisoceras) saussureanus von Hauer (non Pictet), 1861: 644, pl. 2 (fig. 1).
Anisoceras armatus Stoliczka (non J. Sowerby), 1866: 174, pl. 81 (figs 8-10), pl. 82 (fig. 1).
Hamites perarmatus (Pictet & Campiche) Neumayr, 1875a: 30; 18755: 898.
? Anisoceras vraconense Renz, 1968: 75, pl. 16 (fig. 2), fig. 27b.
Anisoceras perarmatum simplex Renz, 1968: 75, pl. 13 (fig. 7), pl. 14 (fig. 4), fig. 27k.
? Anisoceras pseudopunctatum Pictet & Campiche, 1861: 74, pl. 52 (figs 1-3). Breistroffer,
1947: 62. Renz, 1968: 79, pl. 16 (figs 1, 3, 5). Wiedmann & Dieni, 1968: 72.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 197
Fig. 12. A-H. Anisoceras perarmatum Pictet & Campiche. A-B. USNMNH 236928.
C-D. USNMNH 236929. E-F. USNMNH 236944. G-H. USNMNH 236940, I-J. Anisoceras
sp. juv., USNM 236745. x1.
198 ANNALS OF THE SOUTH AFRICAN MUSEUM
A B C D
Fig. 13. A-B. Anisoceras armatum (J. Sowerby), USNMNH 236934. C-D. Anisoceras
perarmatum Pictet & Campiche, USNMNH 236931. x1.
Material
Twenty-seven specimens, SAM-PCA2940, 2950, 2956, 3115, 3122, 3143,
3148, 3152, 3193, 3214, 3243, 3283, 3285, 3299, 3339, 3352, 3355, 3358, 3361 and
3391, all from Egito, SAM-—PCA4801, 4804 from Cabo Ledo and USNMNH
236928-29, 236931, 236933, 236940, and 236944 from Porto Amboim, either
retaining recrystallized shell or preserved as composite internal moulds.
Description
Whorl section varies from almost circular in specimens replaced by calcite
to slightly elliptical in those crushed individuals preserved as composite internal
moulds.
Ornament comprises small dorsolateral to lateral tubercles and the septate
bases of prominent ventrolateral spines, between which strong ribs are looped
in pairs. There are sporadic non-tuberculate intercalatories between looped ribs.
The main ribs are also looped across the venter, and split into fine riblets,
thirteen per three dorsolateral tubercles, on the dorsum. On the body chamber
there may be one to two simple ribs (SAM-PCA3143, 3148) between looped
ribs, whilst on the final shaft of USNMNH 236940, the button-and-loop
ribbing is lost, and all ribs become single.
Discussion
Amongst contemporaneous species of Anisoceras, only A. saussureanum and
A. armatum closely approach this species. Differences are noted under the
discussion of A. armatum.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 199
kat
Fig. 14. A-C. Anisoceras perarmatum Pictet & Campiche. Dorsal, lateral and ventral views of
SAM-PCA3115. D-E. Anisoceras cf. armatum (J. Sowerby). A fragment of a body chamber,
SAM-PCA3287, which may belong here. F. Anisoceras sp. Oblique lateral view of SAM-
PCA3220. x0,75S.
200 ANNALS OF THE SOUTH AFRICAN MUSEUM
Anisoceras vraconense Renz (1968: 75, pl. 16 (fig. 2), fig. 27b) differs from
A. perarmatum in having most ribs simple and much weaker dorsolateral
tubercles. This species may be based upon a fragment of the final shaft of
A. perarmatum, since the ribbing simplifies on the body chamber.
Anisoceras charlottense Anderson (1958: 209, pl. 11 (fig. 3)) has a circular
whorl section and very prominent looped ribs separated by three to four simple,
finer intercalatories. It may be conspecific with the low Upper Albian A. salei
Clark (1958: 1079, pl. 140 (fig. 3)).
Anisoceras perarmatum simplex Renz (1968: 75, pl. 13 (fig. 7), pl. 14 (fig. 4),
fig. 27k) is simply a variant based on a small body chamber fragment.
Anisoceras pseudopunctatum Pictet & Campiche (1861: 74, pl. 52 (figs 1-3))
is based on material which, like A. perarmatum simplex, has also lost button-and-
loop ribbing, and may equally be an intraspecific variant.
Occurrence
Anisoceras perarmatum is a typical dispar Zone species known from
England, France, Switzerland, Sardinia, Tunisia, Nigeria, Madagascar, southern
India, Texas, and Angola.
Anisoceras (Anisoceras) armatum (J. Sowerby, 1817)
Figs 13A—B, 14D-E, 16A, C, E, I, 17-19
Hamites armatus J. Sowerby, 1817: 153, pl. 168. De Haan, 1825: 152, no. 2. Buckland, 1837:
65, pl. 44 (figs 9-10). Brown, 1837: 2, pl. 2 (fig. 6). Romer, 1840: 94, pl. 15 (fig. 2).
D’Orbigny, 1842: 547, pl. 135.
Non Hamites armatus Mantell (non J. Sowerby), 1822: 121, pl. 23 (figs 3-4) (= A. plicatile).
Baculina armata (J. Sowerby) Fleming, 1828: 250.
? Hamites undulatus Brown (non Forbes), 1837: pl. 2 (fig. 11).
Non Hamites armatus J. de C. Sowerby (non J. Sowerby), 1850: pl. 29 (fig. 13)
(= A. perarmatum).
? Hamites armatus J. Sowerby, Dixon, 1851: pl. 29 (fig. 13).
Anisoceras armatum (J. Sowerby) Pictet & Campiche, 1861: 62, pl. 48 (figs 1-2, 4, 6). Spath,
1939: 543, pl. 59 (fig. 6), pl. 60 (fig. 1), pl. 61 (figs 9-11), pl. 62 ( ?fig. 5), fig. 191. Swensen,
1963: 66, pl. 3 (fig. 4), pl. 4 (?fig. 6). Clark, 1965: 25, pl. 5 (fig. 4), pl. 6 ( ?fig. 6), fig. 7a.
Renz, 1968: 75, pl. 15 (figs 1, 3), figs 27d, 28a.
? Hamites (Anisoceras) armatus J. Sowerby, von Hauer, 1861: 644, pl. 1 (figs 9-10).
Non Anisoceras armatum Stoliczka (non J. Sowerby), 1866: 174, pl. 81 (figs 8-10), pl. 82 (fig. 1)
(= A. perarmatum).
Hamites (Anisoceras) armatus J. Sowerby, Kossmat, 1895: 149.
? Anisoceras armatum (J. Sowerby) Choffat, 1905: 41, pl. 1 (fig. 6).
Non Hamites (Anisoceras ?) armatus Pervinquiére (non J. Sowerby), 1907: 84, pl. 4 (figs 2-3)
(= ? A. exoticum).
Anisoceras aff. armatum (J. Sowerby) Adkins, 1920: 69.
? Hamites cf. armatus J. Sowerby, Passendorfer, 1921: 237.
Anisoceras cf. armatum (J. Sowerby) Bose, 1923: 143, pl. 10 (figs 22-24). Haughton, 1924: 94.
Clark, 1958: 1080, pl. 139 (fig. 2).
Non Anisoceras aff. armatum (J. Sowerby) Spath, 19256: 190 (= A. raynaudi).
Anisoceras picteti Spath (non Matheron), 1926a: 432. Spath, 1939: 554, pl. 59 (fig. 4), pl. 61
(fig. 8), pl. 63 (figs 3, 8), fig. 194. Renz, 1968: 76, pl. 13 (figs 8-9), pl. 14 (figs 6-9), pl. 15
(fig. 4), figs 27c, 28f.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 201
Fig. 15. A-B. Anisoceras sp. Oblique dorsolateral and lateral views of SAM-PCA3301 from
Egito. C-F. Anisoceras perarmatum Pictet & Campiche. C-D. Ventral and lateral views of
SAM-PCA3154. E-F. Lateral and ventral views of SAM-—PCA4801. G-—H. Anisoceras sp.
An indeterminate body chamber fragment from Egito, SAM-PCA2942. I-J. Mortoniceras
(Angolaites) simplex (Choffat). Lateral and ventral views of SAM-—PCA3249. A-D, G-H x 0,75,
E-F, I-J x1.
202 UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL
? Hamites aff. armatus J. Sowerby, Scott, 1926: 80. Bose, 1928: 146. Adkins, 1928: 24.
? Anisoceras cf. armatum (J. Sowerby) Passendorfer, 1930: 667.
? Non Hamites (Anisoceras) armatus Collignon (non J. Sowerby), 1932: 20, pl. 4 (fig. 1).
Anisoceras Saussureanum var. spinosa Haas, 1942: 192, pl. 46 (figs 1-3), fig. 30.
Non Anisoceras armatum Haas (non J. Sowerby), 1942: 189, fig. 29 (= A. tropicale).
Anisoceras jacobi Breistroffer, 1946: 310; 1947: 62. Wiedmann & Dieni, 1968: 67, pl. 6 (fig. 13),
fig. 41.
? Anisoceras aff. picteti Spath, Kennedy, 1971: 13, pl. 3 (fig. 6), pl. 7 (fig. 10).
? Non Anisoceras armatum (J. Sowerby) Kennedy, 1971: 14, pl. 5 (fig. 11).
Material
Fourteen specimens, SAM-—PCA2932, 2947, 2953, 3174, 3364, and 3398
from Egito, USNMNH 236930, 236934, 236936-37, 236939, and 9236952 from
Porto Amboim, and SAM-—PCA4606 and 4610 from Cabo Ledo.
Description
Shell form suggests initial coiling in an open planispiral, straightening in
maturity. The whorl section is approximately circular.
The ornament is rather variable, comprising slightly rursiradiate to slightly
prorsiradiate looped ribs, tetween which are generally one, sometimes two,
simple intercalatories. Looped ribs are ornamented with small pointed dorso-
lateral tutercles and the septate bases of prominent ventrolateral spines.
All rits are of more or less equal strength. Across the dorsum the ribs divide into
fine riblets and are accompanied by intercalatories.
In USNMNH 236939, which is taken to represent an early portion of the
spire of the species, main ribs are weakly looped and are separated by two to
three intercalatories. In USNMNH 236952, the largest specimen (which is,
however, still septate), the whorl section is slightly compressed, oval, and there
is generally only one intercalatory between looped ribs, although even these are
sometimes absent.
Discussion
Anisoceras armatum is a widely-cited but poorly understood species, and in
consequence the holotype is refigured here (Fig. 17). As noted by Spath
(1939: 546), this is a crushed, composite internal mould which shows the
following features: the penultimate shaft has distinct looped ribs separated by
only one non-tuberculate intercalatory which is of approximately the same
strength as the looped ribs. At this stage there are about five ribs in a distance
equal to the whorl height (allowing for post mortem crushing). On the hook, the
preservation is poor, but the looped ribs clearly break up so as all to become
single on the final non-septate shaft. At this stage there are still only about
five ribs in a distance equal to the whorl height, and most of the ribs are of
approximately the same strength. The dorsolateral tubercles are still present in
maturity, although weak and irregularly developed.
It is clear from the above description, and comparison with Sowerby’s
original illustration (cf. Spath 1939, fig. 191), that this species has been mis-
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 203
Fig. 16. A, C, E, 1. Anisoceras armatum (J. Sowerby). A. SAM-—PCA5470. C. SAM-PCAS5471.
E. SAM-PCAS5473. I. SAM-PCAS5472. B. Anisoceras perarmatum Pictet & Campiche.
Lateral view of SAM-PCA3339, showing change of ornament on the body chamber.
D. Hamites duplicatus Pictet & Campiche. Lateral view of SAM-PCA2955. F. Hamites
virgulatus Brongniart. Lateral view of SAM-PCA3158, showing venefzianus-type ribbing.
G. Mariella (Mariella) gresslyi (Pictet & Campiche). SAM-PCA3133. H. Hamites virgulatus
Brongniart. Lateral view of SAM-PCA2959, showing rather dense subvirgulatus-type ribbing.
J-K. Anisoceras phillipsi sp. noy. Lateral and dorsal views of SAM-PCA 4799. B 0,75,
other x1.
204 ANNALS OF THE SOUTH AFRICAN MUSEUM
interpreted. Indeed, Sowerby’s figure bears no great resemblance to the original
and shows too many intercalated ribs both on the penultimate and final shafts,
while the differentiation of the ribs on the final shaft is less prominent than is
shown in Sowerby’s drawing.
Anisoceras jacobi Breistroffer (nom. nov. pro Anisoceras picteti Spath (non
Matheron) 1939: 554, pl. 59 (fig. 4), pl. 61 (fig. 8), pl. 63 (figs 3, 8), fig. 194) was
diagnosed as follows: ‘Like A. armatum, but more coarsely ornamented, with
Fig. 17. Anisoceras armatum (J. Sowerby). Sowerby’s (1817, pl. 178) original
figured specimen from the Upper Greensand of Roak, near Benson,
Oxfordshire. Oxford University Museum K675a. x 1.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 205
A B C
Fig. 18. A-C. Anisoceras armatum (J. Sowerby), USNMNH 236952. x1.
the intermediate ribs (one, rarely two) as prominent as the main ribs that meet
at the strong tupercles. Suture-line similar to that of A. armatum.’
These are the exact features which characterize the penultimate shaft of the
holotype of the contemporaneous A. armatum, and the authors regard A. jacobi
as a junior subjective synonym of A. armatum.
Anisoceras armatum and A. perarmatum are contemporaneous species
which differ in that the latter generally lacks intercalatories between the
206 ANNALS OF THE SOUTH AFRICAN MUSEUM
looped ribs in maturity, although there are intermediates between the two
species, e.g. USNMNH 236952 (Fig. 18).
Spath (1939) noted the very close resemblance between A. saussureanum
(Pictet) and A. armatum and considered immature growth stages indistinguish-
able. In maturity, A. saussureanum was said to differ by its distant tuberculation,
with commonly two to three intercalatories between looped ribs, whilst coming
from a lower level in the Albian. Spath (1939) appeared to place much weight,
in his separation of the above two species, on the supposedly lower horizon of
t
\ v
h
itis
ail
77)
/
£.
som
aka
F G H
Fig. 19. Anisoceras armatum (J. Sowerby). A-B. USNMNH 236930. C-D. USNMNH
236936. E-F. USNMNH 236937. G-H. Doubtful juvenile, USNMNH 236939. x1.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 207
A. saussureanum, although Breistroffer (1940, 1947) and Renz (1968) record it
from the uppermost Albian of France and Switzerland respectively. Moreover,
A. saussureanum spinosum (Haas) is recorded from the uppermost Albian of
Angola and Mexico (fide Haas 1942). The holotype of A. saussureanum spinosum
comes from the same locality as the Washburn collection and the authors have
no hesitation in regarding it as a junior subjective synonym of A. armatum.
However, Haas’s identification clearly focuses on the difficulties of separating
A. armatum and A. saussureanum and, in view of the fact that they are contem-
porary species in the uppermost Albian, they probably do not bear specific
separation. However, until the type and topotype material of A. saussureanum
are restudied with regard to their intraspecific variation, it seems preferable to
retain these two well-known species separate.
Anisoceras tropicale (Meunier) (1887: 62, pl. 1 (fig. 5)) is based upon a very
poorly preserved internal mould which was tentatively referred to A. armatum
by Choffat (1905: 41) and Haas (1942: 191), although Spath (1939: 558)
considered it closer to A. pseudoelegans. Meunier’s specimen shows three fine
intercalatories between looped ribs and, judging from the locality, a valley to
the north of Lobito, is somewhat older than A. armatum. This is supported by
the example of A. armatum figured by Haas (1942, fig. 29), which the authors
would assign to A. tropicale; it occurs on the reverse side of the holotype of
Mortoniceras vokesi (Haas), topotype material of which comes from the
mid-Upper Albian zone of Elobiceras elobiense (Cooper 1978) at Lobito, in
association with Puzosia cuvervillei (Meunier).
Anisoceras exoticum Spath (1939: 555, pl. 59 (fig. 7), pl. 60 (fig. 4), pl. 63
(fig. 2), fig. 195) differs from A. armatum in having four to six intercalatories
between main ribs, with very feebly-developed dorsolateral tubercles. The
ribbing of this species is also attenuated across the siphonal line.
In Anisoceras oldhamianum (Stoliczka) (1865: 135, pl. 83 (figs 1-4), pl. 92
(fig. 1)) all the ribs of the early growth stages are tuberculate and presumably
looped, whilst on the straight shaft there is a fine, non-tuberculate rib separating
looped ribs. The early whorls are coiled in a shallow, open helix.
Occurrence
Anisoceras armatum is at present known with certainty only from the
uppermost Albian of southern England, France, Switzerland, Sardinia, Angola,
Texas, and Mexico.
Anisoceras haasi sp. nov.
Figs 20-22
Idiohamites (?) indet. sp., Haas, 1942: 195, pl. 45 (fig. 6).
Idiohamites spiniger Haas (non J. Sowerby), 1942: 195, pl. 46 (fig. 4), fig. 31a—b.
Idiohamites indet. sp., Haas, 1942: 197, pl. 45 (fig. 7), fig. 31c.
Idiohamites aff. subspinigero Haas (non Spath), 1942: 197, pl. 46 (fig. 5), fig. 31d-e.
ANNALS OF THE SOUTH AFRICAN MUSEUM
208
‘TX ‘sode}S YIMOIS I[PPIUI puv ATIvSd JO UOTJONI}SUODEY “AOU ‘ds IspDY SDAQIOSIUP “OZ “BIZ
inh
ee oe,
fo
we eo mmctcqea
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 209
Fig. 21. Anisoceras haasi sp. nov. A-C. USNMNH 236923. D, F, H. USNMNH 236924,
E, G. USNMNH 236922. I-J. USNMNH 236925. x1.
210 ANNALS OF THE SOUTH AFRICAN MUSEUM
Material
Seven specimens, USNMNH 236922-7, all retaining recrystallized shell and
all from Porto Amboim, together with a single crushed internal mould,
SAM-PCA2952 from Praia-Egito.
Type material
USNMNH 236922 is designated holotype; the remaining specimens from
Porto Amboim are paratypes.
Etymology
Named for Otto Haas, pioneer of ammonite studies on the Angola
Cretaceous.
Diagnosis
A densely ribbed species of Anisoceras with 8-15 ribs in a distance equal to
the whorl height; 2 or 3 ribs are looped from small, acute dorsolateral tubercles
to prominent rounded bases of septate ventrolateral spines, commonly with
3-4 looped between tubercles across the venter; there are between 2 and 9 fine
intercalatories separating looped ribs. Whorl section circular to elliptical,
compressed.
Description
Up to 13 mm whorl-width: at this stage, the shell is coiled in a loose planispire,
and has a slightly depressed to circular cross-section. Ornament comprises fine,
slightly rursiradiate ribs, about as wide as the interspaces and effaced across the
dorsum. There are periodic broad bulges which follow the course of the ribbing
and are ornamented with very weak, pointed dorsolateral tubercles and the
septate bases of large, prominent, ventrolateral spines. The fine ribs commonly
arise in pairs from the dorsolateral tubercles and are joined by a third rib to
meet the ventrolateral spine-base. Three to four ribs are looped across the
venter connecting the ventrolateral spines. There are four to nine fine inter-
calated ribs between adjacent bulges.
14-16 mm whorl-width: beyond 13 mm whorl width, the shell begins to
uncoil, developing an almost straight shaft. At the same time, the whorl section
becomes slightly compressed (“/H = 0,83-0,86) and elliptical. The bulges
become more regular with generally three in a distance equal to the whorl
height, and commonly with two to three fine ribs separating adjacent bulges.
Ribbing remains rursiradiate at this stage and the dorsolateral tubercles have
migrated up flank to a low lateral position.
Discussion
The lectotype of A. pseudoelegans (Renz 1968, pl. 14 (fig. 12)) differs from
the Angolan material in having a strongly compressed whorl section with
flattened flanks and maximum width near the dorsolateral tubercles, whilst it is
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 211
Fig. 22. Anisoceras haasi sp. nov. Dorsal and lateral views of a
slightly crushed composite internal mould, SAM-—PCA2952,
from Egito. x1.
more coarsely ribbed, with ribs arising in bundles from distinctly bullate dorso-
lateral tubercles. Juveniles of A. pseudoelegans also appear to be much more
coarsely ribbed than the present material (compare Renz 1968, pl. 14 (figs 10-11)).
According to Kennedy (1971: 12, pl. 3 (figs 12-13), pl. 4 (figs 1-3)),
Anisoceras plicatile is a Middle Cenomanian species with 12 fine ribs in a
distance equal to the whorl height, and a circular whorl section. 2 or 3 ribs are
looped between the midlateral and ventrolateral tubercles and are separated by
1-3 non-tuberculate intercalatories. Anisoceras plicatile differs from the present
material, therefore, in having much more swollen main ribs and a much more
prominent lateral tubercle which is at the middle of the flank, not dorso-
lateral as in A. haasi. The Angolan species also has more numerous, fine
intercalatories, whilst the ribbing is not as coarse in maturity as it is in A. plicatile
(compare Kennedy 1971, pl. 2 (fig. 12)).
Anisoceras bendirei (Adkins) (1920: 8, pl. 11 (fig. 1)) from the late Albian of
Texas differs from the Angolan material in being much more coarsely ribbed,
with the lateral tubercle higher on the flanks.
Anisoceras raynaudi (Boule, Lemoine & Thévenin 1907: 170, pl. 4 (figs 7-8),
fig. 38) is a finely ribbed species in maturity, which differs from A. Aaasi in
having fewer intercalatories (only 2-3) between looped ribs in the early growth
stages, a more prominent dorsolateral tubercle and, judging from material from
Catuane, southern Mozambique, in the South African Museum, Cape Town, in
212 ANNALS OF THE SOUTH AFRICAN MUSEUM
having the early whorls coiled in an open helical spire up to 20 cm in height.
It is also an earlier species (orbignyi-varicosum Subzones).
Anisoceras nanaense (von Hauer) (1861, pl. 1 (figs 11-12)) differs from the
present species in lacking dorsolateral tubercles, and in having prorsiradiate ribs
with only two to three intercalatories between tuberculate ribs.
Anisoceras arrogans (Giebel) (1852: 305) (nom. nov. pro Hamites elegans
d’Orbigny (non Parkinson) 1842: 542, pl. 133 (figs 1-5)) can be distinguished
from A. haasi in its lack of dorsolateral tubercles, with only two to four non-
tuberculate intercalatories, and in the coarse, distant ribbing of the body
chamber.
Occurrence
Anisoceras haasi sp. nov. is at present known only from the uppermost
Albian of Angola.
Anisoceras phillipsi sp. nov.
Figs 16J-K, 23-24
Material
Eight fragments, SAM—PCA2974, 3179, 3183, 3211-3212, 3220 and 3222,
all preserved as composite internal moulds, from Praia-Egito, and SAM-—
PCA4799 from the Quissama Ridge at Cabo Ledo.
Type material
The specimen illustrated as Figure 23, SAM-—PCA3183 from Praia-Egito is
designated as holotype. All other specimens cited above are paratypes.
Etymology
The species is named for Denis Phillips of the British Museum (Natural
History) who, during many years, has given both authors invaluable assistance
and advice in connection with their researches.
Diagnosis
A large species of Anisoceras with subrectangular, depressed to ovate whorl
section in maturity. Prominent dorsolateral clavae give rise to two, occasionally
only one, rursiradiate ribs which meet the well-developed ventrolateral clavae
singly or in pairs. There are no intercalated ribs. Across the venter the ribs are
looped or single.
Description
In the smallest example, SAM-—PCA3179, the whorl section is elliptical,
compressed, although it may have suffered lateral compaction. All the other
larger fragments show a strongly depressed, subrectangular whorl section
(Fig. 24) prior to the final hook, and an ovate whorl section afterwards.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 213
Fig. 23. Anisoceras phillipsi sp. nov. Dorsal, ventral and lateral views of the holotype,
SAM-PCA3183, preserved as an internal mould. 0,75.
214 ANNALS OF THE SOUTH AFRICAN MUSEUM
(\O)
Seq
Fig. 24. Anisoceras phillipsi sp. nov. Whorl sections. A-B. SAM-—PCA2974. C. SAM-
PCA3183. D. SAM-—PCA3222. x1.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 215
On the dorsolateral shoulder, prominent clavae give rise to commonly two,
sometimes only one, coarse, rursiradiate flank ribs which, on curved fragments,
are strongly convex adorally. The flank ribs meet the well-developed ventro-
lateral clavi either singly or in pairs, with six to eight ribs in a distance equal to
the whorl height. Across the venter, ribs are single or looped.
Discussion
The only species with which the present form may usefully be compared is
Anisoceras arrogans (Giebel) (= Anisoceras campichei Spath 1942: 559, pl. 63
(figs 6-7), fig. 197) which differs in lacking the dorsolateral clavae and in the
regular nature of its ribbing across the venter.
Occurrence
Anisoceras phillipsi sp. nov. is known only from the uppermost Albian of
Angola.
Anisoceras cf. arrogans (Giebel, 1852)
Figs 25-26
Compare
Hamites elegans d’Orbigny (non Parkinson), 1842: 542, pl. 133 (figs 1-5).
Hamites arrogans Giebel, 1852: 305.
Anisoceras pseudoelegans Pictet & Campiche, 1861: pl. 50 (figs 6-7 only).
Anisoceras campichei Spath, 1926: 432; 1939: 559, fig. 197 only.
Metahamites (?) arrogans (Giebel) Spath, 1939: 559.
Anisoceras aff. campichei Spath, 1939: pl. 63 (figs 6-7).
Anisoceras arrogans (Giebel) Wiedmann & Dieni, 1968: 69, pl. 7 (fig. 10),
pl. 8 (figs 5, 7, 11), figs 46-50.
Description
This species is known only from large fragments. In SAM-—PCA3205, the
whorl section is slightly compressed, subquadrate, but the remaining material
shows some variation in this character.
Ornament comprises prominent, single flank ribs arising from weak
umbilical tubercles and passing slightly prorsiradiate across the flanks to the
bases of large, septate ventrolateral spines. Ribs are either single or looped across
the venter, and the rib pattern is shown in Figure 26. There are six to seven flank
ribs in a distance equal to the whorl height. Across the dorsum, fine ribs are
looped between the weak dorsolateral tubercles, with generally an intercalated
rib between pairs, so that there are about fifteen ribs across the dorsum per
five dorsolateral tubercles.
Discussion
The present specimen is closest to Anisoceras arrogans (Giebel)
(= A. campichei Spath, 1939: 559, fig. 197 only) from which it differs in its
subquadrate whorl section, slightly prorsiradiate flank ribs, and the fact that all
216 ANNALS OF THE SOUTH AFRICAN MUSEUM
ribs across the venter arise from ventrolateral tubercles. Anisoceras arrogans
also does not show the peculiar zigzagging of the ribs across the venter (see
Fig. 26) seen in the Angolan material. However, since A. arrogans and the
present material represent different ontogenetic stages, the differences may not
be as great as they first appear, and consequently the present material is
identified as A. cf. arrogans (Giebel).
Fig. 25. Anisoceras cf. arrogans (Giebel). Dorsal, ventral and lateral views of SAM—PCA3205.
x 0,75.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 217
Fig. 26. Anisoceras cf. arrogans (Giebel). Schematic
rib pattern across the venter of SAM-—PCA3205.
x 0,75.
Anisoceras phillipsi sp. nov. differs from the present material in the posses-
sion of distinctly clavate dorsolateral tubercles, from which ribs frequently arise
in pairs.
Occurrence
Anisoceras arrogans is known from the uppermost Albian of Switzerland,
France and Sardinia, and may also be present in southern England and Angola.
Anisoceras aff. exoticum Spath, 1939
Fig. 27
Compare
Anisoceras saussureanum Pictet & Campiche (non Pictet), 1861: 118, pl. 1 (fig. 2).
Anisoceras exoticum Spath, 1939: 555, pl. 59 (fig. 7), pl. 60 (fig. 4), pl. 63 (fig. 2),
fig. 195.
Material
A single specimen, SAM-PCA3174, preserved as a composite internal
mould, from Praia-Egito.
218 ANNALS OF THE SOUTH AFRICAN MUSEUM
Description
The specimen represents a slightly crushed fragment of a straight shaft in
which the whorl section was originally probably almost circular. Ornament
comprises slightly rursiradiate main ribs between which are three to seven fine,
secondary ribs, the density of which increases adorally. The fragment lacks
obvious tuberculation.
Fig. 27. Anisoceras aff. exoticum Spath. A body chamber fragment,
SAM-PCA3174. 0,75.
Discussion
The specimen is a body chamber fragment and, since ornament frequently
changes drastically on the body chamber of Anisoceras, reference to Spath’s
species is somewhat tentative.
Occurrence
Anisoceras exoticum is known from England, Switzerland, Sardinia, and
possibly Angola.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 219
Anisoceras aff. subarcuatum Spath, 1939
Fig. 28
Compare
Anisoceras subarcuatum Spath, 1939: 560, pl. 65 (fig. 1), pl. 66 (fig. 1), fig. 198.
Material
A single specimen, SAM-—PCA3143, from the dispar Zone at Praia-Egito.
Description
A short fragment shows affinities with Spath’s species and may belong here.
The whorl section is slightly compressed, oval. Ornament comprises slightly
tursiradiate, strengthened, simple main ribs which bifurcate or trifurcate across
the dorsum and are ornamented with dorsolateral and ventrolateral tubercles.
Between main ribs are two to three, slightly weaker, somewhat irregular
secondary ribs, which occasionally bifurcate across the venter. All the secondary
ribs lack tubercles.
Discussion
Body chamber fragments of Anisoceras are notoriously difficult to identify
and hence full determination of the present specimen must await further material.
Fig. 28. Anisoceras aff. subarcuatum Spath. Ventral, lateral and dorsal views of SAM-PCA3143.
x 0,75.
220 ANNALS OF THE SOUTH AFRICAN MUSEUM
Occurrence
Anisoceras subarcuatum is known with certainty only from the low Upper
Albian (varicosum and auritus subzones) of southern England, but may also be
present in the uppermost Albian of Angola.
Anisoceras aff. spathi (Wiedmann, 1962)
Fig. 29
Compare
Anisoceras subarcuatum Spath, 1939: 560, pl. 63 (fig. 5 only).
Idiohamites spathi Wiedmann, 1962a: 188.
Material
A single fragment, SAM-—PCA3223, from the upper part of the dispar Zone
at Praia-Egito, preserved as a composite internal mould.
Fig. 29. Anisoceras aff. spathi (Wiedmann). Ventral and lateral views of SAM—PCA3223.
0315:
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 221
Description
The single fragment available represents part of the crozier. It has a
compressed, elliptical whorl section with simple rursiradiate ribs. On the adoral
portion of the fragment all ribs are relatively fine, of even strength, and lack
tuberculation. On the adapical portion, every second rib is strengthened slightly
and bears weak ventrolateral tubercles.
Discussion
The absence of lateral or dorsolateral tubercles led Wiedmann (1962a) to
assign the present species to Idiohamites. However, the holotype is a body-
chamber fragment which merely shows the radical change in ornament on the
body chamber displayed by many Anisoceras species.
Occurrence
Anisoceras spathi is known from the low Upper Albian of southern England
and Spain, and may be present in the dispar Zone of Angola.
Anisoceras sp. indet.
Fig. 14F
Material
A single, crushed fragment, SAM-—PCA3220, from Praia-Egito, preserved
as a composite internal mould.
Description
Only the greater part of one flank, one row of ventrolateral tubercles and
much of the venter of the specimen concerned is preserved.
Rather fine (?looped) ribs on the flanks are slightly narrower than the
interspaces, with at least 10 in a distance equal to the whorl height. The flank
ribs join the ventrolateral clavi in pairs whereas, across the venter, the ventro-
lateral tubercles are connected by 2-3 ribs so that for 16 flank ribs, there are
20 ribs across the venter. The whorl section gives the appearance of having been
strongly depressed.
Discussion
The specimen under discussion is too poorly preserved for specific identifi-
cation, but may be regarded as belonging to the plicatile-haasi plexus.
Occurrence
Upper Albian, dispar Zone, Praia-Egito.
222 ANNALS OF THE SOUTH AFRICAN MUSEUM
Genus Idiohamites Spath, 1925
Type species Hamites tuberculatus J. Sowerby, 1818
Discussion
Wiedmann (1962a) separated Idiohamites from Anisoceras because the
former was considered to have the early whorls coiled in a helix, whereas in
Anisoceras the juvenile whorls were considered to be planispirally coiled. The
differences are not, however, so clear cut, since Anisoceras of the type referred to
A. raynaudi (Boule, Lemoine & Thévenin) by Forster (1975) (= Anisoceras
saussureanum quadrifasciatum Klinger) from the low Upper Albian of Catuane,
Mozambique, and now in the South African Museum, show the early whorls to
be coiled in an open helix up to 20 cm high. The straight final shafts of this
species are figured by Forster (1975, pl. 4 (fig. 8)). Moreover, Matsumoto
(1959, pl. 28 (fig. 1), pl. 29 (fig. 2)) has also figured an Anisoceras in which the
early growth stages are helically coiled.
Nor is the presence or absence of lateral tubercles a diagnostic feature since
some Anisoceras, e.g. A. auberti (Pervinquiére) and A. gracile Renz, lack them.
Moreover, Renz (1968) has recently figured a number of specimens which he
considers transitional between Jdiohamites and Anisoceras in this respect
(cf. pl. 12 (figs 7-8), pl. 13 (figs 3, 6)). In addition both genera show a suture line
with a fairly shallow, bifid external lobe (E), an asymmetrically bifid first lateral
saddle (E/L), a bifid first lateral lobe (L) which is deeper than the external lobe,
and a second lateral saddle (L/U) which is bifid, and a bifid second lateral
lobe (U) which is almost as deep as the first. Clearly the two genera are very
closely related although, at present, the typically smaller size, almost complete
absence at any growth stage of looped flank ribs, and the helically coiled
juvenile whorls, are considered to be sufficient for the generic separation of
Idiohamites from Anisoceras.
Idiohamites dorsetensis Spath, 1926
Figs 30, 31G
Anisoceras alternatus Pictet & Campiche (non Mantell), 1861: 71, pl. 51 (figs 1, 3-4).
Idiohamites dorsetensis Spath, 1926b: 432; 1939: 596, pl. 62 (figs 2-3), pl. 63 (figs 1, 9, 15),
pl. 65 (fig. 2), fig. 215. Renz, 1968: 70, pl. 11 (figs 39-40), pl. 12 (figs 3-4), figs 25a-d, f,
26a-d.
? Idiohamites aff. turgidus robustus Spath, Renz, 1968: 72, pl. 11 (figs 33-37), figs 25k—I, 26f-h.
? Idiohamites elegantulus laticostatus Renz, 1968: 73, pl. 11 (figs 38, 41-42), pl. 12 (figs 1-2),
figs 25m, 26i-m.
Material
Three specimens in the South African Museum, SAM-—PCA4803 and two
unnumbered fragments, retaining recrystallized test, together with USNMNH
236951, preserved as a composite internal mould from Porto Amboim, and a
single specimen from Cabo Ledo, SAM-PCA5469.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 223
Fig. 30. A-—D. Idiohamites dorsetensis Spath. A-B. Ventral and lateral views
of SAM-PCA4803. C-—D. Ventral and lateral views of SAM-PCAS5S469. x1.
Description
All the fragments show the shell to be loosely coiled in a single plane, with
an elliptical, compressed whorl section. Ornament comprises simple, strong,
slightly prorsiradiate ribs, of which there are four to five in a distance equal to
the whorl height. Every second or third (or occasionally adjacent) rib bears very
small, sharp, pointed lateral tubercles and more prominent ventrolateral spines.
At small growth stages, tubercles are connected across the venter by simple ribs
but, on the mature body chamber, they may be weakly looped.
224 ANNALS OF THE SOUTH AFRICAN MUSEUM
Discussion
Until populations of individuals are available for study, the differences
between several named species of /diohamites appear suspiciously small. The
authors assign their material to J. dorsetensis because of the very close similarities
to the types, and because it is the oldest available name for material of this age.
Occurrence
Idiohamites dorsetensis is known from the uppermost Albian of southern
England, France, Switzerland and Angola.
Idiohamites cf. elegantulus Spath, 1939
Fig. 31A—D
Compare
Idiohamites elegantulus Spath, 1939: 599, fig. 216.
Material
One specimen, USNMNH 236950, a composite internal mould from Porto
Amboim.
Discussion
The specimen consists of a body chamber hook with a maximum whorl
height of 12 mm. There are six fine prorsiradiate ribs in a distance equal to the
whorl height, the majority bearing ventral tubercles, suggesting reference to
Spath’s species.
Occurrence
Stoliczkaia dispar Zone of England, and possibly Angola.
Idiohamites pygmaeus sp. nov.
Fig. 31J-N
Material
Eight specimens, USNMNH 236942-49, all retaining recrystallized shell
and all from Porto Amboim.
Type material
USNMNH 236942, a complete adult, is designated holotype; the remaining
specimens are paratypes.
Etymology
From the Latin adjective pygmaeus, pygmy-like, dwarfish; derived from the
Greek pygmaios. The pygmaioi were a fabulous dwarfish race of antiquity,
especially in Africa; at war with the cranes, they were constantly defeated
(Pliny).
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 225
L
Fig. 31. A-D. Idiohamites cf. elegantulus Spath. A-B. USNMNH 236948. C-D. USNMNH
236938. E-F. Mariella (Mariella) cirumtaeniatus (Kossmat), USNMNH 236958. G. Idiohamites
cf. dorsetensis Spath,h USNMNH 236951. H-I. Borissiakoceras sp. noy.? aff. reymenti
(Brunnschweiler), USNMNH 236980. J-N. Jdiohamites pygmaeus sp. nov. J-L. USNMNH
236942. M-N. USNMNH 236943, x1.
226 ANNALS OF THE SOUTH AFRICAN MUSEUM
Diagnosis
A very small Jdiohamites with a known maximum length of only 52 mm,
characterized by helicoid early whorls with three quadrituberculate rursiradiate
ribs in a distance equal to the whorl height, a straight shaft with four prorsi-
radiate ribs in a distance equal to the whorl height and ribbing tending to be
rectiradiate at the aperture.
Description
Shell small, initially coiled in a low, open helical spire, straightening in
maturity before recurving on the body chamber to form a crozier.
Earliest whorls ornamented with rather robust ribs, narrower than the
interspaces and with three in a distance equal to the whorl height, which cross
the venter obliquely so that the four rows of tubercles on the main ribs are
asymmetrically arranged. There is a regular alternation of tuberculate and non-
tuberculate ribs, which, at this stage, are slightly rursiradiate. On the first
straight shaft, which commences at a whorl height of 6 mm, the ribbing becomes
prorsiradiate, with still only three ribs in a distance equal to the intercostal
whorl height. With the uncoiling of the shell, the tuberculation is no longer
asymmetrical and main ribs are ornamented with small pointed dorsolateral and
ventrolateral tubercles. The rib direction changes to rectiradiate on the final
shaft. In maturity, the intercostal whorl section is oval, depressed, although it is
almost circular at small diameters.
Discussion
This species is characterized by its small size at maturity. Idiohamites
dorsetensis 1s adult at much larger diameters, with a slightly compressed whorl
section, and closer ribbing (four to six in a distance equal to the whorl height).
Idiohamites elegantulus is even more densely ribbed (seven in a distance equal to
the whorl height), also with a compressed whorl section. /diohamites recticostatus
Renz is a much larger species with an almost circular whorl section and four
rectiradiate ribs in a distance equal to the whorl height. It closely approaches the
present material, but its very much larger size (septate at 23 mm whorl height)
is distinctive.
Occurrence
Uppermost Albian of Angola only.
Family Hamitidae Hyatt, 1900
Genus Hamites Parkinson, 1811
Type species Hamites attenuatus J. Sowerby, 1814
Discussion
Recent discussions of the family are to be found in Wiedmann & Dieni
(1968) and Klinger (1976), and further discussion on the material available here
is unnecessary.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 227
Hamites virgulatus Brongniart, 1822
Figs 16F, H, 32B—C
Hamites virgulatus Brongniart (in Cuvier & Brongniart), 1822: pl. 0 (fig. 6).
Hamites (Stomohamites) virgulatus (Brongniart ?) Pictet & Campiche, Spath, 1941: 635, pl. 71
(figs 7-10), pl. 72 (fig. 11), fig. 230 (with synonymy). Renz, 1968: 65, pl. 11 (figs 9-11),
fig. 23b-d (with synonymy).
Hamites (Stomohamites) subvirgulatus Spath, 1941: 645, fig. 234. Renz, 1968: 66, pl. 11
(figs 13-14), figs 23e, 24a (with synonymy).
Stomohamites brongniarti Breistroffer, 1947: 77.
Hamites (Hamites) virgulatus Brongniart, Wiedmann & Dieni, 1968: 53, pl. 5 (figs 1-2, 10),
pl. 7 (figs 1-2), figs 21-27. Klinger, 1976: 60, pl. 23 (figs 4-5, 8), figs 8 I,n—o, 11i (with
synonymy).
Hamites venetzianus Pictet (in Pictet & Roux), 1847: 134, pl. 14 (fig. 6).
Hamites (Stomohamites) venetzianus Pictet, Spath, 1941: 638, pl. 71 (figs 11-13), fig. 231.
Renz, 1968: 67, pl. 11 (figs 15-16), figs 23f, 24b (with synonymy).
Material
Ten composite internal moulds, SAM-—PCA2959, 2963-64, 2966, 2971,
3118, 3157-58, and 3371 from the upper part of the dispar Zone at Praia-Egito,
three fragments retaining recrystallized tests from Porto Amboim, USNMNH
236955-7, and SAM-PCA4603 from Cabo Ledo.
Description
The whorl section is elliptical compressed, with three to six prorsiradiate
ribs in a distance equal to the whorl height. The ribs are effaced across the
dorsum.
Discussion
The writers follow Wiedmann & Dieni (1968) in regarding H. brongniarti,
H. venetzianus, and H. subvirgulatus as synonyms of H. virgulatus, although
there may be some justification for retaining venetzianus at the varietal level for
those variants in which the ribs are as thick as, or thicker than, the interspaces.
Hamites duplicatus Pictet & Campiche, 1861
Figs 16D, 32A
Hamites virgulatus Pictet (non Brongniart) (in Pictet & Roux), 1847: 391, pl. 14 (figs 7, 9 only).
Hamites duplicatus Pictet & Campiche, 1861: 98.
Hamites (Stomohamites) duplicatus Pictet & Campiche, Spath, 1941: 640, pl. 72 (figs 12-16),
fig. 232. Renz, 1968: 68, pl. 11 (figs 19-21), fig. 23h-k.
Material
A single composite internal mould from Praia-Egito, SAM-—PCA2955.
Description
The whorl section is oval, compressed, with a somewhat flattened dorsum.
Ornament comprises fine, rursiradiate ribs, seven in a distance equal to the
whorl height, which are very weak across the dorsum.
228 ANNALS OF THE SOUTH AFRICAN MUSEUM
Discussion
The writers are not convinced by Wiedmann & Dieni’s (1968) argument for
the inclusion of H. duplicatus in the synonymy of H. virgulatus and, for the
present, maintain it as distinct.
Occurrence
Hamites duplicatus is known from the Upper Albian of England, France,
Switzerland, and Angola.
Superfamily TURRILITACEAE Meek, 1876
Family Turrilitidae Meek, 1876
Subfamily Turrilitinae Meek, 1876
Genus Mariella Nowak, 1916
Type species Turrilites bergeri Brongniart, 1822
Discussion
Kennedy (1971) and Klinger & Kennedy (1978) have recently provided a
comprehensive discussion of the taxonomic standing of Mariella, including in it
three subgenera, viz. M. (Mariella), M. (Plesioturrilites) and M. (Wintonia).
As noted by Clark (1965: 49), however, M. (Wintonia) and M. (Plesio-
turrilites) differ only in that the former has an early, straight shaft which then
passes into the helical coiling of M. (Plesioturrilites), although specimens of
M. (Wintonia) graysonensis (Adkins), the only known species, cannot be
distinguished from M. (Plesioturrilites) bosquensis (Adkins) in the absence of
this straight shaft. Klinger & Kennedy (1978) suggest the ‘shaft’ is an artefact of
preservation and treat Plesioturrilites as a synonym of Wintonia.
Mariella (Mariella) circumtaeniatus (Kossmat, 1895)
Figs 31E-F, 39H
Turrilites gresslyi Stoliczka (non Pictet & Campiche), 1865: 186, pl. 87 (figs 1-5, ? non 2).
Turrilites circumtaeniatus Kossmat, 1895: 141, pl. 18 (figs 4-5); Boule, Lemoine & Thévenin,
1907: 57, pl. 13 (fig. 4).
Non Turrilites circumtaeniatus Scott (non Kossmat), 1926: 145, pl. 1 (figs 10-11)
(= M. worthensis).
Paraturrilites aff. circumtaeniatus (Kossmat) Collignon, 1963: 46, pl. 258 (fig. 1120).
Non Turrilites circumtaeniatus Kossmat, Woods, 1917: 11, pl. 5 (figs 2-3) (= M. thomsoni
Henderson).
Material
A single specimen USNMNH 236958, with recrystallized shell preserved,
from Porto Amboim.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 229
@yonc
Fig. 32. A. Hamites duplicatus Pictet & Campiche.
Whorl section of SAM-PCA2955. B-C. Hamites
virgulatus Brongniart. Whorl sections. B. SAM-
PCA2959. C. SAM-PCA3158. x1.
Description
Coiling sinistral, with a very acute apical angle (approximately 20°). Flanks
rounded intercostally, with two rows of prominent, rounded tubercles and a
third hidden in the spiral seam. There are nine to eleven tubercles per half-whorl
situated on very weak, slightly oblique ribs. The upper row of tubercles is
situated slightly above mid-flank, and the central row midway between the
upper row and the lower spiral suture. Ribbing is conspicuous only on the area
between the upper row of tubercles and the upper spiral suture, where rather fine
ribs commonly connect in pairs to the upper row of tubercles, frequently with an
intercalated rib between tubercles.
Discussion
Amongst contemporaneous species of Mariella, M. (M.) cantabrigiensis
(Jukes-Browne) (Spath 1937: 518, pl. 57 (fig. 36), pl. 58 (figs 1-2), figs 18la—b,
182d-e) approaches the present species most closely. It differs in its much larger
spiral angle (30° or more) and in lacking the fine looped ribs on the upper, outer
face of the whorls.
M. (Mariella) nobilis (Jukes-Browne) (Spath 1937: 520, pl. 58 (figs 10-11),
fig. 182) differs in having more prominent single ribs and subdued tuberculation.
M. (Mariella) bergeri (Brongniart) (1822: 395, pl. 7 (fig. 3)), of which
M. (M.) miliaris (Pictet & Campiche) (1861: 136, pl. 51 (fig. 5)) may be only an
extreme variant, has four rows of tubercles exposed on the outer face of the
whorls.
M. (Mariella) dubourdieui (Collignon) (1963: 46, pl. 285 (fig. 1121))
resembles M. (M.) circumtaeniatus in the possession of looped ribs, but also has
four rows of tubercles exposed on the outer face of the whorls.
M. (Mariella) worthensis (Adkins & Winton) (1920: 44, pl. 7 (figs 10-11, 13))
from the uppermost Albian Pawpaw Formation of Texas, closely resembles
M. (M.) circumtaeniatus, but appears to lack the fine looped ribs of Kossmat’s
species, as does M. (M.) hillyi (Dubourdieu) (1953: 46, pl. 4 (figs 1-3)).
Occurrence
This species is known from the uppermost Albian of India, Madagascar,
Zululand, Angola, and New Zealand.
230 ANNALS OF THE SOUTH AFRICAN MUSEUM
Mariella (Mariella) nobilis (Jukes-Browne, 1877)
Fig. 33A-B
Turrilites intermedius Pictet & Campiche, 1861: 127, pl. 57 (fig. 15 only).
Turrilites escherianus Pictet & Campiche (von Pictet), 1861: 130, pl. 56 (figs 6-8 only).
Turrilites nobilis Jukes-Browne, 1877: 493, pl. 21 (fig. 1).
Mariella nobilis (Sukes-Browne) Breistroffer (in Besairie), 1936: 147. Spath, 1937: 520, pl. 58
(figs 10-11), fig. 182a—c. Breistroffer, 1940: 147. Clark, 1965: 40, pl. 10 (figs 2-4), pl. 11
(figs 4-5). Renz, 1968: 89, pl. 17 (figs 28, 40), figs 311, 32k.
Mariella nobilis var. cruciana Breistroffer (in Besairie), 1936: 148. Spath, 1937: 521. Breistroffer,
1940: 148. Renz, 1968: 90, pl. 17 (fig. 34), figs 31b, 32d.
Mariella aff. nobilis (Jukes-Browne) Spath, 1937: 520, pl. 58 (fig. 21).
Paraturrilites nobilis (Sukes-Browne) Breistroffer, 1947: 60; 1953: 1350.
Paraturrilites nobilis var. cruciana (Breistroffer) Breistroffer, 1947: 60.
Paraturrilites nobilis var. brownei Breistroffer, 1947: 96.
Mariella nobilis brownei (Breistroffer) Renz, 1968: 90, pl. 17 (figs 35-36, 38-39), figs 31i, 32e.
Material
Two specimens, SAM-—6531 (from an unknown location on the Angolan
littoral), and a single fragment in the Washburn collection, USNMNH 237019
from Porto Amboim.
Description
These two fragments of M. (Mariella) bear prominent oblique ribs,
narrower than the interspaces, estimated at totalling twenty-six per whorl. They
are ornamented by three rows of weak tubercles; the tubercles of the adoral row
are weakly clavate, whereas those of the other two rows are weakly rounded.
Discussion
M. (Mariella) nobilis var. cruciana Breistroffer (in Besairie 1936: 148) was
separated from the typical form by its denser ribbing (30 ribs per whorl, as
against 26-28) and more prominent tubercles. M. (Mariella) nobilis var. brownei
Breistroffer (1947: 96) was distinguished by its sparser ribbing (24-26 per whorl)
and weaker tuberculation.
As noted by Spath (1937: 521), M. (Mariella) escheriana (Pictet) (in Pictet &
Roux 1847: 154, pl. 15 (fig. 11)) closely resembles the present species and there
even appear to be intermediates. Pictet’s species was distinguished by its denser
ribbing (35-40 ribs per whorl), and flattened flanks with only two rows of
tubercles.
The closest species to M. (M.) nobilis is M. (M.) cantabrigiensis (Jukes-
Browne) (Spath 1937: 518, pl. 57 (fig. 36), pl. 58 (figs 1-2), figs 18la—b, 182d-e)
from the late Albian of western Europe. Not only are they strictly contem-
poraneous, but Spath (1937: 519) also admits to the presence of intermediates.
Typically, however, it differs in its sparser ribbing and more prominent tubercles.
M. (Mariella) worthensis (Adkins & Winton) (1920: 44, pl. 7 (figs 10-11, 13))
is very close to M. (M.) nobilis but is generally more coarsely ribbed (14-28 ribs
per whorl) with four rows of tubercles per whorl.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 231
M. (Mariella) gresslyi, M. (M.) cantabrigiensis, M. (M.) nobilis, M. (M.)
escheriana, M. (M.) worthensis and M. (M.) hillyi are a contemporaneous group
of very closely allied species whose intraspecific variation clearly needs
documenting before the true taxonomic status of the species involved can be
resolved.
Occurrence
M. (Mariella) nobilis is known only from the upper Upper Albian of
southern England, Texas, and Angola.
Fig. 33. A-B. Mariella (Mariella) nobilis (Jukes-Browne), SAM-6531. C-D. Puzosia
(Puzosia) sp., SAM-6407. x1.
Mariella gresslyi (Pictet & Campiche, 1861)
Fig. 16G
Turrilites gresslyi Pictet & Campiche, 1861: 132, pl. 57 (figs 11-13). Neumayr, 1875a: 901.
Renevier, 1890: 339. Pervinquiére, 1910: 54. Bose, 1923: 147. Diener, 1925: 83. Collignon,
1929: 65, pl. 1 (fig. 15).
Turrilitoides (2) gresslyi (Pictet & Campiche) Breistroffer, 1936: 65.
Mariella gresslyi (Pictet & Campiche) Spath, 1937: 516, pl. 58 (figs 3-4), fig. 180. Breistroffer,
1940: 149. Renz, 1968: 89, pl. 17 (figs 30a—b, 32-33), figs 31c, 32a—b.
Paraturrilites gresslyi (Pictet & Campiche) Breistroffer, 1947: 60. Collignon, 1963: 47, pl. 258
(fig. 1122).
? Turrilites gresslyi Pictet & Campiche, Boule, Lemoine & Thévenin, 1907: 39, pl. 6 (fig. 2).
Mariella gresslyi bifurcata Renz, 1968: 89, pl. 17 (fig. 42a—b), figs 3le, 32c.
? Turrilites cantabrigiensis Jukes-Browne, 1877: 493.
? Mariella cantabrigiensis (Jukes-Browne) Spath, 1937: 518, pl. 57 (fig. 36), pl. 58 (figs 1-2),
figs 18la—b, 182d-e.
232 ANNALS OF THE SOUTH AFRICAN MUSEUM
Material
A single specimen, SAM-—PCA3133, from low down in the coastal cliffs
immediately north of the estuary at Praia-Egito, and thus from a level below
the main Stoliczkaia occurrence.
Description
The specimen is rather poorly preserved, but retains recrystallized test.
The shell is sinistrally coiled, with a moderately large apical angle. The outer
faces of the whorls are gently convex intercostally, and angular, polygonal
costally. Three rows of tubercles are visible on rather oblique ribs. The upper
row of tubercles are conical whereas the middle row is distinctly clavate. The
nature of the tubercles of the lower row was difficult to discern, but they, too,
appear to be clavate.
Discussion
The features of the present specimen are those of M. gresslyi and the
authors have no hesitation in assigning the Angolan example to this species.
Mariella cantabrigiensis (Jukes-Browne) (Spath, 1937: 518, pl. 57 (fig. 36),
pl. 58 (figs 1-2), figs 180h, 18la—b, 182d-e) closely approaches M. gresslyi, but
was said to differ in having all tubercles rounded, whereas the lower two rows
are clavate in M. gresslyi. However, the two species are strictly contemporaneous
and occur side by side, whilst Spath (1937) admits to transitions between the
two. Population studies may show M. cantabrigiensis to be an intraspecific
variant of M. gresslyi.
Mariella gresslyi bifurcata Renz (1968: 89, pl. 17 (fig. 42), figs 3le, 32c) is
based on a single specimen which shows the ribs on the base of the spire
bifurcating from the lowest (adoral) row of tubercles.
Mariella nobilis (Jukes-Browne) (Spath 1937: 520, pl. 58 (figs 10-11),
figs 18lc, 182a—c) differs from the present species in having very subdued
tubercles, with ribs dominant.
Occurrence
Mariella gresslyi is currently known from southern England, France,
Switzerland, Algeria, Angola, Zululand, and Madagascar.
Mariella cf. oehlerti (Pervinquiére, 1910)
Figs 3H, 34
Compare
Turrilites gresslyi Boule, Lemoine & Thévenin (non Pictet & Campiche),
1907: 57, pl. 13 (fig. 2). Collignon, 1929: 65, pl. 6 (fig. 15).
? Turrilites cf. gresslyi Spath (non Pictet & Campiche), 1921: 289.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 233
Turrilites oehlerti Pervinquiére, 1910: 53, pl. 5 (figs 14-17). Collignon, 1929: 65,
pl. 6 (figs 16-17); 1964: 15, pl. 320 (figs 1398-1399).
? Mariella malgachensis Breistroffer, 1940: 79.
? Turrilites bergeri Choffat (non Brongniart), 1903: 15, pl. 1 (figs 4-6).
? Mariella bergeri var. conduciensis Breistroffer, 1940: 149.
Mariella (Mariella) oehlerti (Pervinquiére), Férster 1975: 190, pl. 7 (figs 7-8),
fig. 52. Klinger & Kennedy 1978: 31, pl. 3 (fig. E), pl. 4 (fig. E), pl. 6
(figs H, N), pl. 7 (fig. G), pl. 8 (figs G-H), figs la—b, 7b, d, 8g.
Material
A single composite internal mould, SAM-—PCA4798, from the Quissama
Ridge at Cabo Ledo.
Fig. 34. Mariella (Mariella) cf. oehlerti (Pervinquiére),
SAM-PCA4798. x1.
Description
The shell is a high-spired, sinistrally-coiled turriliticone, with a spiral angle
of about 22°. The whorls are just touching with the outer face gently convex, and
somewhat flattened. The adapical shoulder is abrupt and subangular (Fig. 3H)
whilst the adoral shoulder is evenly rounded. The outer face is ornamented with
234 ANNALS OF THE SOUTH AFRICAN MUSEUM
four rows of prominent tubercles arranged on weak, oblique ribs, of which there
are about thirty per whorl. The upper row of tubercles is the most prominent,
and they are slightly bullate and situated slightly above mid-whorl. The tubercles
of the two middle rows are conical and are separated by a weak spiral groove.
The tubercles of the lower row are the smallest and are situated in the whorl
seam. On the adoral face of the final whorl there are prominent ribs. The
spacing of the tubercles is subequal on the penultimate whorl of the present
specimen, but on the final whorl the middle two rows are distinctly closer
together than the others.
Discussion
At present the species M. gresslyi—M. cantabrigiensis—M. oehlerti—
M. circumtaeniatus are not well differentiated.
Mariella gresslyi (Pictet & Campiche) (1861: 132, pl. 57 (figs 11-13)) is
typically an uppermost Albian species which differs from the present form and
M. oehlerti in that the rows of tubercles are typically clavate. However, Spath
(1937: 519) records passage form between M. gresslyi and the contemporaneous
M. cantabrigiensis (Jukes-Browne) (Spath 1937: 518, pl. 51 (fig. 36), pl. 58
(figs 1-2)), the latter distinguished from Pictet & Campiche’s species in having
rounded tubercles of equal size, and thus very close to M. oehlerti and the
Angolan material. Since, however, the present specimen is much larger than
known material of M. gresslyi and M. cantabrigiensis, differences may be due
to a comparison of different ontogenetic stages. Clearly, however, this problem
cannot be resolved in the present paper.
Mariella circumtaeniatus has only three rows of tubercles, generally with
conspicuous looping of the ribs on the adapical shoulder of the whorls.
Mariella dorsetensis (Spath) (= Turrilites bergeri Sharpe (non Brongniart)
1857: 65, pl. 26 (fig. 11 only)) differs from the present material in having fewer
ribs per whorl and in having the third and fourth (adoral) rows of tubercles
approximated, whilst ribs are absent on the base of the spire.
Mariella gallienii (Boule, Lemoine & Thévenin) (1906: 60, pl. 14 (figs 5—6))
differs from the present specimen in that the tubercles of the four rows are
distinctly bullate.
Occurrence
Mariella oehlerti is known from the Lower Cenomanian of Algeria,
Madagascar, Mozambique, Zululand, and possibly Japan, and possibly the
uppermost Albian of Angola and Switzerland.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 235
Suborder AMMONITINA Hyatt, 1889
Superfamily HOPLITACEAE Douvillé, 1890
Family Desmoceratidae Zittel, 1895
Subfamily Puzosiinae Spath, 1922
Genus Puzosia Bayle, 1878
Subgenus Puzosia Bayle, 1878
Type species Ammonites planulatus J. de C. Sowerby, 1827
Puzosia (Puzosia) sp.
Fig. 33C-D
Material
A single crushed specimen, SAM-6407, retaining recrystallized shell, from
Porto Amboim.
Description
The shell is compressed, moderately involute, with about 60 per cent of the
preceding whorl covered by the outer whorl. The umbilicus is moderately
narrow (22% of the diameter), with steep umbilical walls and evenly rounded
umbilical shoulders. The whorl section is oval, compressed (W/H = 0,83) with
an evenly rounded venter. There are an estimated five constrictions per half-
whorl, preceded by a strong ventral rib. Flank and venter also bear faint,
ill-preserved, fine ribs.
Measurements
No. D H W W/H U
SAM-6407 a7 18(49) 15(41) 0,83 +8(22)
Discussion
The specimen cannot be usefully compared with the large number of late
Albian puzosiids known, although if the estimated number of constrictions is
correct, it must approach forms such as P. crebrisulcata Kossmat (1898: 116,
pl. 17 (fig. 4), pl. 18 (fig. 2) and P. malandiandrensis Collignon (1963: 66, pl. 265
(fig. 1156)).
Puzosia (Puzosia) cf. sharpei Spath, 1923
Fig. 35
Compare
Ammonites planulatus Sharpe (non Sowerby), 1854: 29, pl. 12 (fig. 4 only).
Puzosia sharpei Spath, 1923: 46, pl. 1 (figs 11-12), fig. 11b. Renz, 1968: 21,
pl. 1 (figs 4, 8), figs 6b, 7e (with synonymy).
Material
A single fragment, SAM-PCA3141, preserved as an internal mould, from
the dispar Zone of Praia-Egito.
236 ANNALS OF THE SOUTH AFRICAN MUSEUM
Description
The single specimen is a fragment of about one-quarter whorl. The shell
was moderately evolute, with a fairly wide, shallow umbilicus and evenly
rounded umbilical shoulders. The flanks are broad, subparallel and only
slightly convex, with a broadly rounded venter. There are two deep constrictions
on the fragment, separated along the venter by a distance slightly greater than
the whorl height. The constrictions are initially prorsiradiate, but soon recurve
before flexing strongly forwards to form a chevron across the venter.
Discussion
The Angolan material differs from P. sharpei in that the constrictions do
not show as strong a geniculation, and are therefore not as strongly falcate.
Wiedmann & Dieni (1968) included this species in the synonymy of Puzosia
provincialis (Parona & Bonarelli), a lower Middle Albian species. The material
figured by these authors (1968) as P. provincialis shows relict lappet structures
and are, therefore, microconchs. The holotype of P. sharpei, on the other hand,
is still septate at 83 mm diameter and appears to be a macroconch. Consequently,
the authors regard the inclusion of P. sharpei into the synonymy of P. provincialis
as premature, and prefer to maintain Spath’s species as distinct, for the time
being.
Occurrence
Puzosia sharpei is known with certainty only from southern England,
France, Switzerland, and possibly Angola, where it is typical of the dispar Zone.
Fig. 35. Puzosia (Puzosia) cf. sharpei Spath. Lateral and ventral
views of SAM-PCA3141. x1.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 237
Fig. 36. Desmoceras (Desmoceras) latidorsatum (D’Orbigny). The original of D’Orbigny’s
(1841, pl. 80 (fig. 5)) figured specimen from the Middle Albian of France, in the D’Orbigny
collection, Natural History Museum, Paris. x1.
Subfamily Desmoceratinae Zittel, 1895
Genus Desmoceras Zittel, 1884
Subgenus Desmoceras Zittel, 1884
Type species Ammonites latidorsatus Michelin, 1838
Desmoceras (Desmoceras) latidorsatum perinflatum subsp. nov.
Figs 37-38, 39D-F
Desmoceras latidorsatum var. inflata Breistroffer, Haas, 1952: 2, figs 1, 3-10.
Desmoceras reynesianum Haas, 1952: 4, figs 2, 11-13.
Material
Twenty-one specimens, SAM-6414, SAM-PCA2931, 2934, 2968, 3170 and
3172 from Egito, preserved as composite internal moulds, and USNMNH
236961-75 from Porto Amboim with recrystallized shell generally preserved.
Type material
USNMNH 236970 from Porto Amboim is designated holotype; the
remaining specimens are paratypes.
238 ANNALS OF THE SOUTH AFRICAN MUSEUM
Etymology
From the Latin, per—exceedingly, very much, inflatus—swollen; applying
to the strongly inflated shell form.
Diagnosis
A rather small, late Albian (dispar Zone) subspecies of D. latidorsatum in
which the majority of the population comprises strongly inflated individuals
(“/H = 1,10-1,50) which correspond to the D. latidorsatum var. inflatum of
previous workers.
Description
Very involute, cadicone, with a narrow, crater-like umbilicus (16-20% of
diameter). Umbilical walls steep, with gently rounded shoulders. Flanks
flattened (USNMNH 236972) to strongly convex (USNMNH 236964), with
maximum width just below mid-flank. Venter broadly rounded. Shell smooth,
K
Fig. 37. Desmoceras (Desmoceras) latidorsatum perinflatum subsp. nov. A-B. USNMNH
236966. C-E. The holotype USNMNH 236970. F-G. USNMNH 236973. H-J. USNMNH
236768. K-L. USNMNH 236967. M-O. USNMNH 236964. x1.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 239
except for weak constrictions on some specimens (USNMNH 236970). Where
present, constrictions begin at the umbilical seam and pass strongly forwards to
the umbilical shoulder where they recurve strongly, flexing forwards again just
above mid-flank to form a prominent U-shaped tongue across the venter.
Measurements
No. D H W W/H U
USNMNH 236961 21 +9(43) +12(57) 1,33 +4(19)
USNMNH 236962 14 7(50) S561) . “1,23 2,3(16)
USNMNH 236964 27 13(48) 18,3(68) 1,41 +5(18)
USNMNH 236965 21 E5(S4) 14(67) 122 2
USNMNH 236966 23 9,5(41) 13(56) 137 +4,5(20)
USNMNH 236967 20 10,2(51) 12(60) 1,18 +4(20)
USNMNH 236968 12,3 6,2(50) 8,3(67) 1,34 +2(16)
USNMNH 236969 42 22(52) +24(57) 1,09 +8(9)
USNMNH 236970 24 11(46) 15(63) 1,36 4,5(19)
USNMNH 236971 14,6 7,5(51) 9(62) E20 rs
USNMNH 236972 15 6(40) 8,5(57) 1,42 2,7(18)
= 10,5 5(48) 6,5(62) 1,30 2,0(19)
USNMNH 236973 13 +6(46) 8,1(62) 1,35 ?
USNMNH 236975 +14 6,5(46) 8,2(59) 1,26 2,2(16)
Intraspecific variation
The wide range of intraspecific variation shown by D. (Desmoceras)
latidorsatum has long been recognized (D’Orbigny 1941, Pictet (in Pictet &
Roux) 1848, Kossmat 1897, Jacob 1908, Fallot 1910, Spath 1923, Wiedmann &
Dieni 1968) and it would appear that species separation is possible only at the
population level since gross intraspecific variation exceeds subtle differences
between successive faunas. Consequently, the following names, which are used
both at the varietal and subspecific levels by various authors, are based merely
upon individuals within these populations and are of no taxonomic value:
media (Jacob), complanata (Jacob), inflata (Breistroffer), obesa (Reynés),
petkovici Breistroffer, Jongesulcata (Collignon) and angusteumbilicata Haas.
Jacob (1908) studied the intraspecific variation within Desmoceras latidor-
satum from the Balme de Rencurel, a fauna contemporaneous with D’Orbigny’s
(1841) material (Fig. 36) from the ‘Argile 4 Hoplites dentatus Sow.’ of Aube, and
of early Middle Albian age. Within this assemblage, Jacob (1908) recognized the
typical form, as well as his varieties media and complanata which are more
compressed than typical D. Jatidorsatum. In contrast, the present collection,
together with that described by Haas (1952) shows that inflated variants,
normally assigned to the variety inflata Breistroffer, form the vast majority of
the populations from the S. dispar Zone of Angola (Fig. 38), whereas compressed
240 ANNALS OF THE SOUTH AFRICAN MUSEUM
0
ater a —
0 40
SME = a
Fig. 38. Plot of inflation and umbilical ratio against diameter for Desmoceras
latidorsatum perinflatum subsp. nov. Dots = present Angolan material;
open triangles = material described by Haas (1952), and Wiedmann & Dieni
(1968); black triangle = Desmoceras reynesianum (after Haas 1952); square =
Desmoceras latidorsatum latidorsatum (Michelin), (after D’Orbigny 1841).
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 241
forms are entirely absent. It would seem, therefore, that there has been a genuine
shift within the population structure of D. latidorsatum through time, from
predominantly compressed individuals in the Middle Albian to predominantly
inflated variants in the uppermost Albian. The observation that certain
individuals throughout this range can be assigned to Desmoceras latidorsatum s.s.
suggests that the differences are not of specific importance. Since the Angolan
population comprises mainly strongly inflated individuals (W/H = 1,10-1,50),
the differences are sufficient for subspecific separation and the entire Angolan
population (it is the characters of the population which define the subspecies) is
assigned to D. latidorsatum perinflatum subsp. nov.
Desmoceras reynesianum Haas (1952: 4, figs 2, 11-13) was proposed to
replace Ammonites obesus Reynés (non Stoliczka) and is characterized by its
extreme inflation. The Angolan material assigned to this species, however, differs
in being of uppermost Albian age and, as can be seen from Figure 38, merely
represents extreme variants within the present population, and consequently
Haas’s (1952) material is included in the synonymy of D. latidorsatum perinflatum
subsp. nov.
Discussion
Population studies will probably show that D. collignoni Breistroffer (in
Besairie 1936: 170, pl. 16 (fig. 2), fig. 10d), D. inane (Stoliczka) (1865: 121,
pl. 59 (figs 13-14)) and D. chirichense (Pervinquiére) (1907: 152, pl. 6 (figs 17-20))
do not bear separation from D. /atidorsatum. Desmoceras barryae Anderson
(1958: 214, pl. 12 (fig. 2)) and D. merriami Anderson (1902: 103, pl. 6 (figs
135-138)), which was treated as a variety of D. latidorsatum by Breistroffer
(1947: 61), are probably better referred to the subgenus Pseudouhligella.
Occurrence
Desmoceras latidorsatum perinflatum subsp. nov. is currently known with
certainty only from Angola.
Superfamily ACANTHOCERATACEAE Hyatt, 1900
Family Lyelliceratidae Spath, 1921
Discussion
Wright (in Arkell et al. 1957) included the following genera within the
Lyelliceratidae—Prolyelliceras, Lyelliceras, Tegoceras, Neophlycticeras, Stolicz-
kaia (with Faraudiella as a subgenus), Budaiceras and Salaziceras. More recently,
Casey (1965) introduced the subgenus Stoliczkaia (Villoutreysia), and proposed
the new genus Paradolphia for forms from the S. dispar Zone of southern
England said to te transitional between Stoliczkaia and Forbesiceras. Matsumoto
& Inoma (1975: 277) have also introduced the subgenus Stoliczkaia
(Shumarinaia).
242 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 39. A. Stoliczkaia (Stoliczkaia) tenuis Renz. Lateral view of SAM—PCAS5478. B-C.
Stoliczkaia (Stoliczkaia) sp. A fragmentary internal mould, SAM—PCA4805, from Cabo Ledo.
D-F. Desmoceras latidorsatum perinflatum subsp. noy. D-E. Ventral and lateral views of
SAM-PCA2934. F. Lateral view of SAM—PCA2931. G. Mortoniceras (Angolaites) gregoryi
(Spath). Lateral view of a crushed fragment, SAM—PCA3168. H. Mariella aff. circumtaeniatus
(Kossmat). SAM—PCA3130, from low in the sea cliffs (lower part of dispar Zone) at Praia-Egito.
eile
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 243
Amedro (1976) has recently suggested that Paradolphia is better regarded as
a subgenus of Neophylicticeras (along with Protissotia and Eotropitoides), whilst
there is a case for regarding both Paracalycoceras and Cottreauites, both of
which derive from Stoliczkaia, as Lyelliceratidae rather than Acanthoceratidae.
The collection of Stoliczkaia to be discussed below shows a remarkable
similarity to the earliest Forbesiceras, i.e. the beaumontianum-largilliertianum
group (compare Juignet & Kennedy 1977, pl. 6 (fig. 1)), and both from a
stratigraphic and morphological point of view is most likely to have provided
the ancestor to Forbesiceras. The close similarity between Forbesiceras and
Stoliczkaia of the dispar—clavigera group suggests that the monogeneric sub-
family Forbesiceratinae is superfluous, and that Forbesiceras should be trans-
ferred to the Lyelliceratidae. A study of the early ontogenetic stages of
Forbesiceras has led Casey (1965) to suggest that Neopulchellia (Collignon 1929)
was based upon pyritic nuclei of Forbesiceras.
Genus Stoliczkaia Neumayr, 1875
Type species Ammonites dispar d’Orbigny, 1841
Discussion
Stoliczkaia occupies a key position in the evolution of the mid-Cretaceous
Acanthocerataceae, as it appears to be the origin of both the Mantelliceratinae
and Acanthoceratinae which in turn gave rise to the remaining Upper Cretaceous
acanthoceratids.
More than a score of specific names have been applied to the genus, but
there has been no sound account of intraspecific variation, nor of the apparent
dimorphism present, some subgenera and species reaching a large size and
becoming feebly ribbed at maturity (e.g. S. (S.) dispar), others remaining small
with strong ribs throughout (S. (Shumarinaia)). Furthermore, because of their
transitional position between Lyelliceratidae, Acanthoceratinae and Mantelli-
ceratinae, there are a number of forms whose position is equivocal. Indeed, the
authors find themselves in disagreement over the precise position of some of
these passage forms.
The subgenera of Stoliczkaia are as follows:
1. Stoliczkaia (Stoliczkaia) (type species Ammonites dispar d’Orbigny). Typically
rather involute, compressed ammonites with straight or slightly curved primary
ribs, with shorter intercalated ribs between. Primary ribs may be weakly bullate,
and during early growth stages bear ventrolateral tubercles or clavi, whilst the
venter may be flat or slightly raised. In middle growth tuberculation disappears
and ribs extend across a rounded venter; at maturity ribs broaden, become
irregular, and may become effaced on the adult body chamber. Typical repre-
sentatives of the type species S. (S.) dispar are shown as Figures 40 and 41.
2. Stoliczkaia (Faraudiella) (type species Ammonites blancheti Pictet &
Campiche). Small Stoliczkaia in which distinct siphonal, and sometimes ventro-
244 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 40. Stoliczkaia (Stoliczkaia) dispar (d’Orbigny). The holotype, Renaux collection,
Faculté des Sciences, Montpellier, from the Upper Albian of Ventoux, Vauceuse, France.
Slightly reduced.
lateral tubercles persist on to the body chamber. Typical representatives are
shown in Figure 42.
3. Stoliczkaia (Shumarinaia) (type species S. (Shumarinaia) hashimotoi
Matsumoto & Inoma). Small, with simple suture line and coarse ribbing
throughout.
A fourth subgenus, Villoutreysia was proposed by Casey (1965: 435,
fig. 161; type species S. (V.) villoutreysi Casey) (Fig. 43) for what he described
as Hypacanthoplites homoeomorphs diagnosed as ‘Stoliczkaia with broad,
square venter and strong persistent ribbing, differing from Mantelliceras in much
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 245
Fig. 41. Stoliczkaia (Stoliczkaia) dispar (d’Orbigny). Juvenile specimen in C. W. Wright
collection WW 72344, from the dispar Zone Ammonite Bed of the Dorset Coast. x1.
A B c D
Fig. 42. Stoliczkaia (Faraudiella) sexangulata (Seeley). Seeley’s original specimen, Sedgwick
Museum, Cambridge, B53, from the Cambridge Greensand, Cambridge. 1.
246 ANNALS OF THE SOUTH AFRICAN MUSEUM
earlier loss of ventral tubercles, squarer whorls and, generally, narrow and
shallower umbilicus’. The holotype of S. (V.) villoutreysi is illustrated here as
Figure 43; after an examination of the large collections of Stoliczkaia in the
Paris Museums the authors have concluded that it is not separable subgenerically
from Stoliczkaia sensu stricto.
The genus Paradolphia Casey was proposed (Casey 1965: 461, pl. 77
(figs 5—6)) for the type species P. prisca Casey (illustrated here as Fig. 44) for
intermediates between Stoliczkaia and Forbesiceras but, as noted above, is
possibly a subgenus of Neophlycticeras.
Paracalycoceras (type species Ammonites wiestii Sharpe 1857: 47, pl. 21
(fig. 3)) is an enigmatic genus from the Lower Cenomanian of southern England,
known with certainty only from two specimens—the lost holotype (fide Kennedy
1971) and an extant specimen in the collections of C. W. and E. V. Wright
(Fig. 45). Kennedy (1971: 79) diagnosed the genus as follows: ‘Medium-sized,
somewhat involute ammonites. Inner whorls slightly compressed, with long ribs
bearing umbilical bullae, and lower and upper ventrolateral tubercles separated
by 1, 2, or sometimes more shorter ribs. There is a distinctly raised siphonal area,
and an incipient siphonal tubercle on all ribs. Outer whorl with a broad venter,
and broad, distant, flexuous rursiradiate ribs, irregularly long and short.’
Both morphologically and in the observed ontogenetic changes, Paracaly-
A B c
Fig. 43. Stoliczkaia (Stoliczkaia) villoutreysi Casey. Holotype, O. de Villoutreys collection,
Uppermost Albian, Monte Carlo Tunnel, Monte Carlo. x1.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 247
Fig. 44. Neophylicticeras (Paradolphia) prisca (Casey). A—C. Holotype,
Norwich Castle Museum 61.18(1679). D-E. Paratype, Sedgwick Museum,
Cambridge, B93303. Both from the Cambridge Greensand, Cambridge.
A-C x1, D-E x2.
coceras is virtually indistinguishable from certain species of Stoliczkaia, from
which it is obviously descended. Little more can be said until topotype material
is studied, but it may prove more satisfactory to regard it as a subgenus of
Stoliczkaia.
Kennedy (1971: 80) considered that the genus Coftreauites (Collignon
1929), based upon pyritic nuclei, ‘may be wholly or partly a synonym of
Paracalycoceras’. Again, this question cannot be resolved until Paracalycoceras
is better known or mature Cottreauites are described. It seems very likely,
however, that some Coftreauites are juveniles of either Stoliczkaia or Paracaly-
coceras, and that it should be considered a nomen dubium.
248 ANNALS OF THE SOUTH AFRICAN MUSEUM
C D
Fig. 45. Stoliczkaia (Paracalycoceras) wiestii (Sharpe). C. W. Wright collection 3556, Lower
Cenomanian, Cenomanian Limestone, Bed A2, White Cliff, Seaton, Devon. x1.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 249
Subgenus Stoliczkaia (Stoliczkaia) Neumayr, 1875
Renz (1968: 46) recognized four species groups within Stoliczkaia sensu
stricto:
1. The group of S. dispar with ventrolateral tubercles only on the inner whorls
and weak ribbing on the body chamber, comprising S. dispar (d’Orbigny),
S. dorsetensis Spath and S. tenuis Renz.
2. The group of S. africana which retains ventrolateral tubercles on to the body
chamber, with well-developed, broad, falcate ribs, and includes S. africana
(Pervinquiére) and S. flexicostata Breistroffer. Matsumoto & Inoma (1975) have
proposed the subgenus Shumarinaia for this group.
3. The group of S. notha, which lacks ventrolateral tubercles whilst retaining
strong ribbing on to the body chamber, and comprises S. notha (Seeley) and
S. clavigera Neumayr.
4. The group of S. /evis which lacks ventrolateral tubercles and is virtually
without ornament. Only S. /evis Renz, based on a unique holotype, is assigned
to this group.
Stoliczkaia (Stoliczkaia) tenuis Renz, 1968
Figs 46-53, 54A-F, 55, 68E
Stoliczkaia tenuis Renz 1968: 48, pl. 6 (figs 6, 12), fig. 16b, f.
Material
37 specimens, USNMNH 236981-237012a—b, 237014-5, together with
3 specimens in the collections of the South African Museum, SAM-—PCA5477-78
and 6811, all retaining part or all of their recrystallized shell, from Porto
Amboim, and 7 specimens, SAM-—PCA2938-39, 2944, 3169, 3373 and 5475-5476,
preserved as composite internal moulds from Egito.
Description
Up to 20 mm diameter: shell compressed (“/H = 0,50-0,63), very involute
with a deep, narrow umbilicus (11-17% of diameter). Umbilical walls steep,
almost vertical, with evenly rounded umbilical shoulders. Flanks broad,
slightly convex to flat, with maximum width below mid-flank, converging
towards the narrow venter. Venter slightly convex to almost tabulate, and
weakly raised along the siphonal line. Ornament comprises thirteen to sixteen
prorsiradiate ribs per half-whorl, generally alternating long and short. The ribs
are more or less strongly flexed and bear distinct ventrolateral tubercles. The
latter are joined across the venter by convex ribs.
Up to a diameter of 12 mm in USNMNH 237005 (10 mm in USNMNH
237012) ribs appear to be absent, although there are weak tubercles possibly
marking their position along the ventrolateral shoulders. In USNMNH 237010,
the main ribs are ornamented by rather distinct umbilical bullae.
250 ANNALS OF THE SOUTH AFRICAN MUSEUM
21-40 mm diameter: shell compressed (W/H = 0,53-0,68), very involute
(umbilicus 15-17% of diameter), with a high rectangular whorl section. Ribbing
denser (twenty to twenty-four ribs per half-whorl), not infrequently with two
intercalatories between main ribs. The ventrolateral tubercles are generally still
prominent at this stage.
Fig. 46. Stoliczkaia (Stoliczkaia) tenuis Renz. Reconstructed juvenile and middle growth
stages. Xl.
41-70 mm diameter: the ventrolateral tubercles are commonly lost between
40-45 mm diameter (37 mm in USNMNH 236996) and the ribs pass uninter-
ruped across the venter, sometimes with a slight thickening in the ventrolateral
position. The shell becomes slightly more inflated (W/~ = 0,58-0,69), with
distinctly convex flanks and a rounded venter. The flank ribs coarsen consider-
ably and there are commonly one or two intercalatories between long ribs,
although in USNMNH 236994 and USNMNH 236987 there are probably more
long ribs than intercalatories. Where there are two intercalatories between long
ribs they may be of markedly different lengths (USNMNH 236984). Immediately
prior to the aperture, all ornament is lost and the body chamber becomes smooth
(USNMNH 236981).
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 251
V Ww X
Fig. 47. Stoliczkaia (Stoliczkaia) tenuis Renz. A-C. USNMNH 236999. D-F. USNMNH
237009. G-I. USNMNH 237006. J-—L. USNMNH 237010. M-O. USNMNH 237015.
P-R. USNMNH 237014. S—U. 236997. V-X. USNMNH 237003. x1.
252 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 48. Stoliczkaia (Stoliczkaia) tenuis Renz. A-C. USNMNH 236988. D-F. USNMNH
237003. G-I. USNMNH 236990. x1.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 253
Fig. 49. Stoliczkaia (Stoliczkaia) tenuis Renz. A-C. USNMNH 236945. D-F. USNMNH
237004. G-I. USNMNH 237000. J-—L. USNMNH 237001. M-N. USNMNH 237007. x1.
ANNALS OF THE SOUTH AFRICAN MUSEUM
254
Fig. 50. Stoliczkaia (Stoliczkaia) tenuis Renz. A-C. USNMNH 236985. D-F. USNMNH
236984. x1.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL
Measurements
No.
USNMNH
USNMNH
USNMNH
USNMNH
USNMNH
USNMNH
USNMNH
USNMNH
USNMNH
USNMNH
USNMNH
USNMNH
USNMNH
USNMNH
USNMNH
USNMNH
USNMNH
USNMNH
USNMNH
USNMNH
USNMNH
USNMNH
USNMNH
USNMNH
Intraspecific variation
236983
236985
236986
236987
236988
236989
236990
236992
236994
236995
236996
236997
236998
236999
237000
237001
237002
237003
237004
237005
237006
237008
237010
237012
38
H
32(48)
36(55)
29(52)
35(52)
33(48)
26(52)
27,5(50)
13,5(52)
+19(50)
26(52)
30(54)
17,5(47)
18(54)
19(56)
19(56)
14(54)
15(50)
15,7(52)
10,8(53)
+16,5(55)
17(54)
15(54)
9,5(54)
9(52)
9(51)
6(49)
11(52)
7(47)
8(47)
4(42)
Ww
+22(33)
26(40)
17,6(31)
+23,5(35)
?
18(36)
+19(34)
7,2(28)
+10(26)
15,2(30)
18,7(34)
+12(32)
9(26)
+10(29)
11(32)
7,5(29)
9(30)
8,3(28)
5,5(27)
8,5(28)
10(32)
8(29)
6(34)
+4,5(26)
5(28)
3,4(28)
6(29)
4(27)
4,5(26)
3(32)
W/y
0,69
0,72
0,61
0,67
9
0,69
0,69
0,53
0,53
0,58
0,62
0,68
0,50
0,52
0,58
0,54
0,60
0,53
0,51
0,51
0,59
0,53
0,63
0,50
0,56
0,57
0,54
0,57
0,56
0,75
255
U
10(15)
+13(20)
+9(16)
+9,7(14)
11,1(16)
+9(18)
+10(18)
?
7,7(15)
7,9(14)
?
6(17)
5,5(16)
5(15)
?
2
4,7(16)
2,5(12)
;
+4,5(14)
4(14)
2,7(15)
+3(17)
2(11)
?
3(14)
2,2(15)
2,5(15)
+1,5(16)
The large number of well-preserved specimens available to the authors
permits a better understanding of the intraspecific and ontogenetic variation
in this species. The marked ontogenetic change in ornament shown by this
species has been outlined above. However, Figure 55 also shows that there is
considerable variation in the degree of inflation of the whorls (W/H = 0,50-0,72),
with a distinct tendency for the whorls to become more inflated at large
diameters. Futhermore, Figure 55 shows that not only is there some variation
in the width of the umbilicus, but there is also a slight tendency for the shell to
become more evolute with growth.
256 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 51. Stoliczkaia (Stoliczkaia) tenuis Renz. A-D. USNMNH 236994. x1.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 257
Discussion
As shown above, the present material is rather variable, and the type of
S. (S.) tenuis and the specimen referred to as S. (S.) aff. tenuis by Renz (1968,
pl. 6 (fig. 12)) fall within this range.
When mature, S. tenuis closely resembles S. (S.) clavigera Neumayr
(= Ammonites dispar Stoliczka (non d’Orbigny) 1865: 45, pl. 85 (figs 1-3 only))
(see Fig. 56), from which it appears to differ only in being consistently more
compressed. Further work may show that the two merit only subspecific
separation.
S. (Stoliczkaia) dispar (d’Orbigny) (1841: 143, pl. 45 (figs 1-2)) is a widely-
cited but much misinterpreted species. Consequently, the holotype is here
photographically figured for the first time (Fig. 40), as well as a typical specimen
from the dispar Zone of Dorsetshire, England (Fig. 41). S. (Stoliczkaia) dispar
differs from the present species in having far more (up to nine) intercalatories
between long ribs, whilst the latter are ornamented with weak umbilical bullae.
The venter loses its ventrolateral tubercles and becomes rounded at smaller
diameters and the body chamber ornament is also different; in S. (S.) dispar,
ribbing rapidly weakens in maturity (at least on the internal mould) and all that
Fig. 52. Stoliczkaia (Stoliczkaia) tenuis Renz. A-C. USNMNH 236987. x1.
258 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 53. Stoliczkaia (Stoliczkaia) tenuis Renz. A-B. Lateral and ventral views of SAM-—
PCA3169. C—D. Ventral and lateral views of SAM—PCA5475. E. Lateral view of SAM-—
PCAS5476. F. Lateral view of SAM-PCA2939. G-H. Lateral and ventral views of SAM-—
PCA2938. I. Lateral view of SAM-—PCA3373. A-B, I x0,75, C-H x1.
259
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL
A-B. Lateral and ventral views of
SAM-PCA6811. C—D. Ventral and lateral views of SAM-PCA2944. E. Lateral view of
SAM-PCAS5477. F. Ventral view of SAM-—PCA5478. G-H. Mortoniceras (Angolaites)
simplex (Choffat). Ventral and lateral views of SAM—PCA3107. A-B, E-H x1, C-D 0,75.
Fig. 54. A-F. Stoliczkaia (Stoliczkaia) tenuis Renz.
260 ANNALS OF THE SOUTH AFRICAN MUSEUM
!
" OTH EIGHT —______._g
a a oes ™
UMBILICUS (%) —___g
0 70
DIAMETER (am) ==
Fig. 55. Plots of inflation and umbilical ratio against diameter for S. tenuis. Dots = present
material; squares = holotype and paratype (after Renz 1968).
remains are the weak umbilical bulges of the long ribs. Furthermore, the upper
half of the flanks are concave, forming a broad, spiral depression.
S. (Stoliczkaia) notha (Seeley) (1865: 232; Spath 1929: 335, fig. 110)
(Fig. 57) differs from S. tenuis in having a rounded venter at all growth stages.
Stoliczkaia dorsetensis Spath (1929: 337, pl. 31 (fig. 2), pl. 33 (fig. 1)) is more
inflated and more densely ribbed than the Angolan material, as well as having
more (three to nine) intercalatories between long ribs on the inner whorls.
S. (Stoliczkaia) villoutreysi Casey (Fig. 43) has more intercalatories between
long ribs (up to seven), whilst the venter is tabulate in maturity, when it
homoeomorphs Hypacanthoplites.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 261
S. (Stoliczkaia) argonautiformis (Stoliczka) (1864: 87, pl. 46 (figs 1-2)) has
strongly converging flanks, subdued ribbing and a more narrowly arched venter
than the present species. Stoliczkaia tetragona Neumayr (nom. nov. pro
Ammonites dispar Stoliczka (non d’Orbigny), 1864: 85, pl. 45 (fig. 2)) (Fig. 58)
differs from the Angolan material in being much more inflated, although the
style of ornament is similar in adults.
Stoliczkaia grandidieri Boule, Lemoine & Thévenin (1907: 34, pl. 8 (fig. 8)),
S. gardonica (Herbert & Meunier-Chalmas) (1875: 116, pl. 4 (figs 1-2));
S. rhamnonota (Seeley) (1865: 233, pl. 11 (fig. 7); Spath 1929: 333, fig. 109)) and
S. blancheti (Pictet & Campiche) (1859: 188, pl. 23 (figs 2, 6); Renz 1968: 46,
pl. 5 (fig. 21)) are all referable to the subgenus Faraudiella, whilst S. africana
Pervinquiére (1907: 388, pl. 12 (fig. 10)), S. flexicostata Breistroffer (nom. nov.
pro A. dispar Pictet & Campiche (non d’Orbigny) 1860: 264, pl. 38 (fig. 4);
Renz 1968: 49, pl. 6 (fig. 9)), S. hashimotoi Matsumoto & Inoma, and S. asiatica
Matsumoto & Inoma (1975) are all Shumarinaia.
As suggested by Matsumoto & Inoma (1975), Stoliczkaia uddeni Bése
(1927: 211, pl. 4 (figs 12-15)), is probably a synonym of S. texana Cragin
(1893: 235, pl. 44 (fig. 1)) which itself is a close relative of S. crotaloides
(Stoliczka) (1864: 88, pl. 46 (fig. 3)) (Fig. 59). The latter species differs from the
Angolan material in maintaining single prorsiradiate ribs to the peristome, with
only rare intercalatories, and in the flattened venter of the inner whorls; they
may represent a distinct Cenomanian offshoot.
‘Stoliczkaia’ razafimbeloi Collignon (1968: 29, pl. 6 (fig. 7), pl. 7 (fig. 4))
and ‘S.’ vendegiesi Collignon (1968: 31, pl. 7 (fig. 5)) both differ from the
present material in the development of distinct upper and lower ventrolateral
tubercles on the body chamber and are thus transitional to Graysonites. They
do not appear to be referable to the genus Stoliczkaia.
Stoliczkaia patagonica Stoyanow (1949: 128, pl. 26 (figs 3-4)), S. excentrum-
bilicata Stoyanow (1949: 129, pl. 26 (figs 5-6)) and S. scotti Stoyanow (1949:
129, pl. 26 (figs 7-8)) are all from the same stratigraphic level and locality and it
is doubtful whether more than one species is represented. In this material,
flexuous main ribs are ornamented with umbilical bullae and separated by two
to four intercalatories. The ribs pass strongly across the venter and appear to
lack ventrolateral tubercles. These ‘species’ are very close to S. dorsetensis
Spath.
Stoliczkaia adkinsi Bose (1927: 193, pl. 18 (figs 9-17)) differs from the
present material in having more strongly differentiated long and short ribs, the
former with distinct umbilical tubercles.
‘Submantelliceras’ worthense (Adkins) (1920: 93, pl. 1 (figs 11—13)) from the
Pawpaw Formation of Tarrant County, Texas, may be based upon juveniles of
Stoliczkaia. It differs from the present material in the possession of umbilical
tubercles and in the (?) earlier loss of ventrolateral tubercles.
Some of the Mantelliceras (Submantelliceras) saxbii (Sharpe) figured by
Thomel (1972: 16-17, pl. 1 (figs 8-12 only)) are clearly based upon Stoliczkaia
262 ANNALS OF THE SOUTH AFRICAN MUSEUM
A B
Fig. 56. Stoliczkaia (Stoliczkaia) clavigera Neumayr. Copy of Stoliczka 1864, pl. 45 (fig. 1-1a).
xi
of the dispar—dorsetensis group and differ from the Angolan material in having
more intercalatories between long ribs.
Stoliczkaia praecursor Anderson (1958: 246, pl. 12 (fig. 1)) is too poorly
figured and described for proper comment, but appears to differ from the
present species in being more inflated, more coarsely ribbed, and in the presence
of umbilical bullae at large diameters. It seems to be allied to S. tetragona
Neumayr.
In his original account of this species, Renz compared S. tenuis with
Mantelliceras martimpreyi (Coquand), and, indeed, as demonstrated by the
present population S. (S.) tenuis confirms that some Stoliczkaia have sub-
mantellicerine nuclei, as suggested by Kennedy (1971). In particular, the authors
are impressed by their close similarity to pyritic nuclei such as those figured by
Collignon (1929, pl. 3 (figs 4-5)) from Diego-Suarez, Madagascar, as Acantho-
ceras (Mantelliceras) martimpreyi Coquand. The latter differ only in having
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 263
C D
Fig. 57. Stoliczkaia (Stoliczkaia) notha (Seeley). A-B. Holotype, Sedgwick Museum,
Cambridge, B40. C-D. BMNH C4811, type of the variety u/tima Spath. Both from the
Cambridge Greensand, Cambridge. 1.
264 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 58. Stoliczkaia tetragona Neumayr. Copy of Stoliczkaia 1864, pl. 45 (fig. 2-2a). x1.
distinctly differentiated lower ventrolateral tubercles. However, there can be
little doubt that records of Mantelliceras (Thomel 1972) and Submantelliceras
(Adkins 1920) from the Upper Albian are based upon juveniles of Stoliczkaia.
Kennedy & Hancock (1971) have shown Submantelliceras martimpreyi
(Coquand) to be a junior subjective synonym of M. saxbii (Sharpe), from which
the authors’ material differs in being more compressed, lacking distinct lower
ventrolateral tubercles and in showing a marked, and characteristic, change of
ornament on the body chamber. However, the fact that Thomel (1972) assigned
late Albian species of Stoliczkaia to M. saxbii merely serves to emphasize the
close relationship between these two genera and suggests that the origin of
(at least) compressed Mantelliceras and Utaturiceras lies close to Stoliczkaia of
tenuis type. The writers would also point to the close similarity of compressed
variants to juvenile Forbesiceras (see Juignet & Kennedy 1977), generally
described as Neopulchellia (a subjective synonym), and evidence for the descent
of Forbesiceras from the S. (S.) tenuis group is to be published elsewhere.
Occurrence
S. (Stoliczkaia) tenuis is so far known only from the Upper Albian of
Switzerland and Angola.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 265
Fig. 59. Stoliczkaia crotaloides (Stoliczka). Copy of Stoliczka 1864, pl. 46 (fig. 3-3a).
xT
Family Brancoceratidae Spath, 1934
Subfamily Brancoceratinae Spath, 1934
Genus Hysteroceras Hyatt, 1900
Type species Ammonites varicosus J. de C. Sowerby, 1824
Discussion
Hysteroceras is typically a low Upper Albian micromorph genus, clearly
descended from the earlier Brancoceras, and has generally been regarded as
characteristic of, and largely confined to, the Mortoniceras inflatum Zone.
However, micromorph brancoceratids persist into the highest levels of Albian
where they have generally been referred to as Spathiceras or Mortoniceras
(Cantabrigites).
Whitehouse (1927: 110) introduced Spathiceras without formal diagnosis,
merely naming as type of the genus Hystrichoceras antipodeum Etheridge (1902:
47, pl. 7 (figs 6—-7)). The holotype of S. antipodeum is from the Upper Albian of
Point Charles, near Darwin, Northern Australia, where it occurs associated with
Desmoceras latidorsatum (Michelin) (= D. carolensis Etheridge 1902, pl. 7
(figs 2-5)), Idiohamites cf. spinulosus (J. Sowerby) (= Ancyloceras (?) sp. ind.,
Etheridge 1902, pl. 7 (figs 14-15)), Scaphites eruciformis Etheridge (very close to
S. simplex Jukes-Browne), Hamites cf. virgulatus (Brongniart) (= Hamites (?)
sp. ind., Etheridge 1902, pl. 7 (figs 12-13)), Aucellina gryphaeoides (J. de C.
Sowerby) (= A. incurva Etheridge), together with the genera Beudanticeras,
Labeceras, Myloceras, Anisoceras and Ptychoceras (Whitehouse 1928). Because
266 ANNALS OF THE SOUTH AFRICAN MUSEUM
V W X Y z
Fig. 60. Hysteroceras? spp. juv. A-B. USNMNH 237018. C-E. USNMNH 237017.
F. Hysteroceras? cf. ootatoorense (Stoliczka), USNMNN 237016. G—H, K-Z. Aysteroceras
antipodeum (Etheridge), a series of limonitic specimens from northern Australia. G—H.
BMNH C26539. K-L. BMNH C26551. M-N. BMNH C26523. O-Q. BMNH (C26552.
R-T. BMNH C26548. U-V. BMNH C26518. W-X. BMNH C26546. Y-Z. BMNH C35269.
I_J. Hysteroceras ? ootatoorense (Stoliczka). Copy of Stoliczka 1865, pl. 32 (fig. 2). All x1.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 267
Whitehouse (1928: 279) considered the fauna to be ‘. . . typical of the substuderi
Zone of the Upper Albian’, the name Spathiceras has been applied to all
flat-sided, weakly tuberculate micromorph mortoniceratinids of uppermost
Albian age. However, the genera Labeceras, Myloceras and Beudanticeras are
nowhere known to range above the Mortoniceras inflatum Zone, and the
faunas recorded by Etheridge (1902) and Whitehouse (1927) seem to be typical
low Upper Albian (approximately varicosum Subzone) assemblages. It is,
perhaps, not coincidental, therefore, that S. antipodeum (Fig. 60K-—Z) bears a
remarkable resemblance to Hysteroceras of the binum-subbinum group, from
which it differs only in being consistently more evolute (umbilicus 40-45 % of the
diameter). The differences are not sufficient for generic separation, and
Spathiceras is considered a junior subjective synonym of Hysteroceras. There is
no good evidence for Cenomanian occurrences of the genus.
We are left, therefore, only with Hysteroceras for those uppermost Albian
mortoniceratinid micromorphs with an evolute shell, and undivided ventro-
lateral tubercles. Either these represent a continuation of the Hysteroceras stock,
or they represent an unnamed homoeomorphic development. Until better and
larger collections are known, the authors prefer to follow Renz (1968) and refer
their material to Hysteroceras.
Hysteroceras ? cf. ootaturense (Stoliczka, 1865)
Fig. 60E-F, I-J
Compare
Ammonites ootaturensis Stoliczka, 1865: 56, pl. 32 (fig. 2).
Mortoniceras ootaturense (Stoliczka) Pervinquiére, 1910: 64, pl. 6 (figs 2-5).
Spathiceras ootaturense (Stoliczka) Spath, 1934: 444, 445, fig. 160h. Breistroffer,
1940: 75.
? Pervinquieria (2) sp. nov. Breistroffer, 1940: 75.
Material
A single specimen, USNMNH 237016, with recrystallized shell preserved,
although somewhat corroded, from Porto Amboim.
Description
Shell small, evolute, with a wide, shallow umbilicus, steep umbilical walls
and evenly rounded umbilical shoulder. The whorl section is subquadrate,
compressed with flattened flanks. Ribbing is very weak on the inner flank, but
strengthens markedly over the ventrolateral shoulder, whilst there is a prominent
keel.
Discussion
Stoliczka’s (1865) type (Fig. 60I—J) comes from a locality ‘near Odium’,
from where he also records other typical uppermost Albian species such as
M. (Mariella) bergeri (Brongniart), M. (M.) circumtaeniatus (Kossmat),
268 ANNALS OF THE SOUTH AFRICAN MUSEUM
Anisoceras perarmatum Pictet & Campiche, D. (Desmoceras) latidorsatum
(Michelin), and Lechites gaudini (Pictet). Ammonites ootaturense is probably,
therefore, of latest Albian age, although Stoliczka’s record of Turrilites costatus
Lamarck and Neoptychites xetra (Stoliczka) from the same locality suggests the
presence of beds as high as the Lower Turonian in the vicinity.
Hysteroceras antipodeum (Etheridge) (Fig. 60G-—H, K-—Z) differs from the
Angolan specimen in being somewhat older, with a wider umbilicus, flatter
flanks and more prominent umbilical tubercles. The specimen of Schloenbachia
rostratus var. antipodeus (Etheridge) figured by Etheridge (1909, pl. 67 (figs 3—-4))
(non Etheridge 1902) was referred to Dipoloceras bouchardianum (d’Orbigny) by
Stieler (1920), but was renamed Prohysteroceras richardsi var. nitidum by
Whitehouse (1926). However, Etheridge’s specimen appears indistinguishable
from Dipoloceras quadratum Spath (1921: 278, pl. 25 (fig. 3)).
Hysteroceras ? nanum Renz (1968: 63, pl. 11 (fig. 4), fig. 22c—d) differs from
H? ootaturense in having prominent umbilical tubercles at an early stage. The
specimen of H. semileve Haas recorded by Renz (1968: 63, pl. 11 (fig. 6),
fig. 22g-h) from the ‘Unteren Vraconnien’ appears to be based upon a larger
fragment of his H.? nanum.
Hysteroceras ? tunisiense (Spath) (nom. nov. pro Mortoniceras inflatum var.
orientalis (?) Pervinquiére (non Kossmat) 1907: 229, pl. 11 (fig. 2)) differs from
H ? ootaturense and the present specimen in having sharp, flexuous prorsiradiate
ribs which bifurcate from distinct umbilical bullae at only 17 mm diameter.
Hysteroceras ? wenoense (Adkins) (1928: 229, pl. 20 (fig. 13)) differs from
the authors’ material in its sharp ribbing and distinct umbilical tubercles. The
specimen figured by Renz (1968: 62, pl. 11 (fig. 5)) as Hysteroceras cf. subbinum
Spath may possibly belong here.
‘Algericeras’ boghariense (Coquand) (Pervinquiere 1907: 240, pl. 11 (fig. 16))
differs from the present specimen in having a quadrate whorl section (W/H = 1,00)
with dense, fine, straight ribs, about thirty-two per half whorl, which arise in
pairs from umbilical bullae and terminate in ventrolateral tubercles. It is said
to be of Cenomanian age but appears merely to be based upon pyritic nuclei of
Mortoniceras.
Occurrence
Hysteroceras ? ootaturense is known from the Upper (? uppermost) Albian
of southern India, and possibly the uppermost Albian of France, Algeria and
Angola.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 269
Subfamily Mortoniceratinae Spath, 1925
Genus Mortoniceras Meek, 1876
Subgenus Durnovarites Spath, 1932
Type species Subschloenbachia perinflata Spath, 1921
Discussion
Wiedmann & Dieni (1968: 142) have divided the subgenus Durnovarites
into two species groups:
1. The group of M. (D.) subquadratum, characterized by four rows of tubercles
on the ribs. To this group may be assigned M. (D.) subquadratum Spath,
M. (D.) quadratum Spath, M. (D.) perinflatum (Spath), M. (D.) postinflatum
Spath, M. (D.) depressum (Spath), M. (D.) adkinsi (Young), M. (D.) vraconense
Renz, M. (D.) spinosum (van Hoepen non Pervinquiére), M. (D.) subnanum
(Breistroffer), M. (D.) ishiguaense Reyment, M. (D.) levecostatum Reyment,
M. (?D.) crassicornutum (Reyment), M. (D.) lowrii McLearn, M. (D.) downii
McLearn, M. (D.) rerati Collignon and M. (D.) haueri Collignon.
2. The group of M. (D.) spinosum, with only three rows of tubercles and sub-
ordinate ribbing (appears merely to be based upon juveniles which have still to
develop the fourth row of tubercles). To this group belong M. (D.) spinosum
(Pervinquiére), M. (D.) kentronotum Spath, M. (D.) lemoinei (Spath), M. (D.)
neokentroides Wiedmann & Dieni and M. (D.) aubersonense Renz.
Mortoniceras (Durnovarites) perinflatum (Spath, 1922)
Figs 3G, 61, 62D-I, 63-64
Ammonites inflatus Pictet & Campiche (non J. Sowerby), 1860: 178, pl. 21 (fig. 5), pl. 22 (fig. 3).
Inflaticeras (‘Subschloenbachia’) perinflatum Spath, 1922: 113.
? Inflaticeras (Subschloenbachia) depressum Spath, 1922: 114, figs B, 2a—d.
Inflaticeras (Subschloenbachia) quadratum Spath, 1922: 115.
Pervinquieria quadrata (Spath) Spath, 19265: 423.
? Pervinquieria depressa (Spath) Spath, 1928: 51.
Mortoniceras (Durnovarites) perinflatum (Spath) Spath, 1933: 430, pl. 40 (fig. 2), fig. 150.
Wiedmann & Dieni, 1968: 143, pl. 14 (figs 3-4), fig. 92. Renz, 1968: 51, pl. 8 (figs 3, 5, 8),
pl. 9 (figs 1-2), figs 17a, 18c, 19c, f. Marcinowski & Naidin, 1976: 109, pl. 6 (figs 1-2).
Mortoniceras (Durnovarites) quadratum (Spath) Spath, 1933: 432, pl. 45 (fig. 3), pl. 46 (fig. 6),
pl. 49 (fig. 12). Reyment, 1955: 38, pl. 6 (figs 4-5), pl. 7 (fig. 3), fig. 15. Wiedmann &
Dieni, 1968: 145, fig. 93.
Mortoniceras (Durnovarites) postinflatum Spath, 1933: 433, pl. 40 (figs 3-5), pl. 46 (figs 3, 7),
pl. 47 (fig. 6). Renz, 1968: 53, pl. 8 (figs 1-2, 6), figs 17b, d, 18b, 19a—b, d. Marcinowski &
Naidin, 1976: 109, pl. 7 (fig. 1), pl. 8 (fig. 1).
? Mortoniceras (Durnovarites) depressum (Spath) Collignon (in Besairie), 1936: 195. Reyment,
1955: 37, pl. 7 (fig. 4), fig. 14.
Durnovarites adkinsi Young, 1957: 6, pl. 1 (figs 3, 6).
Mortoniceras (Durnovarites) vraconense Renz, 1968 : 54, pl. 7 (figs 6-7, 11), fig. 19e. Marcinowski
& Naidin, 1976: 111, pl. 9 (fig. 1).
Material
Two specimens, USNMNH 237021-22, both with recrystallized shell
preserved from Porto Amboim, together with three specimens, SAM—PCA4802,
4576 and 4587, from Cabo Ledo.
270 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 61. Mortoniceras (Durnovarites) perinflatum (Spath). The holotype, Pictet collection,
Natural History Museum, Geneva. From the Upper Albian of Vraconne, Switzerland. x1.
Description
The shell is moderately inflated and rather evolute (umbilicus 32-36% of
the diameter), with a wide, shallow umbilicus and steep umbilical walls. The
whorl section is almost quadrate (W/H = 0,93-1,05), with subparallel flanks.
The umbilical shoulder is evenly rounded intercostally. Ribs begin at the
umbilical seam and are rectiradiate to the umbilical shoulder where they termi-
nate in weak bullae. The latter give rise to 1-2 prorsiradiate flank ribs, 38-43 per
whorl and broader than the interspaces. All ribs are ornamented by a midlateral
tubercle and closely spaced upper and lower ventrolateral tubercles. On the
venter, the ribs pass forwards, finally becoming effaced in the sulci bordering the
siphonal keel. The ribs show spiral ornament which is especially prominent on
the ventrolateral tubercles.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 271
Fig. 62. A-C. Mortoniceras (Durnovarites) subquadratum Spath, USNMNH_ 237023.
D-I. Mortoniceras (Durnovarites) perinflatum (Spath). D-F. USNMNH 237021. G-I.
USNMNH 237022. x1.
272 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 63. Mortoniceras (Durnovarites) perinflatum (Spath). Ventral and lateral views of
SAM-PCA4802. x1.
Measurements ;
No. D H WwW W/a U
USNMNH 237021 ao 22,5(42) ? 2 17(32)
ue 44 +19(43) +20(45) 1,05 ?
USNMNH 237022 35 +16(46) +15(43) 0,93 12,5(36)
Discussion
Renz (1968) showed M. (D.) quadratum to be based upon juveniles of
M. (D.) perinflatum, whilst M. (D.) vraconense appears to comprise hyponodose
adults which the authors do not consider to bear specific separation from
M. (D.) perinflatum.
Similarly, M. (D.) postinflatum differs from the strictly contemporaneous
M. (D.) perinflatum only by its more prominent ventrolateral tubercles and more
inflated whorls (W/H = 1,20—-1,65). The authors do not regard the differences as
sufficient for specific separation and place M. (D.) postinflatum within the
synonomy of M. (D.) perinflatum, although the name might usefully be retained
at the varietal level.
Durnovarites adkinsi Young was separated from M. (D.) perinflatum on the
basis of its slightly rounder ribs, less tumid flanks, and denser-ribbed inner
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 273
Fig. 64. Mortoniceras (Durnovarites) perinflatum
(Spath). The holotype of Durnovarites spinosus
van Hoepen, SAM—D3154, from the Upper Albian
of the Mzinene Formation, Zululand. x1.
whorls. The differences appear to be those between individuals, and the Texas
species is considered to be referable to M. (D.) perinflatum.
Mortoniceras (Durnovarites) depressum (Spath) was said to differ from
M. (D.) quadratum in its far more depressed whorl section, with rounded instead
of parallel flanks. It closely approaches M. (D.) postinflatum and may, therefore,
be no more than an extreme variant of M. (D.) perinflatum.
Occurrence
Mortoniceras (Durnovarites) perinflatum is known from the uppermost
Albian of Switzerland, Poland, southern England, Sardinia, Texas, Nigeria and
Angola. The authors have also seen comparable material from Zululand.
Mortoniceras (Durnovarites) subquadratum Spath, 1933
Figs 62A-C, 65C-D
? Subschloenbachia meunieri Spath, 1922: 115.
? Mortoniceras (Durnovarites) meunieri (Spath), Spath 1932: 399.
Mortoniceras (Durnovarites) subquadratum Spath, 1933: 435, pl. 42 (fig. 5), pl. 43 (fig. 1),
pl. 44 (fig. 6), pl. 45 (fig. 5), pl. 47 (figs 2-4), pl. 48 (fig. 2). Reyment, 1955: 38. Dieni &
Massari, 1963: 798. Wiedmann & Dieni, 1968: 142, pl. 13 (fig. 9), fig. 90. Renz, 1968: 55,
pl. 7 (figs 8, 10), pl. 10 (figs 1-4, 7-8), fig. 17f,-f.. Marcinowski & Naidin, 1976: 110,
pl. 6 (fig. 3).
Mortoniceras (Durnovarites) subquadratum var. tumida Spath, 1933: 435, pl. 48 (fig. 4).
Wiedmann & Dieni, 1968: 143, pl. 13 (fig. 10), fig. 91.
Mortoniceras (Durnovarites) subquadratum var. crassicostata Spath, 1933: 432, pl. 42 (fig. 9).
? Mortoniceras (Pervinquieria) sp. juv., Spath, 1933: 412, pl. 41 (fig. 7).
? Pervinquieria (Cantabrigites ?) subnana Breistroffer, 1947: 91.
Pervinquieria (Durnovarites) subquadrata (Spath) Breistroffer, 1947: 61.
Durnovarites spinosum van Hoepen, 1951: 324, figs 380-383.
? Mortoniceras (Durnovarites) levecostatum Reyment, 1955: 38, pl. 7 (fig. 2).
? Mortoniceras (Durnovarites) subnanum (Breistroffer) Renz, 1968: 56, pl. 10 (figs 5—6), fig. 17g.
Material
A single specimen, USNMNH 237023, with recrystallized shell preserved
from Porto Amboim, and one, SAM-PCA3235, from Praia-Egito, preserved as
an internal mould.
ANNALS OF THE SOUTH AFRICAN MUSEUM
274
1% 60UttVJd-WVSs yUuA}eAeU E JO” SE EE OF TED —
SMOIA [BIOIL] pue [eIUDA “AOU “ds JUOUsI]]09 (SajtADAOUANG) SD41BIIUOLAOW “J-A “SETEVOI-WYVS JO SMOIA [RIVA Pure [PINE T “yyedg winjoaponbqns
(Sajtapaouing) spsaUojsoBM “G-D “OSTEVIdM-WYS JO SMOIA JerJoyeT pue [eIUSA “QeyoyD) xajdiuis (sajipjosup) svda1uoJAOw “G-VY °S9 “BI
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 275
Description
The shell is very evolute (umbilicus 50° of diameter), compressed, with a
slightly depressed, subquadrate whorl section (W/H = 1,11). The umbilicus is
wide, shallow, with steeply inclined walls and evenly rounded umbilical
shoulders. The flanks are flattened, with maximum width close to the umbilical
shoulder, and converge slightly to the broad venter.
There are 10 prominent, somewhat bullate umbilical tubercles per half-
whorl, from which arise 1-2 rectiradiate to slightly rursiradiate ribs. Where
single there is frequently an adjacent intercalated rib, so that there are 19 ribs
per half-whorl. The ribs are thick, robust, about as wide as the interspaces, and
are ornamented by closely spaced double ventrolateral tubercles. The lower
ventrolateral tubercle is sharp and prominent, whereas the upper ventrolateral
tubercle is more weakly developed and clavate. On the final third of the outer
whorl (which is entirely septate), there is a weakly developed midlateral tubercle.
The well-developed siphonal keel is separated from the upper ventrolateral clavi
by prominent sulci. The ribs on the adoral portion of the outer whorl show
weakly developed spiral ornament.
Measurements
No. D H W W/a U
USNMNH 237023 46 13,5(29) +15(33) 1,11 23(50)
Discussion
Mortoniceras (Durnovarites) meunieri (Spath) (1922: 115; 1932: 399)
(nom. nov. pro Ammonites inflatus Meunier (non J. Sowerby) 1887: 61, pl. 1
(fig. 2)) is an evolute species with about 30 coarse, rectiradiate to slightly
rursiradiate ribs arising singly or in pairs from umbilical tubercles, each
ornamented with a prominent midlateral tubercle and a (?) double ventrolateral
tubercle. Meunier (1888) figured his specimen only in lateral view, without
description, and hence comparison is difficult. If it is, indeed, a M. (Durnovarites),
then not only might it be a synonym of M. (D.) subquadratum, but it also has
priority over that name.
This species differs from M. (D.) perinflatum (Spath) in its much wider
umbilicus (43-50% as against 28-36%) and its typically less depressed whorl
section (W/H = 1,03-1,16). However, adults of M. (D.) subquadratum are
unknown and until population studies are undertaken it is not known whether
the differences are truly of specific importance.
Renz (1968) included Durnovarites spinosum van Hoepen (non Pervinquiére)
in the synonomy of M. (D.) subquadratum, an assignment with which the authors
concur; the type is re-illustrated here as Figure 64.
Mortoniceras (Durnovarites) ishiaguense Reyment (1955: 38, pl. 7 (fig. 1))
is very close to M. (D.) subquadratum but is apparently much more densely
ribbed. Since the Nigerian species is based upon a mature individual, it is not
276 ANNALS OF THE SOUTH AFRICAN MUSEUM
directly comparable with M. (D.) subquadratum at the present time. Mortoniceras
(Durnovarites) levecostatum Reyment (1955: 38, pl. 7 (fig. 2)) is based upon a
somewhat distorted composite internal mould showing about twenty-six coarse,
robust ribs per whorl and with a wide umbilicus: Judging from Reyment’s
(1955) description, it may not bear separation from M. (D.) subquadratum.
Occurrence
Mortoniceras (Durnovarites) subquadratum is known from southern
England, Poland, Switzerland, Sardinia, Zululand, and Angola.
Mortoniceras (Durnovarites) collignoni sp. nov.
Figs 65E-F, 66-67, 68B—D, 69
Material
About a hundred specimens, in the South African Museum, Cape Town,
from Praia-Egito. All are preserved as composite internal moulds.
Type material
SAM-PCA3227 is designated as holotype; paratypes are SAM-—PCA2975,
3189, 3199, 3202, 3217, 3259, 3277, 3294, 3309, 3317 and 3407.
Etymology
For the late General Maurice Collignon who contributed so much to our
knowledge of ammonite systematics, and who helped the authors with their
studies in many ways.
Diagnosis
A densely ribbed species of M. (Durnovarites) characterized by a dramatic
change in shell morphology on the adult body chamber. The phragmocone has
a subrectangular, compressed whorl section and is ornamented with 36-42
rectiradiate to prorsiradiate ribs per whorl which frequently bifurcate from
umbilical bullae and are all ornamented with midlateral and double ventro-
lateral tubercles. On the adult body chamber, however, all tuberculation is
rapidly lost and the simple ribs develop a strong convex-adoral curvature, while
the whorl section now becomes strongly compressed and lanceolate.
Description
Almost all the material has been crushed to varying degrees.
The coiling is evolute (umbilicus about 24-30% of diameter), with a more
or less compressed whorl section from an early growth stage (Fig. 69). Up to the
body chamber, the intercostal whorl section is generally subrectangular,
compressed. On the body chamber, the flanks become strongly convergent and
the whorl section eventually becomes lanceolate. The umbilicus is wide, shallow,
with a steep umbilical wall on the inner whorls, and an evenly rounded umbilical
shoulder. On the outer whorl, the umbilical wall becomes sloping and the
umbilical shoulder is gently rounded.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 277
Fig. 66. Mortoniceras (Durnovarites) collignoni sp. nov. A reconstruction based on the
holotype and paratype material. 0,66.
—eghap = aly mamas @
eal
ANNALS OF THE SOUTH AFRICAN MUSEUM
278
‘TX 3 ‘990 E-W “8LZTEVOd-WYS ‘odAjered &
JO MOIA [BINT “OD SLZZTEVOd-WYVS ‘edAi0[0Y 9Y} JO SMOIA [e.I]UBA PUR [eJaIeT “g—YW “AOU ‘ds 1u0USI//09 (Sa1ADAOUANG ) SDADIIUOJAOJY *“D-Y
“L9 “SIF
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 279
Fig. 68. A. Mortoniceras (Angolaites) simplex (Choffat). Ventral view of SAM-PCA3142.
B-D. Mortoniceras (Durnovarites) collignoni sp. nov. B. Lateral view of a paratype,
SAM-PCA3269. C-D. Lateral and ventral views of a paratype, SAM-PCA3182. E. Stoliczkaia
tenuis Renz. Lateral view of SAM-—PCA3313. A-B x0,66, C-E x1.
280 ANNALS OF THE SOUTH AFRICAN MUSEUM
In the earliest observed growth stages, the ribbing is generally simple,
slightly prorsiradiate, with frequent intercalatories. All ribs are ornamented by
distinct lateral and upper and lower ventrolateral tubercles, while long ribs
arise from fairly prominent bullae. Even at this stage, spiral notching is evident
on the tubercles. In the middle growth stages, the connections between the
intercalated ribs and the umbilical bullae strengthen, and many ribs are seen to
bifurcate from the latter, while the lateral and upper and lower ventrolateral
tubercles become more swollen. At large growth stages, the ribbing again
becomes simple but very subordinate to the now very swollen and prominent
lateral and ventrolateral tubercles. At this stage, the lower lateral tubercle is
somewhat clavate and the upper lateral and ventrolateral tubercles strongly so.
The ventrolateral tubercles are now prominently raised above the narrow,
sunken, keeled venter. On the last portion of the body chamber there is a
great change in ornament. The whorl section changes from subrectangular to
lanceolate, with the disappearance of all tubercles, and the ribs become strongly
convex.
There are generally 36—42 ribs per whorl in the middle growth stages, about
as wide as the interspaces, with somewhat fewer in juveniles and on the outer
whorl.
Discussion
The body chamber ornament of this species is characteristic.
Mortoniceras (Durnovarites) perinflatum (Spath) (Renz 1968: 51, pl. 9
(fig. 1)) differs from M. (D.) collignoni sp. nov. in having a strongly depressed
whorl section in maturity, whilst M. (D.) subquadratum Spath (1933: 435, pl. 37
(fig. 6)) differs from the Angolan species in being more evolute (umbilicus
40-48 % of the diameter) and in apparently lacking the dramatic change in body
chamber ornament shown by M. (D.) collignoni.
Mortoniceras rostratum (J. Sowerby) (Fig. 70) differs from the present
species in having sparser, more distant ribbing, whilst the ribs of the body
chamber retain four rows of tubercles almost to the peristome.
Mortoniceras (Durnovarites) ishiaguense Reyment (1955: 38, pl. 7 (fig. 1))
differs from the present species in being more evolute, with less compressed
whorls, and in apparently lacking the characteristic change in the body ornament
seen in M. (D.) collignoni. Mortoniceras levecostatum Reyment (1955: 38, pl. 7
(fig. 2)) is from the same locality and horizon as M. (S.) ishiaguense but was said
to differ in being more distantly ribbed, with more irregular ornament. The
differences may not be of specific significance.
Howarth (1965) considered Neokentroceras curvicornu crassicornutum
Reyment (1955: 41, pl. 4 (figs 7-8)) a species of Durnovarites, but Reyment
(1955) records this form in association with a typical N. curvicornu Spath, and
hence it is much older than typical M. (Durnovarites).
Mortoniceras (Styphloceras) lowrii McLearn (1972: 72, pl. 30 (figs 1-3),
pl. 39 (figs 3-4)) and M. (S.) downii McLearn (1972: 73, pl. 31 (figs 1-3)) are
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 281
Fig. 69. Mortoniceras (Durnovarites) collignoni sp. nov. Whorl sections. A. SAM—PCA3202.
B. SAM-PCA3309. C. SAM-—PCA3278. D. SAM-—PCA3257. E. SAM-PCA3309. x1.
D
both species of Durnovarites, thus indicating the presence of uppermost Albian
strata at the Skidegate Inlet, British Columbia. Mortoniceras (Durnovarites)
lowrii differs from the present species in having very depressed inner whorls,
somewhat more distant ribbing, and in lacking the modifications of the body
chamber ornament seen in the Angolan species. Mortoniceras (Durnovarites)
downii is based upon body chamber fragments. In its swollen, clavate tuber-
culation it approaches M. (D.) collignoni, but it appears to be more inflated, and
shows no sign of the body chamber becoming lanceolate.
Mortoniceras (Durnovarites) depressum (Spath) (1922: 114, figs B, 2a—d) is
based upon a body chamber fragment of a specimen about 50 mm in diameter.
At this stage, the whorls are very depressed (W/y = 1,41) and the fragment
appears to have been very evolute. Slightly prorsiradiate ribs arise from
prominent umbilical bullae and are indistinctly bifurcating, so as to appear
alternating long and short. There are four rows of tubercles, with maximum
width at mid-flank. This species would seem to be closely allied to M. (D.)
subquadratum var. tumidum Spath; it differs from the Angolan material in being
282 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 70. The holotype of J. Sowerby’s Ammonites rostratus, from the Upper Greensand of
Roak, near Benson, Oxfordshire. Oxford University Museum K835. 0,75.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 283
more coarsely ribbed and (?) more evolute, with a strongly depressed whorl
section.
Mortoniceras ( Durnovarites) baueri Collignon (1963: 159, pl. 305 (fig. 1311))
differs from M. (D.) collignoni sp. nov. in its coarse ribbing, depressed whorls,
wide umbilicus and prominent umbilical tubercles. Its body chamber ornament
is not known. Mortoniceras (Durnovarites) rerati Collignon (1963: 162, pl. 307
(figs 1312—1313)) differs from the Angolan material in much the same respects,
but does not have the swollen umbilical tubercles of M. (D.) baueri. It very
closely approaches M. (D.) ishiaguense.
Mortoniceras (Durnovarites) subdepressum Collignon (in Besairie 1936: 196,
pl. 21 (figs 4-5)) is based upon a unique fragment with a very depressed whorl
section. It was said to differ from M. (D.) depressum in having straighter ribs
which are not projected forwards on the venter. It is more coarsely ribbed with
a more depressed whorl section than M. (D.) collignoni.
Occurrence
Mortoniceras (Durnovarites) collignoni is known only from the uppermost
Albian of Angola.
Subgenus Angolaites Spath, 1932
Type species Subschloenbachia gregoryi Spath, 1922
Discussion
Angolaites was separated as a subgenus of Mortoniceras (Spath 1932: 380)
for “. . . serpenticones, with single costation from a very early stage, and two
peripheral tubercles, close together’. The characters of the subgenus are con-
sistent, making it an easily recognized and useful taxon.
Amongst mortoniceratinids, only Drakeoceras Young, 1957, and Canta-
brigites Spath, 1933, have the same closely spaced ventrolateral tubercles whilst
also lacking flank tubercles. Drakeoceras appears, however, to be a Goodhallites
derivative characterized by its much narrower umbilicus and high whorls. The
micromorph Cantabrigites is a contemporaneous form, abundant in western
Europe where Angolaites is unknown, while the extreme rarity of Cantabrigites
in Angola makes it unlikely that they represent sexual dimorphs.
Mortoniceras (Angolaites) gregoryi (Spath, 1922)
Figs 39G, 71, 72C, 73D
Subschloenbachia gregoryi Spath, 1922: 127, pl. 3 (fig. 1).
Mortoniceras (Angolaites) gregoryi (Spath) Reyment, 1955: 37, pl. 4 (fig. 13), pl. 6 (fig. 3).
Material
Three specimens, SAM-PCA3110, 3145 and 3235, from Praia-Egito,
together with seven specimens from the Quissama Ridge of Cabo Ledo, SAM-—
PCA4601, 4608, 4611, 4626, 4685, 4712, and 4813, all preserved as composite
internal moulds.
284 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 71. Mortoniceras (Angolaites) gregoryi (Spath). The holotype, from the uppermost
Albian at Catumbella, BMNH C20066. x1.
Description
The shell is very evolute, compressed, with a wide, shallow umbilicus
(42-47% of the diameter). The umbilical walls are gently rounded and the
flanks are slightly convex intercostally, converging somewhat towards the
venter, with greatest width slightly above the umbilical shoulder.
Ribs begin very weakly on the umbilical wall and pass radially outwards to
the umbilical shoulder where they terminate in fairly prominent bullae, about
seventeen per whorl. Each bulla gives rise to one to two flank ribs, with frequent
intercalatories which become more abundant in maturity. Across the flanks the
ribs are prorsiradiate and all are ornamented with closely spaced double ventro-
lateral tubercles. There are thirty-nine ventrolateral tubercles on the outer
whorl, and the upper ventrolateral tubercles are spirally notched. The venter is
moderately narrow, with shallow sulci on either side of the siphonal keel.
‘TX “SPIEWOd-WVS JO MaIA [eI9}e'T “(YyredS)
NN 1408048 (Sajwjosup) svdauojsop *D ‘OOTEWOd-WYS JO SMOIA eNUDA pue [e1eT “eYOyD) xajduus (sayivjosuy) svsaquojsop ‘g-y “ZL “B14
85
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL
286 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 73. A—C. Mortoniceras (Angolaites) simplex (Choffat). A. Lateral view of SAM-—PCA3142.
B. Lateral view of SAM-—PCA3153. C. Lateral view of SAM-—PCA3114. D. Mortoniceras
(Angolaites) gregoryi (Spath). Lateral view of SAM—PCA3147. A x0,66, B-D x1.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 287
Measurements
No. D H W W/a U
SAM-PCA3145 85 25(29) 20(24) 0,8 40(47)
e 58 18(31) —(—) — 27(47)
SAM-PCA3235 67 22(32) 20(30) 0,9 28(42)
Discussion
Mortoniceras (Angolaites) gregoryi differs from M. (A.) simplex (Choffat)
and M. (A.) vicina (Haas) in the common occurrence of bifurcating and inter-
calated ribs, and in having far fewer umbilical bullae.
Occurrence
Mortoniceras (Angolaites) gregoryi is known with certainty only from the
Upper Albian of Angola and Nigeria.
Mortoniceras (Angolaites) simplex (Choffat, 1905)
Figs 3A—D, 54G-H, 72A-B, 73A-C, 74-77
Schloenbachia simplex Choffat, 1905: 35, pl. 4 (fig. 3).
Pervinquieria simplex var. tenuis Haas, 1942: 81, pl. 16 (fig. 1), figs 7e, 8a.
Inflaticeras sp. n. aff. gregoryi Spath, 1922: 127, pl. 3 (fig. 2).
Pervinquieria vicina Haas, 1942: 82, pl. 16 (fig. 2), fig. 8b.
Pervinquieria vicina var. evoluta Haas, 1942: 83, pl. 16 (fig. 3), fig. 8c.
Material
9 specimens, SAM—PCA3107, 3116, 3142, 3146, 3150, 3153, 3166, 3200 and
3249, from Praia-Egito, together with 30 specimens from the Quissama Ridge at
Cabo Ledo, SAM-PCA4575, 4578-79, 4581-82, 4584-85, 4588, 4590, 4593-94,
4596, 4605, 4609, 4613, 4615-16, 4618, 4628, 4631, 4640, 4718, 4756, 4770, 4774,
4863, 4867-69, and 4874, all preserved as composite internal moulds.
Description
This species is abundant at Egito, with adult specimens attaining a diameter
of 170 mm (SAM-PCA3142).
The shell is evolute, compressed, with the outer whorls only covering the
preceding whorls to the top of the lower ventrolateral tubercles. The umbilicus
is shallow and wide (41-47% of the diameter), with steep umbilical walls and
well-rounded umbilical shoulders.
Ribbing begins at the umbilical seam, is rather faint at first, strengthening
as it passes radially outwards to the umbilical shoulder. The ribs may strengthen
slightly on the umbilical shoulder, but true umbilical tubercles are absent. In the
immature growth stages, the flank ribs are slightly sinuous but on the final whorl
they become adorally concave. Rare intercalated ribs occur only during the very
early ontogenetic stages, and there are about twenty ribs per whorl. Lateral
tubercles are lacking, but all ribs are ornamented with closely spaced double
288 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 74. A-B. Mortoniceras (Angolaites) simplex (Choffat). Ventral and lateral views of
SAM-PCA3116. C—D. Mortoniceras (Angolaites) cf. simplex (Choffat). Lateral and ventral
views of SAM-PCA3179. x1.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 289
Fig. 75. Mortoniceras (Angolaites) vicina (Haas). The holotype BMNH-C20067. x1.
ventrolateral tubercles which are spirally notched, especially the upper ventro-
lateral tubercle. The narrow, rounded venter is keeled, with smooth sulci on
either side. At large growth stages, the ventrolateral tubercles stand somewhat
above the level of the keel.
Measurements
No. D H W W/H U
SAM-PCA3142 173 48(28) 41(24) 0,85 81(47)
si 135 45(33) 35(26) 0,77 56(41)
SAM-PCA3107 91 31(34) 24(26) 0,77 37(41)
SAM-PCA3200 — 43(—) 33(—) 0,76 —
SAM-PCA3249 — 33(—) 26(—) 0,78 —
SAM-PCA3150 — 39(—) 35(—) 0,89 —
SAM-PCA3153 — 22(—) 19(—) 0,86 —
SAM-PCA3146 iss si) 26(—) 0,84 a
290 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 76. Mortoniceras (Angolaites) simplex (Choffat). Whorl sections. A. SAM-—PCA3166.
B. SAM-PCA3116. x1.
Discussion
Spath (1922) briefly discussed and figured a specimen which he considered
to differ from M. gregoryi in having coarser, more distantly ribbed inner whorls
(Fig. 75). This specimen was renamed Pervinquieria vicina by Haas (1942); it is
in the British Museum (BMNH C20067) and, so far as the writers are able
to judge, differs from M. (A.) simplex only in its coarser, more distant ribbing
with five ribs in a distance equal to the whorl height, whereas in M. (A.)
simplex there are eight to nine. The differences are slight, and within the range
of variation seen in M. (A.) simplex from Egito; they are not regarded as of
specific significance.
Occurrence
Mortoniceras (Angolaites) simplex is known only from Angola.
Genus Cantabrigites Spath, 1933
Type species Mortoniceras (Cantabrigites) cantabrigense Spath;
by original designation
Discussion
The first appearance of the name Cantabrigites Spath (1932: 380) was as a
nomen nudum, the diagnosis and description of the type species appearing only a
year later (Spath 1933: 436). Consequently, the valid date of introduction for
Cantabrigites is 1933.
Spath (1933) proposed Cantabrigites as a subgenus of Mortoniceras for
‘dwarf-forms with reduced, generally single and almost untuberculate costation
and greatly simplified suture-line’. In maturity, many typical Mortoniceras
commonly exceed 200-300 mm in diameter, whereas Cantabrigites is mature at
diameters of less than 80 mm. It would appear, therefore, to be a genuine
micromorph taxon. The fact that Cantabrigites is restricted to one level in the
Upper Albian, viz. the dispar Zone, suggests that it is not the microconch of
Mortoniceras. Cantabrigites differs so greatly from Mortoniceras vespertinum
(Morton), the type of the genus, that the authors consider the differences
sufficient for generic separation.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 291
Fig. 77. Mortoniceras (Angolaites) simplex (Choffat). Whorl sections. A. SAM-—PCA3249.
B. SAM-PCA3147. C. SAM-PCA3150. D. SAM-PCA3144. E. SAM-PCA3200.
F. SAM-PCA3153. G. SAM-PCA3111. H. SAM-PCA3179. I. SAM-PCA3200. x1.
292 ANNALS OF THE SOUTH AFRICAN MUSEUM
Cantabrigites ? curvatum Renz, 1968
Figs 78-79
Cantabrigites curvatum Renz, 1968: 61, pl. 11 (figs la—b, 2a-c), figs 20g, 21f.
Material
A single specimen, SAM-PCA3177, from a horizon some way below the
main occurrence of Stoliczkaia at Egito, in road gravels.
Description
The shell is evolute, with a wide, moderately deep umbilicus and a slightly
compressed, subquadrate intercostal whorl section. The umbilical shoulders
are well rounded and the flanks convex, with maximum width at about midflank.
Ribs begin at the umbilical seam and pass radially outwards to small, but
distinct, bullae on the umbilical shoulder. The ribs are rather thick, robust, and
vary from slightly rursiradiate to slightly prorsiradiate across the flanks. Where
no umbilical bullae are present, some ribs are intercalated at the level of the
umbilical shoulder, so that there are eleven ribs per half whorl, of which six
arise from umbilical bullae. Each rib is ornamented with a prominent, obliquely
clavate, ventrolateral horn which projects backwards. There appears to be the
faintest swelling just below the ventrolateral horns which may represent a very
weakly developed lower ventrolateral tubercle. There is a distinct siphonal keel.
Measurements
No. D H W W/H U
SAM-PCA3177 29 11(38) 9(31) 0,82 13(41)
Discussion
The Angolan specimen is indistinguishable from the holotype from
Ste Croix, Switzerland, although the writers are in some doubt as to the generic
identity of this species, since it closely resembles some of the earlier Neokentro-
ceras spp. known from Angola. As, however, this species is known only from
three small specimens and Neokentroceras is typical of the low Upper Albian of
Angola and Nigeria, tentative assignment to Cantabrigites seems preferable, it
being an homoeomorphous development only.
Fig. 78. Cantabrigites? curvatum Renz. Lateral
and ventral views of SAM-—PCA3177. Sill
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 293
Fig. 79. Cantabrigites? curvatum Renz. Whorl
section of SAM-PCA3177. x2.
Mortoniceras (Durnovarites) neokentroides Wiedmann & Dieni (1968: 146,
pl. 13 (fig. 5)) was, as its specific name implies, considered to resemble
Neokentroceras. It differs from the present species in having well-developed
upper and lower ventrolateral tubercles which are elongated at right angles to
the keel.
Mortoniceras ? nanum Spath (1933: 411, pl. 43 (fig. 6), pl. 46 (figs 4-5),
fig. 141) (Fig. 80) resembles the present species, but has double ventrolateral
tubercles and lacks the posteriorly directed horns of C? curvatum.
Occurrence
Cantabrigites ? curvatum Renz is known only from Switzerland and Angola.
Fig. 80. Mortoniceras? nanum Spath. The holotype,
BMNH-C72726, from the Cambridge Greensand. x1.
Genus Drakeoceras Young, 1957
Type species Drakeoceras drakei Young, 1957
Discussion
Young (1957) erected the new genus Drakeoceras for Goodhallites-like
forms in which the ventrolateral tubercles are doubled. As such, therefore,
Drakeoceras bears the same relationship to Goodhallites that Angolaites does to
294 ANNALS OF THE SOUTH AFRICAN MUSEUM
Mortoniceras, and further work may show that it is best treated as a subgenus of
Goodhallites. Drakeoceras differs from M. (Angolaites) in being higher whorled,
with Goodhallites-like :nner whorls and a more quadrate whorl section in
maturity, as well as retaining prominent umbilical bullae and bifurcating ribs
onto the body chamber.
Drakeoceras cf. dellense Young, 1957
Figs 3E-F, 81-82
Compare
Drakeoceras dellense Young 1957: 25, pl. 7 (fig. 1), pl. 10 (figs 4-5, 8-10),
figs 2j, 3a, h-j.
Material
Six specimens, SAM-—PCA4662, 4673, 4705, 4733, 4786, and 4800, from the
Quissama Ridge at Cabo Ledo, all poorly preserved limestone steinkerns, appear
to belong here.
Description
The shell is strongly compressed and moderately involute (umbilicus
23-25 % of the diameter). The umbilicus is fairly narrow and shallow, with steep
umbilical walls and evenly rounded umbilical shoulder. The broad flanks are
slightly convex and converge towards the narrowly arched venter. The whorl
section is compressed, elliptical (¥/H = 0,70-0,94).
Fig. 81. Drakeoceras cf. dellense Young. Ventral and lateral views of
SAM-PCA4800. x1.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 295
Ornament comprises about twelve weak umbilical bullae per half-whorl,
from which ribs commonly arise in pairs, or singly with an intercalated rib
between long ribs. The ribs are initially strongly prorsiradiate, but recurve just
below midflank. The ribs are broader than the interspaces and there are about
twenty-four per half-whorl at the venter. All ribs are ornamented with very weak
(? due to abrasion) lower and distinct upper ventrolateral tubercles. There is a
prominent siphonal keel.
Fig. 82. Drakeoceras cf. dellense Young. Eroded suture at 65 mm diameter. Approx. x 3.
Discussion
The Angolan specimen is closest to D. dellense from which it differs only in
being somewhat more involute. It is significant that in Texas D. dellense is
associated with M. (Durnovarites) perinflatum (Spath) (= D. adkinsi Young)
and is probably, therefore, strictly contemporaneous with the Angolan speci-
mens. Differences from other species of Drakeoceras are noted by Young (1957).
Occurrence
Drakeoceras dellense is currently known only from the dispar Zone of Texas
and probably Angola.
296 ANNALS OF THE SOUTH AFRICAN MUSEUM
Family Binneyitidae Reeside, 1927
Genus Borissiakoceras Arkhangel’skii, 1916
Type species Borissiakoceras mirabile Arkhangel’skii, 1916
Discussion
As currently diagnosed (Wright in Arkell et al., 1957), the Binneyitidae is a
family of micromorph ammonites characterized by their compressed, flat-sided
form, narrow umbilicus and greatly simplified suture. Considered to be
descended from the typically Middle Albian Falciferella, the earliest recorded
binneyitids are from the Middle Cenomanian of the Western Interior (Cobban
1961), although we know of specimens of comparable age from western Europe
(Kennedy & Juignet 1973) and Zululand. The time separating the last appearance
of Falciferella and the first appearance of Borissiakoceras has been something
of a problem in this phylogenetic scheme. However, Brunnschweiler (1959) has
recorded species of ‘Falciferella’ from the late Albian of Australia which appear
to bridge this gap. Our present record of Borissiakoceras from the uppermost
Albian of Angola closes the gap even further, and, by extending the record of
Borissiakoceras as far back as the late Albian, provides a direct morphological,
chronological and phylogenetic link with the Middle Albian Falciferella.
Cobban (1961: 747) diagnosed Borissiakoceras as follows: ‘This genus is
characterized by the small size of the conchs which are ordinarily compressed
and moderately evolute to somewhat involute. The venter is rounded to flat.
Most shells are smooth but a few have raised falcoid growth lines or faint closely
spaced falcoid ribs. Nodes, when present, are on the ventrolateral shoulder. The
suture has a broad ventral lobe indented by a broad, shallow ventral saddle. The
first lateral saddle is bifid and as wide as the ventral lobe. The first lateral lobe is
narrow, bifid, and about half as wide as the ventral lobe. The second lateral
saddle is trifid and higher and broader than the first. The second lateral lobe is
about half as large as the first and tends to be bifid. The auxiliaries, which are
small and shallow, may be divided into bifid lobes and saddles.’
Casey (1954: 27) introduced Falciferella (type species: F. milbournei Casey,
1954: 274, pl. 7 (figs 1-5), fig. 3) as follows: ‘Micromorph platycones with
narrow, sharp-rimmed umbilicus and tabulate venter, feebly carinate in early
youth. Test with strongly falcoid lineation or sub-costation, and a faint spiral
groove at the middle of the sides. Mouth border plain. Suture-line of simplified
Aconeceras pattern, with reduced auxiliary elements.’
In southern England, Falciferella is known only from the Middle Albian
Euhoplites loricatus Zone (intermedius and niobe Subzones) (Owen 1971),
although Brunnschweiler (1959) has recorded two species from the Upper Albian
of Australia. Falciferella breadeni Brunnschweiler (1959: 15, pl. 1 (figs 5—6)) is
associated with Labeceras and Myloceras and is thus of early late Albian age.
In this species, the earliest whorls are smooth, but strong falcate ribs develop in
maturity and form weak folds across the tabulate venter. The suture-line of
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 297
F. breadeni is simpler than in F. milbournei, in which respect it is closer to
Borissiakoceras. It differs, however, in the trifid nature of the first lateral
lobe, but there is a tendency towards trifurcation of this sutural element in
B. orbiculatum Stephenson (cf. Cobban 1961, fig. Sa, f). Although a trifid first
lateral lobe is known in Falciferella, the characters of F. breadeni are largely
those of Borissiakoceras, and it is to the latter genus that we refer ‘F.’ breadeni
and ‘F.” reymenti.
Borissiakoceras sp. nov. ? aff. reymenti (Brunnschweiler, 1959)
Fig. 31H-I
Compare
Falciferella reymenti Brunnschweiler 1959: 15, pl. 1 (figs 5-6).
Material
A single specimen, USNMNH 236980, with recrystallized shell preserved
from Porto Amboim.
Description
Shell small, compressed (in part due to post-mortem deformation), with
broad, flat flanks and a narrowly rounded venter. In places the venter appears to
be fastigiate, but this is probably due to secondary crushing. The umbilicus is
narrow (23% of the diameter) and the shell smooth.
Measurements
No. D H W W/a U
USNMNH 236980 17 +7,3(43) +3(18) 0,41 4(23)
Discussion
The only other species of Borissiakoceras yet known from late Albian
strata are B. breadeni (Brunnschweiler) (1959: 15, pl. 1 (figs 5—6)) and B. reymenti
(Brunnschweiler) (1959: 16, pl. 1 (figs 7-8)) from the Oodnadatta region of
South Australia. Both these species differ from the Angolan specimen in their
much narrower umbilicus (13-14% of diameter) and in possessing rather
prominent falcate ribs. Cobban (1961) has shown, however, that the latter
feature is not consistently developed, even within a single Borissiakoceras
population, with both ribbed and smooth variants occurring side by side.
Consequently differences in ornament of this type may not be of specific
importance.
Smooth variants of Borissiakoceras orbiculatum Stephenson (1955: 64,
pl. 6 (figs 1-4)) closely resemble the present specimen, but are much younger
(Middle Cenomanian). Borissiakoceras compressum Cobban (1961: 747, pl. 87
(figs 19-33), pl. 89 (figs 1-9), fig. 4a-k) also resembles the present species, but is
of Middle Cenomanian age and has a narrower umbilicus (14-17% of the
diameter). Borissiakoceras mirabile Arkhangel’skii (1916: 55, pl. 8 (figs 2-3))
298 ANNALS OF THE SOUTH AFRICAN MUSEUM
differs from the Angolan example in its much wider umbilicus (37% of the
diameter) as well as being a Lower Turonian species. Borissiakoceras reesidei
Morrow (1935: 463, pl. 49 (fig. 7), pl. 50 (fig. 5), fig. 8) is an Upper Cenomanian
species which, like B. orbiculatum, cannot be separated from the unique Angolan
example without knowledge of the suture-line characteristics of the latter.
Occurrence
Upper Albian of Angola; species with which the specimen is compared
came from the Upper Albian of Australia.
AGE OF THE FAUNA
At present there are certain problems concerning the scope and nomen-
clature of the Stoliczkaia dispar Zone and its subzones. The divisions of the
Albian Stage in England were erected by Spath on the basis of his understanding
of the successions at Folkestone and elsewhere in England, and are based on firm
stratigraphic principles. Spath (1943) gave the following zonation through the
Upper Albian of southern England:
Stoliczkaia dispar/ Durnovarites perinflatum
Stoliczkaia dispar Zone | Subzone
Arrhaphoceras substuderi Subzone
Mortoniceras aequatoriale Subzone
Callihoplites auritus Subzone
Hysteroceras varicosum Subzone
Hysteroceras orbignyi Subzone
Mortoniceras inflatum Zone
Whilst accepting Spath’s subdivision of the S. dispar Zone, Breistroffer
(1940) showed that Arrhaphoceras substuderi (Spath) was common to both
the perinflatum/dispar and substuderi Subzones and was, therefore, unsuitable
for use as a subzonal index. Instead, he recognized a lower subzone of
S. (Faraudiella) gardonica—S. (F.) rhamnonota— Mariella gresslyi—Turrilitoides
toucasi. By 1947, however, Breistroffer had realized that S. (F.) gardonica
and S. (F.) rhamnonota were synonyms of S. (F.) blancheti (Pictet & Campiche)
and consequently renamed Spath’s A. substuderi Subzone the ‘Zone 4 Para-
turrilites Gresslyi, Turrilitoides Hugardianus et Stoliczkaia (Faraudiella)
Blancheti’. The upper horizon (Spath’s dispar/perinflatum Subzone) was referred
to a ‘Zone a Pervinquieria (Durnovarites) perinflata, P. (Subschloenbachia)
rostrata, Paraturrilites Bergeri et Stoliczkaia dispar’.
On the basis of newly exposed sections, however, Owen (1976) has recently
shown that Spath’s aequatoriale Subzone is a remanié assemblage which cannot
be distinguished from the underlying subzone of Callihoplites auritus and he has,
therefore, proposed its abandonment. In addition, Owen (1976) recalled Brei-
stroffer’s (1940) otservation that Arrhaphoceras substuderi was unsuitable for
use as a subzonal index in the S. dispar Zone and, because Owen (1976: 492)
quite inexplicably considered S. (F.) blancheti ‘. . . is not sufficiently distinct to
ay,
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 299
be used as a subzonal index’, he proposed to replace this subzonal index with
Mortoniceras rostratum (J. Sowerby). This, however, is a very unfortunate
suggestion because M. rostratum was for many years (because of mis-
identification) the index species for the earlier zone now called after M. inflatum.
In addition, Breistroffer (1947) has subsequently used it as one of the subzonal
indices for Spath’s dispar/perinflatum Subzone. Moreover, it is abundantly clear
that the true characters of M. rostratum are not known since all the material
assigned by Spath (1932, pl. 38 (fig. 4), pl. 39 (fig. 4), pl. 40 (figs 1, 7), pl. 41
(fig. 7)) to this species was renamed Pervinquieria fallax by Breistroffer (1940: 67).
Thus, only Sowerby’s holotype (Fig. 70), now in the Oxford University Museum,
is without doubt assignable to this species. The authors’ have had the oppor-
tunity of studying the holotype of M. rostratum. Its inner whorls are obscured
by matrix whilst the ribs of the body chamber bear four rows of tubercles.
Although a number of species of Mortoniceras s.s. have four rows of tubercles
on the inner whorls, the upper and lower ventrolateral tubercles coalesce on the
body chamber of M. (Mortoniceras) to produce ventrolateral horns and thus
only three rows of tubercles remain on the body chamber. In the writers’
opinion, therefore, M. rostratum is a species of Durnovarites, a subgenus so far
recorded only from Spath’s dispar/perinflatum Subzone, and is unsuited,
therefore, for use as the subzonal index of the lower part of the S. dispar Zone.
Moreover, since Owen (1976) does not state what he takes to represent
M. rostratum, the species to which he is referring is totally unrecognizable. For
this reason, the writers would propose a simple return to Breistroffer’s (1947)
subdivision of the S. dispar Zone into a lower subzone of S. (F.) blancheti,
immediately overlying the Callihoplites auritus Subzone, and an upper subzone
of M. (Durnovarites) perinflatum.
With the possible exception of the lower horizon at Egito, the present
faunas can be referred with confidence to the M. (D.) perinflatum Subzone on
the basis of the presence of S. (Stoliczkaia), M. (Durnovarites) and the hetero-
morphs present. There does, however, seem to be some compositional difference
in the faunas. Thus, at Praia-Egito the fauna comprises :*
Anisoceras perarmatum Pictet & Campiche
. armatum (J. Sowerby)
. haasi sp. nov.
. phillipsi sp. nov.
. ef. arrogans (Giebel)
. aff. exoticum Spath
. aff. subarcuatum Spath
. aff. spathi (Wiedmann)
Hamites virgulatus Brongniart
H. duplicatus Pictet & Campiche
Puzosia cf. sharpei Spath
Desmoceras latidorsatum perinflatum subsp. nov.
MRA AAA DR
* These lists on pp. 299-301 follow the order in the text. Ed.
300 ANNALS OF THE SOUTH AFRICAN MUSEUM
Stoliczkaia tenuis Renz
Mortoniceras (Durnovarites) collignoni sp. nov.
M. (Angolaites) simplex (Choffat)
M. (A.) gregoryi (Spath)
A somewhat lower level at this locality has yielded:
Tetragonites kitchini (Krenkel)
Mariella gresslyi (Pictet & Campiche)
Cantabrigites ? curvatum Renz
The fauna from Cabo Ledo includes the following species:
Anisoceras perarmatum Pictet & Campiche
A. armatum (J. Sowerby)
A. phillipsi sp. nov.
Idiohamites dorsetensis Spath
Hamites virgulatus Brongniart
Mariella cf. oehlerti (Pervinquiére)
Stoliczkaia sp.
Mortoniceras (Durnovarites) perinflatum (Spath)
M. (Angolaites) simplex (Choffat)
M. (Angolaites) gregoryi (Spath)
M. (Mortoniceras) spp.
Drakeoceras cf. dellense Young
This fauna differs from the higher horizon at Praia-Egito in that
M. (Mortoniceras) is still fairly abundant whilst M. (Durnovarites) is rather rare.
This suggests that the Cabo Ledo fauna may be somewhat older than the upper
horizon at Praia-Egito, although the possibility of mixing of different horizons
cannot be wholly dismissed.
The fauna from Porto Amboim comprises:
Phylloceras (Hypophylloceras) seresitense Pervinquiére
Tetragonites collignoni Breistroffer
T. jurinianus (Pictet)
Eogaudryceras italicum Wiedmann & Dieni
Anisoceras perarmatum Pictet & Campiche
A. armatum (J. Sowerby)
A. haasi sp. nov.
Idiohamites dorsetensis Spath
I. cf. elegantulus Spath
I. pygmaeus sp. nov.
Hamites virgulatus Brongniart
Mariella circumtaeniatus (Kossmat)
Desmoceras latidorsatum perinflatum subsp. nov.
Stoliczkaia tenuis Renz
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 301
Mortoniceras (Durnovarites) perinflatum (Spath)
M. (D.) subquadratum Spath
M. (Angolaites) simplex (Choffat)
Hysteroceras ? cf. ootaturense (Stoliczka)
Borissiakoceras sp. nov? aff. reymenti (Brunnschweiler)
The abundance of very compressed Stoliczkaia at Porto Amboim, together
with the relative rarity of mortoniceratinids, suggests that this fauna may be
somewhat younger than the upper horizon at Praia-Egito, although stratigraphic
evidence for this is lacking. None the less, the Stoliczkaia fauna from here bears
a very close resemblance to the ‘Submantelliceras’— Utaturiceras assemblages
that characterize the basal Cenomanian of many regions, although the persistence
of Mortoniceras at this level suggests that it is still best regarded as uppermost
Albian.
When the composition of the Angolan faunas is considered, there are
marked differences from the faunas of the European perinflatum subzone in
addition to the obvious absence of hoplitids. Noticeable in the Angolan
assemblage is the complete absence of the widely distributed Mariella of the
bergeri-miliaris group, Lechites, Stoliczkaia (Faraudiella), Turrilitoides, Scaphites,
Ostlingoceras and nautiloids, together with the rarity of Hamites and Canta-
brigites. Moreover, Stoliczkaia tenuis is rather different from the S. dorsetensis—
notha—dispar plexus which characterizes the uppermost Albian of western
Europe.
Although some differences may be ecological (in particular with respect to
the heteromorphs and the absence of hoplitinids), it seems possible that two
slightly different levels in the uppermost Albian are represented. It is perhaps
noteworthy, therefore, that in Texas, M. (D.) perinflatum is known only from
the Pawpaw Formation (Young 1957), some way below the basal Cenomanian
faunas of the uppermost Main Street and Grayson/Del Rio sequences. It also
occurs below the main level of Stoliczkaia in Poland (Marcinowski & Naidin
1976).
Notwithstanding these differences, the abundance of Stoliczkaia, together
with M. (Durnovarites) perinflatum and Anisoceras of the perarmatum group, is
sufficient to date the Porto Amboim fauna at M. (D.) perinflatum Subzone of the
S. dispar Zone, and to point to the possibility of even further subzonal refinement
of this critical interval.
SUMMARY
The Angolan littoral has yielded rich ammonite faunas referable to the
uppermost Albian zone of Stoliczkaia dispar, and mainly to the upper subzone
of Mortoniceras (Durnovarites) perinflatum. The following species are described :
Phylloceras (Hypophylloceras) seresitense Pervinquiére
Tetragonites (Tetragonites) collignoni Breistroffer
Tetragonites (Tetragonites) kitchini (Krenkel)
302 ANNALS OF THE SOUTH AFRICAN MUSEUM
Tetragonites (Tetragonites) jurinianus (Pictet)
Eogaudryceras (Eogaudryceras) italicum Wiedmann & Dieni
Anisoceras (Anisoceras) perarmatum Pictet & Campiche
Anisoceras (Anisoceras) armatum (J. Sowerby)
Anisoceras (Anisoceras) haasi sp. nov.
Anisoceras (Anisoceras) phillipsi sp. nov.
Anisoceras (Anisoceras) cf. arrogans (Giebel)
Anisoceras (Anisoceras) aff. subarcuatum Spath
Anisoceras (Anisoceras) aff. exoticum Spath
Anisoceras (Anisoceras) cf. spathi (Wiedmann)
Idiohamites dorsetensis Spath
Idiohamites cf. elegantulus Spath
Idiohamites pygmaeus sp. nov.
Hamites virgulatus Brongniart
Hamites duplicatus Pictet & Campiche
Mariella (Mariella) circumtaeniatus (Kossmat)
Mariella (Mariella) gresslyi (Pictet & Campiche)
Mariella (Mariella) cf. oehlerti (Pervinquiére)
Mariella (Mariella) nobilis (Sukes-Browne)
Puzosia (Puzosia) cf. sharpei Spath
Desmoceras (Desmoceras) latidorsatum perinflatum subsp. nov.
Stoliczkaia (Stoliczkaia) tenuis Renz
Hysteroceras ? cf. ootaturense (Stoliczka)
Mortoniceras (Durnovarites) perinflatum (Spath)
Mortoniceras ( Durnovarites) subquadratum Spath
Mortoniceras (Durnovarites) collignoni sp. nov.
Mortoniceras (Angolaites) simplex (Choffat)
Mortoniceras (Angolaites) gregoryi (Spath)
Drakeoceras cf. dellense Young
Cantabrigites ? curvatum Renz
Borissiakoceras sp. nov. ? aff. reymenti (Brunnschweiler)
The majority of these species have not previously been described from
Angola; although precisely correlated with the perinflatum Subzone, there are
differences in composition when compared with European faunas. In part these
reflect differences between biogeographic provinces (e.g. the sparsity or absence
of some heteromorphs). Other differences suggest that it may be possible to
further subdivide the dispar Zone and the fauna also permits discussion of recent
reviews of Upper Albian zonation, especially by Owen (1976). A return to a
twofold division of the dispar Zone into Stoliczkaia (Faraudiella) blancheti and
Mortoniceras (Durnovarites) perinflatum Subzones is suggested.
The chief systematic conclusions from the paper are discussions of intra-
specific variation in Phylloceras (Hypophylloceras), T. (Tetragonites),
D. (Desmoceras), E. (Eogaudryceras) and S. (Stoliczkaia) species.
~w
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 303
ACKNOWLEDGEMENTS
Our best thanks are to Messrs F. Collier and E. G. Kauffman (U.S. National
Museum) for allowing us to study the Washburn Collection. Dr M. K. Howarth,
Dr H. G. Owen and Mr D. Phillips (British Museum), Dr R. Casey and
Mr C. J. Wood (Geological Museum, London), Dr J. M. Hancock (London),
Mr C. W. Wright (Oxford), Dr B. Rickards (Cambridge), Dr J. Sornay (Paris),
and Dr J. P. Lefranc (Montpellier) assisted us in many ways, and their help is
gratefully appreciated. Mr C. W. Wright critically reviewed the manuscript and
saved us from a number of errors, for which we are most grateful.
We also thank the staff of the Department of Geology and Mineralogy,
Oxford, and the Geological Collections of the Oxford University Museum for
their help. The fieldwork for this paper was undertaken while one of us (M.R.C.)
was on the staff of the South African Museum.
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6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-1SA
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author's name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.
In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and date.
7. SPECIAL HOUSE RULES
Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to in the text __
e.g. *... the Figure depicting C. namacolus ...’; *. . . in C. namacolus (Fig. 10)...”
(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names ?
e.g. DuToit but A.L.du Toit; Von Huene but F. von Huene
(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian
Punctuation should be loose, omitting all not strictly necessary
Reference to the author should be expressed in the third person
Roman numerals should be converted to arabic, except when forming part of the title of a
book or article, such as
‘Revision of the Crustacea. Part VIII. The Amphipoda.’ ee ere
Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively. 7
Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.
M. R. COOPER
&
W. J. KENNEDY
UPPERMOST ALBIAN (STOLICZKAIA DISPAR ZONE)
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