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FiscHer, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
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Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
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ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 77 7 Band
June 1979 Junie
Part 10 #£Deel
UPPERMOST ALBIAN (STOLICZKAIA DISPAR ZONE)
AMMONITES FROM THE ANGOLAN LITTORAL
By
M. R. COOPER
&
Wiad) 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 Il 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 Pervinquiere, Anisoceras haasi sp. noy., and Mortoniceras
(Durnovarites) collignoni sp. noy. Other important elements present include Idiohamites
dorsetensis Spath, J. 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 . 3 : : 3 : , ; 6
Systematic palaeontology . 5 3 ; é so iesikata!
Family Phylloceratidae : ; : ; nee lari,
Genus Phylloceras . ; : ; F a wae
Family Tetragonitidae . j ; ; ; a RSI
Genus Tetragonites : : : : 5 Altsil
Family Gaudryceratidae ; ; ; 2 a 6189
Genus Eogaudryceras . : ; , galls)
Family Ancyloceratidae : : p ; 7) 196
Genus Anisoceras . : ; ; : ~ L196
Genus Idiohamites . ; ; 4 : ne 222.
Family Hamitidae . : - : : ; 226
Genus Hamites . : : ; : e226
Family Turrilitidae : : : : : se DRS
Genus Mariella. 3: ; F : 5 DRS
Family Desmoceratidae : ; j : = 235
Genus Puzosia 2 5 : : F 35)
Genus Desmoceras P F : é Se 487)
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 . 3 5 : suc, 2Ad
Genus Stoliczkaia . : : E : . 243
Family Brancoceratidae : : ‘ : 5 2X85)
Genus Hysteroceras? . 4 : : . 265
Genus Mortoniceras . : : : . 269
Subgenus M. (Durnovarites) : : . 269
Subgenus M. (Angolaites) . ; j S88,
Genus Cantabrigites? . ; : ; = 290
Genus Drakeoceras : : : ; Hy293
Family Binneyitidae . s ; : : 9296
Genus Borissiakoceras . : s : . 296
Age of the fauna . : : : : : : a ASS
Summary : : : 5 : : ; : . 301
Acknowledgements : : : ‘ : : e303
References. ; ; ; ; : 5 : = 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; W/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). Bose, 1923: 119,
pl. 7 (figs 15-17). Spath, 19254: 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; 19625: 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 Pervinquiére, 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, Foérster, 1975: 139, fig. 26.
? Phylloceras serum vat. 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 Pervinquiere, Matsumoto, 1942: 674, fig. 2.
Phylloceras cf. seresitense Pervinquiere, 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/H = 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/x U
USNMNH 236897 37 21(57) 13(35) 0,62 2,6(7)
9 26,5 15,5(58) +8,5(32) 0,55
USNMNH 236898 2 30,5(59) 17,5(34) 0,57 3(6)
9 35 19(54) 11(31) 0,58 ?
USNMNH 236900 I) 8(53) 5(33) 0,63 1,1(7)
99 1] 5,5(50) 3,7(34) 0,67 ?
USNMNH 236901 17 10(59) 6,5(38) 0,65 1,0(6)
USNMNH 23690221 12(57) 717) - <0:64umeee
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 179
Fig. 1. Phylloceras (Hypophylloceras) seresitense Pervinquiére. 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. (#.) seresitense boulei Collignon—whorl section very compressed
(W/~ = 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 (W/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, 19626) 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
oo
o
WIDTH
MIGHT COP
0 30 60 90 120
DAME ER) ee ————EEeEEe——E—————
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 Pervinquiere; square = holotype of P. angolaense
Haughton; open triangles = P. seresitense seresitense Pervinquiere; 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. (H.) 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 T. 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 T. jurinianus (Pictet).
Tetragonites (Tetragonites) collignoni Breistroffer, 1940
Ammonites timotheanus Stoliczka (non Mayor), 1865: 146, pl. 73 (figs 34, 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 \UJ
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, 19626, 1973; Collignon 1963, 1964; Murphy 1967a, 1967b; Wiedmann &
Dieni 1968; McLearn 1972; Forster 1975; Kennedy & Klinger 19775). Forster
(1975) has suggested that 7. swbtimotheanus 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 TJ. 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.
A C
D E F
G
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/H = 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. 1.
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 L7G)
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 (non 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,
Ja—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 (W/H = 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 W W/a U
USNMNH 236915 S75) 25,5(49) +28(53) 1,10 10(19)
USNMNH 236917 38,5 20(52) +22(57) 10 +9(23)
a 30 14,5(48) 16(53) 1,10 9(30)
USNMNH 236918 43 19(44) 22,3(52) SIZ 9,8(23)
USNMNH 236919 16 7,5(47) 8,5(53) 1,13 4(25)
USNMNH 236920 18 8(44) 10(55) E25 +4(22)
USNMNH 236921 21 10(48) +10(48) 1,00 5(24)
USNMNH 236977 TRS 6,8(39) 8,5(49) Ws) 4,5(26)
USNMNH 236978 17 7(41) 9(53) 1-29 4,7(28)
T. jurinianus
angolanus
(holotype) 105 56(53) 51(48) 0,91 17(16)
- 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 T. 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
WA
0,70
0 30 60 90
DIAMETER (mm) —————>
40
ok
2) . é
a ~-08, _ 1On 2 fe)
= eJ o OR SRO os = Sees ae ae
Ova ON Pare
=
=)
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 7. 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 (19626: 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 (19626), 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/y = 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.
get Pagy.*
y
4
?
eX
*
f7
ot ul
€.'
Di as
f° 22
é ¢
Pa
£
AS ar
= - -— 7
+ Ps
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. 1.
192
ANNALS OF THE SOUTH AFRICAN MUSEUM
Measurements
No. D H W Wii U
USNMNH 236905 81 37,5(46) 25(31) 0,66 21(26)
i 56 25(45) IP.) O78 We3X(B1)
USNMNH 236906 34 13(38) 15,5(45) 1,19 +10(29)
USNMNH 236907 +47 +21(45) +18(38) 0,86 15,5(33)
USNMNH 236908 71 33,5(47) +27(38) 0,80 19(27)
USNMNH 236909 26 11(42) 11(42) 1,00 8(31)
ss 1955 8(41) 8(41) 100 +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 556i) 6(40) 1,09 5(30)
USNMNH 236912 22 8(36) 9(41) 1S 8(36)
USNMNH 236913 y 535(62) +7,5(44) 1,36 7(41)
USNMNH 236914 25 9(36) +11(44) 22 +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/H = 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
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), EF. (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. (E.) muntaneri Wiedmann.
Of other Upper Albian species of Eogaudryceras (Eogaudryceras),
Wiedmann (1962a: 154) selected a neotype for E. (E.) bourritianum (Pictet)
(in Pictet & Roux 1848: 298, pl. 4 (fig. 1)) and divided it into two chronological
subspecies: FE. (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
LEG
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 yraconense 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
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. 0,75.
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, 19255: 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 0,75,
E-F, I-J x1.
202 UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL
? Hamites aff. armatus J. Sowerby, Scott, 1926: 80. Bése, 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 ?236952 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, I. Anisoceras armatum (J. Sowerby). A. SAM-PCA5470. C. SAM-PCAS5471.
E. SAM-PCA5473. I. SAM-PCA5472. 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 venetzianus-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. nov. Lateral and dorsal views of SAM-PCA 4799. B x0,75,
other <1.
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 K67S5a. 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
/.
ih”
.
€
:
i
N
aor wy ne
Se
pet | at
acme =
A B
;
22th
i
dj 4;
E 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. | (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.
208
Pe ie it i * . * “
pes oe ee paceilh Skat oe 5 ee
aes SS
si sone a teititen “iy,
ANNALS OF THE SOUTH AFRICAN MUSEUM
aes Be,
SE pei ior MUTE IE AE i, 4 “
a
Fig. 20. Anisoceras haasi sp. nov. Reconstruction of early and middle growth stages. x1.
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 (“/yH = 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 21
&
.
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. haasi 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
DID 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
ere
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 PANS)
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 O75:
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL Die |
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
rursiradiate, 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.
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.
<OM/5:
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 22
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.
22D) 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—l, 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
Fi oe
me
é ees
«i Se <
<
Fig. 30. A-—D. Idiohamites dorsetensis Spath. A-B. Ventral and lateral views
of SAM-PCA4803. C-—D. Ventral and lateral views of SAM-PCA5469. 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 Jdiohamites 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, USNMNH 236951. H-I. Borissiakoceras sp. nov.? 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 Idiohamites 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 aperiure.
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. /diohamites
dorsetensis is 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, 111 (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. subyirgulatus 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
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 (non Pictet), 1861: 130, pl. 56 (figs 6-8 only).
Turrilites nobilis Jakes-Browne, 1877: 493, pl. 21 (fig. 1).
Mariella nobilis (Jukes-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 (1428 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,
19293165) pl. Wi(fig., 15):
Turrilitoides (?) 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 31e, 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 Gig. 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), Forster 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.
- aaeeane *
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. gress/yi 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 (“/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 37 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. 1.
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
(W/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,
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/a U
USNMNH 236961 21 +9(43) +12(57) 1,33 +4(19)
USNMNH 236962 14 7(50) 8,5(61) 1,21 2,3(16)
USNMNH 236964 27 13(48) 18,3(68) 1,41 +5(18)
USNMNH 236965 DA i1RS@5) 14(67) 1222
USNMNH 236966 23 9,5(41) 13(56) oy, +4,5(20)
USNMNH 236967 20 10,2(51) 12(60) 1,18 +4(20)
USNMNH 236968 123 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) 1,20 B
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) $s1(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 a 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. latidorsatum. 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
1,60
WIOTH
MEGKT 2 eee
0.70
0 40
DIAMETER e0rmm
40
=
a
= |
re)
=
3
— }
0
0 40
DIAMETER: == ms
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. /atidorsatum 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/q = 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. latidorsatum. 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 Le transitional between Stoliczkaia and Forbesiceras. Matsumoto
& Inoma (1975: 277) have also introduced the subgenus Stoliczkaia
(Shumarinaia).
ANNALS OF THE SOUTH AFRICAN MUSEUM
242
Fig. 39. A. Stoliczkaia (Stoliczkaia) tenuis Renz. Lateral view of SAM—PCA5478. 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.
S<il
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-—tlargilliertianum
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. x1.
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-
Fig. 43. Stoliczkaia (Stoliczkaia) villoutreysi Casey. Holotype, O. de Villoutreys collection,
Uppermost Albian, Monte Carlo Tunnel, Monte Carlo. 1.
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 Cottreauites (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 Cottreauites are juveniles of either Stoliczkaia or Paracaly-
coceras, and that it should be considered a nomen dubium.
248 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 45. Sroliczkaia (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. levis 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 (W/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. n 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/H = 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 p25) |
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. 1.
D2 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 48. Stoliczkaia (Stoliczkaia) tenuis Renz. A-C. USNMNH 236988.
237003. G-I. USNMNH 236990. x1.
D-F. USNMNH
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.
254 ANNALS OF THE SOUTH AFRICAN MUSEUM
E F
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
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)
1(47)
8(47)
4(42)
W
+22(33)
26(40)
17,6(31)
+23,5(35)
5
18(36)
+19(34)
208)
+10(26)
15,2(30)
18,7(34)
LED)
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
OFT
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)
?
4,7(16)
25(12)
?
+4,5(14)
4(14)
2,7(15)
E317)
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
Vp
D tie
A B Cc
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.
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL 259
Fig. 54. A-F. Stoliczkaia (Stoliczkaia) tenuis Renz. A—B. Lateral and ventral views of
SAM-PCA6811. C-D. Ventral and lateral views of SAM-—PCA2944. E. Lateral view of
SAM-PCA5477. 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 x0,75.
260 ANNALS OF THE SOUTH AFRICAN MUSEUM
0,80
WIDTH EIGHT a,
DIAMETER(mm) ——_________ 10
UMBILICUS (%) ——_____ >
0 70
DIAMETER(mm) ——HW—_____ a
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 Bose
(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. \t 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
Fig. 56. Stoliczkaia (Stoliczkaia) clavigera Neumayr. Copy of Stoliczka 1864, pl. 45 (fig. 1-la).
<li
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
: as
\ iS ra
N N ¥
.
q = 2
.
Fig. 59. Stoliczkaia crotaloides (Stoliczka). Copy of Stoliczka 1864, pl. 46 (fig. 3-3a).
<r
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 Ww 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 (?) 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) (Pervinquiére 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, 1926b: 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 (Durnoyarites) subquadratum Spath, USNMNH_ 237023.
D-I. Mortoniceras (Durnovarites) perinflatum (Spath). D-F. USNMNH 237021. G-I.
USNMNH 237022. x1.
Dae, ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 63. Mortoniceras (Durnovarites) perinflatum (Spath). Ventral and lateral views of
SAM-PCA4802. x1.
Measurements
No. D H W W/i U
USNMNH 237021 55 22,5(42) we ? 17(32)
a 44 +19(43) +20(45) 1,05 7
USNMNH 237022 28) +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 ZA
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
‘TX “60EEVOd-WVS ‘odAyexed v Jo
SMOIA [eIOIV] PUL [eIUDA “AOU “ds 10UFY]0I (Saj14DAOUING) SDABIIUOJAOW “A-A “SETEWOd-WYVS JO SMOIA [eIUIA pur [eJoeT “Yyedg wnosponbgns
(saqavaouind) SDAZNUOLIOW “A—D ‘OSTEWOd-WVS JO SMOIA [eae] pue [eIUSA “QeRyoyD) xajduus (sajiwjosup) svsvoUojso~ “F@-Y °S9 “SIF
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL Died
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/H U
USNMNH 237023 46 13,5(29) +15(33) twit 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/y = 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 211
Fig. 66. Mortoniceras (Durnovarites) collignoni sp. nov. A reconstruction based on the
holotype and paratype material. 0,66.
ANNALS OF THE SOUTH AFRICAN MUSEUM
278
‘TX 0990 E-V “8L7EVOd-WYVS “odAjesed &
JO MOIA [RIV] “OD “L7TEVOd-WYVS ‘adAj0OY BY} JO SMIIA [R.IUDA pUe [eINeT “G-Y “AOU “dS 1U0U81//09 (SAJJ4DAOUANG ) SDADIUOJAOPW “J-WY *L9 “B14
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/H = 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.
285
UPPERMOST ALBIAN AMMONITES FROM THE ANGOLAN LITTORAL
‘TX “SPIEVOd-NVS JO MOA [er9}e'T “(yeds)
1408948 (sajvjOSup) svsD2UOMOW *D ‘OOTEWOd-WYS JO SMOIA [eIUGA puL [eIOIeT “GeYOYD) xa duns (sanmosup) sproomouopy “GV °ZL Ba
286 ANNALS OF THE SOUTH AFRICAN MUSEUM
Bigs 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/y U
SAM-PCA3145 85 25(29) 20(24) 0,8 40(47)
= 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. x 1.
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 WwW Why U
SAM-PCA3142 173 48(28) 41(24) 0,85 81(47)
= 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 oe
SAM-PCA3249 — 33(—) 26(—) 0,78 _-
SAM-—PCA3150 — 39(—) 35(—) 0,89 —
SAM-PCA3153 — 22(—) 19(—) 0,86 —
SAM-PCA3146 a Be) 26) 0,84 pe
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. (4.)
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
lala
OFANo
aoe
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 Wa 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. ale
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 (W/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. x3.
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. \t 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. 5a, 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/H 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 a 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) okservation 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
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.
. cf. 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.
mwa A A A A A
* These lists on pp. 299-301 follow the order in the text. Ed.
300 ANNALS OF THE SOUTH AFRICAN MUSEUM
Stoliczkaia tenuis Renz
Mortoniceras (Durnoyarites) 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 (Pervinquieére)
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 ‘Suwbmantelliceras’— 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 (Jukes-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.
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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-naturw. KI. Akad. Wiss. Mainz 1963: 155-264.
WIEDMANN, J. 1973. The Albian and Cenomanian Tetragonitidae (Cretaceous Ammonoidea),
with special reference to the circum-Indic species. Eclog. geol. Helv. 66: 585-616.
WIEDMANN, J. & DIENI, I. 1968. Die Kreide Sardiniens und ihre Cephalopoden. Palaeontogr.
ital. 64: 1-171.
Woops, H. 1917. The Cretaceous faunas of the north-eastern part of the South Island of
New Zealand. Bull. geol. Surv. N.Z. Palaeontology 4: 1-41.
WRIGHT, C. W. & WRIGHT, E. V. 1951. A survey of the fossil Cephalopoda of the Chalk of
Great Britain. Palaeont. Soc. (Monogr.): 1-40.
YOUNG, K. 1957. Upper Albian (Cretaceous) Ammonoidea from Texas. J. Paleont. 31: 1-33.
ZWIERZYCKI, J. 1913. Zur Frage der Unteren Kreide in Portugiesisch-Mozambique. Sber. Ges.
naturf. Freunde Berl. 7: 319-326.
6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
Nnov., syn. nov., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year i
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.’ Ao ene
Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively. ;
Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.
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