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ANNALS OF THE ANNALE VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

VOLUME 82 BAND 82

EXIT HSONIAN

| 1981

\V\ [AN os

LIBRARIES

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

VOLUME 82 BAND

THE TRUSTEES OF THE DIE TRUSTEES VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

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1980

& SET, PRINTED AND BOUND IN THE REPUBLIC OF SOUTH AFRICA BY

THE RUSTICA PRESS (PTY.) LTD., WYNBERG, CAPE

515

LIST OF CONTENTS

Day, J.

Southern African Cumacea. Part 4. Families Gynodiastylidae and Diastylidae.

(Published August 1980.) : ee a Be Sy a

DINGLE, R. V.

Marine Santonian and Campanian Ostracods from a borehole at Richards Bay,

Zululand. (Published June 1980.) . Be:

HENDEY, Q. B. see MUIZON, C. DE

KAUFFMAN, E. G. see KLINGER, H. C.

KENNEDY, W. J. see KLINGER, H. C.

KENSLEY, B.

Marine isopods from Marion, Prince Edward, and Crozet Islands (Crustacea,

Isopoda). (Published August 1980.) a = a : ie

KLINGER, H. C., KAUFFMAN, E. G. & KENNEDY, W. J.

Upper Cretaceous ammonites and inoceramids from the off-shore Alphard Group

of South Africa. (Published September 1980.) Ee a Be

KLINGER, H. C. & KENNEDY, W. J.

Cretaceous faunas from Zululand and Natal, South Africa. A new sextuberculate

texanitid. (Published September 1980.)

MILLARD, N. A. H.

The South African Museum’s Meiring Naude cruises. Part 11. Hydroida. (Pub-

lished June 1980.) ae ae os bes ae bs Be

MUIZON, C. DE & HENDEY, Q. B.

Late Tertiary seals of the South Atlantic Ocean. (Published June 1980.) ..

Ross, G. J. B. see SHAUGHNESSY, P. D.

SHAUGHNESSY, P. D. & Ross, G. J. B.

Records of the subantarctic fur seal (Arctocephalus tropicalis) from South Africa

with notes on its biology and some observations of captive animals.

(Published May 1980.) eh ae ae

Page

155

293

321

NEW GENERIC NAMES PROPOSED IN THIS VOLUME

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Haliana Day, 1980 a 215

Homiphoca de Muizon & Hendey, 1980 .. 3 oh ae ae ah ie 94

Unicapella Dingle, 1980

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Fiscuer, 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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Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.

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TureLe, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Ja: SCHULTZE, L. Zoologische

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ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 82 _ Band

June 1980 Junie

Part 1 Deel

MARINE SANTONIAN AND CAMPANIAN

OSTRACODS FROM A BOREHOLE AT

RICHARDS BAY, ZULULAND

R. V. DINGLE

Cape Town Kaapstad

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MARINE SANTONIAN AND CAMPANIAN OSTRACODS FROM A

BOREHOLE AT RICHARDS BAY, ZULULAND

R. V. DINGLE

Department of Geology, University of Cape Town

(With 33 figures and 4 tables)

LMS. accepted 20 December 1979]

ABSTRACT

36 species, representing 17 genera of ostracods are recorded from Santonian and

Campanian rocks penetrated by borehole BH-9 near Richards Bay, Zululand. 19 of the

species are new, 6 are left in open nomenclature, whilst the remaining 11 species have been

previously recorded from Santonian rocks at Umzamba in Transkei. One new genus,

Unicapella, is erected. The new species are: Cytherelloidea newtoni, C. griesbachi, C. contorta,

Paracypris zululandensis, Bythocypris richardsbayensis, Bairdoppilata andersoni, Amphicytherura

zululandensis, Brachycythere sicarius, Unicapella sacsi, U. reticulata, Cythereis klingeri,

Haughtonileberis vanhoepeni, Hermanites kennedyi, Oertliella pennata, O. africana, Trachyleberis

zululandensis, T. minima, Rayneria nealei, and Gibberleberis elongata.

Population studies reveal five distinctive ostracod assemblages, each dominated by one or

two species, that can be related to the following sedimentary environments (dominant taxa in

parenthesis). In ascending order these are: 1. shallow, restricted water, high energy (Brachy-

cythere longicaudata); 2. shallow, restricted water, low energy (B. longicaudata and Haughtoni-

leberis haughtoni); 3. shallow, open water, low energy (Cythereis klingeri); 4. moderate depth,

open water, low energy (Bairdoppilata andersoni); 5. deep, open water, low energy (B. andersoni

and various Cytherellids). Assemblages 1-3 are Santonian in age, whilst 4-5 correspond

approximately to Campanian I and II, respectively.

CONTENTS

PAGE

Introduction . 1

Summary of the stratigraphy of borehole BH_9, Richards Bay 3

Systematic descriptions . : : : 3

Discussion A : ‘ : : F : : i ¢ . 60

Palaeoecology . ; : ; - . 60

Ostracod assemblages 1- 5 : : : : : . 60

Sedimentary environments . : : : : 05

Biostratigraphy : : : 2 5 p : ee O,

Acknowledgements . : 5 5 ; : : j : . 68

References F ‘ : 2 : ‘ : : 2 : . 69

INTRODUCTION

Although Upper Cretaceous sediments occur extensively on the continental

shelf round southern Africa, onshore exposures of Santonian and Campanian

rocks are confined to small areas along the south-east coast (Dingle 1978). The

best exposures occur in the cliffs at Umzamba (Transkei) and round the shores

Ann. S. Afr. Mus. 82 (1), 1980: 1-70, 33 figs, 4 tables.

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

of False Bay (Zululand), whilst other small outcrops and excavations are known

from Igoda (eastern Cape), Itongazi, and Durban. A further locality has

recently been provided by a borehole at Richards Bay on the Zululand coast

(Maud & Orr 1975) (Fig. 1) where the preservation of fossils is so good that

many of the molluscs have retained their original aragonitic mineralogy and

nacreous sheen. In a region where deep Tertiary weathering has decalcified

many of the natural outcrops, the material from this borehole provides an

excellent opportunity to study small and delicate fossils.

Generally speaking, microfossils from the Upper Cretaceous rocks of

south-east Africa have received little attention. Ostracods are one group which

are poorly documented, with only three publications to date, all dealing with

Santonian assemblages from the lower part of the Umzamba section (Chapman

1904, 1923; Dingle 1969). In providing a regionally applicable biostratigraphic

framework, the recently completed thorough revision of ammonite taxonomy

and zonation by Kennedy & Klinger (1975 et seq.) and Forster (1975) has

removed one of the obstacles to further study of the less age-diagnostic micro-

False Bay

»*

x d

Cretaceous

j sediments

Lebombo

volcanics

Durban

RICHARDS BAY HARBOUR

“~Itongazi

-Umzamba

Fig. 1. Location of BH9, Richards Bay, Zululand. Other Santonian-Campanian localities

are indicated: Umzamba, Itongazi, Durban, and False Bay.

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 3

fossil taxa, and the author is currently undertaking a comprehensive investiga-

tion of the Barremian to Maastrichtian ostracod assemblages from south-east

Africa. The present paper, in describing material from the well-preserved

section in the Richards Bay borehole, will form a stratigraphic and ecologic

standard for comparison with the other less complete, and widely scattered

Senonian sequences.

SUMMARY OF THE STRATIGRAPHY OF BOREHOLE BH-9,

RICHARDS BAY

The borehole is situated 14 km west of Richards Bay township (Fig. 1)

(shown as Borehole W by Maud & Orr 1975, fig. 1), and passed through 159 m

of Upper Cretaceous sediment before entering granitic basement gneiss. 60 m at

the top of the core were weathered and unfossiliferous, and only the section 82 m

to 159 m was available for micropalaeontological analysis by the author.

Lithologically, the section studied consisted of monotonous dark olive-coloured

shelly silts and silty sands, with small pebbles and clay clasts at some levels near

the base. Pyrite and plant debris form minor constituents.

Klinger & Kennedy (1977) have described the ammonites from the BH-9

borehole and recognized three well-defined ammonite assemblages, which they

were able to compare with those from Umzamba, elsewhere in Zululand, and

Madagascar. On the basis of these faunal comparisons the lower part of the

borehole (159-114 m) is allocated a middle and upper Santonian age, whilst the

upper part (114-65 m) belongs to the lower Campanian. Local assemblages

allow a subdivision of the Campanian sequence into two faunas (Campanian I

and II) with the boundary placed at about 90 m. The succession appears to be

complete.

Thirty-three cored samples were available for microfossil analyses between

82,03 m and 150 m, and these gave good coverage from the base o* the borehole

to the lower part of the Campanian II (Fig. 32, Table 1). Each sample consisted

of half a core, about 25 mm radius and about 40 mm in length. Ostracods were

extracted by the normal washing techniques and are illustrated herein by

SEM photographs taken on a Cambridge Instruments Stereoscan 180 of the

Electron Microscopy Unit at the University of Cape Town. Specimens were

mounted on double-sided sellotape and were gold/palladium coated.

SYSTEMATIC DESCRIPTIONS

The ciassification used here is based mostly on the Ostracod Treatise

(Moore 1961), with various additions necessitated by recent work. Morpho-

logical terms have been supplemented by those introduced to cover features

visible at high magnifications (e.g. Sylvester-Bradley & Benson 1971).

TABLE 1

Vertical distribution of ostracod species in the BH9 borehole.

Figures on the left are depths of samples beneath surface in metres. The stratigraphy is based on the ammonite studies of Klinger & Kennedy (1977).

Ostracods

pyvpnoyad vjjadvo2uy

1sops pyjadpaiuy

ipauuay sajupuidayy

WV ‘ds Stuagajojsax

pupo1afo 0)/a1]142O

D140]U0I vaployjasayIaD

Sisuapupjnjnz bansayldorydup

DUlUI Sldaga]AYyods J,

sisuapupjnynz Sldagajdyov. 7,

Juadaoyuva SiiagajiuojYysno Fy

1y9DqSald3 vaplojjasayjaD

Vv “ds 27/2111420

DIDBUO]A S149G2]AIGQID

pjouuad 0112111490

Dplun} vansayido1ydup

DIvIINS DUIOpuoT

J ‘ds pyjasayid a

SIJISSYf SlAagajiuojYysnv FT

Sisuapquipziun vaployjasoyidD

Sisuadogspanyo1a S1Addooy ag

¢€ ‘ds pyjasayidD

1u0S4apubd Div 1ddopalng

1asuly siasayidD

SnuDILALfo S119g2]42gq1D

Sisuapubjopuod asaysdodyoDAg

SNIADIIS AdBYIAIAYIDA

la]Dau DidaUudDYy

py “ds pjjasayidD

7 ds DyjasayidD

1U0]YSnNdY SlAagajuojYysno

Sisuapupjnjnz siaddovivg

Sisuapquipzuin siuddopavg

DIDpPNDIIBUO] asayIAAYyoDA_E

1U0]Mau DaplojjasayjiaD

SisualaysuvA} SiasayjaD

ANNALS OF THE SOUTH AFRICAN MUSEUM

Borehole

depth

82,0

* * * *

87,0

88,0

89,0

92,3

*k Ok *

97,5

100,0

102,2

106,0

110,0

115,2

115,9

118,2

119,8

120,2

123,0

123,5

124,0

125,0

125,5

127,8

* ke ok

* * *

CAMPANIAN

SANTONIAN

x & + *

eee fer eS

22 Yb) eo ee

* ok ke ke

* oe * * * KF F KK HK HK KH K

* * &£ €

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 3)

Abbreviations

RV _ right valve

LV left valve

MPC marginal pore canals

SCT subcentral tubercle

ATE anterior terminal element

PTE posterior terminal element

ME _ median element

AM anterior margin

PM posterior margin

DM dorsal margin

VM ventral margin

NPC _ normal pore canals

MA_ marginal areas

Subclass OSTRACODA Latreille, 1806

Order PODOCOPIDA Miller, 1894

Suborder PLATYCOPINA Sars, 1866

Family Cytherellidae Sars, 1866

Members of this family constitute one of the main elements of the ostracod

fauna (Fig. 32). They are important in the lowermost few metres of the borehole

(up to 21 %) where the genus Cytherelloidea is well represented, and in the upper

half (up to 28 %) where Cytherella occurs abundantly. Between 152 m and 123 m

(assemblages 2-3) the family forms only a minor component (<10%) of the

population.

Not only does the family occur in relatively large numbers, but it is also

diverse (5 species and 4 morphotypes recognized).

Genus Cytherella Jones, 1849

Four morphotypes have been distinguished in BH-9. These probably

represent separate species, but will not be designated until further samples from

the Santonian/Campanian of Zululand have been studied. All four are found

throughout the borehole, but none is common below the Santonian/Campanian

boundary.

Cytherella Sp. 1

Fig. 2A—C

RV and LV differ considerably in shape. In LV the VM and DM are

straight and the PM asymmetrically curved, with a pronounced dorsal slope.

In the RV the DM is ‘pitched’, but not arched. In both valves the posterior ends

carry puntae with stellate outlines.

Range

Santonian to Campanian JJ, but only common in Campanian II.

6 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 2. Cytherella. A. Cytherella sp. 1, SAM—-K5567, BH9 82,03 m, LV. B. Cytherella sp. 1,

SAM-K5568, BH9 82,03 m, RV. C. Cytherella sp. 1, SAM—K5568, BH9 82,03 m, detail of

ornamentation posterior part of LV. D. Cytherella sp. 2, SAM—K5569, BH9 82,03 m, R.V

E. Cytherella sp. 3, SAM—K5570, BH9 82,03 m, RV. F. Cytherella sp. 4, SAM—K5571, BH9

82,03 m, RV.

Scale bars: C = 10p, others = 100y.

MARINE SANTONIAN AND CAMPANIAN OSTRACODS i

Cytherella Sp. 2

Fig. 2D

This form has an oval outline and a compressed anterior region when seen

in dorsal view. The DM is arched and the VM gently convex.

Range

Santonian to Campanian II, but only common in Campanian [-II.

Cytherella Sp. 3

Fig. 2E

An elongate form with a waisted lateral outline. AM symmetrically rounded,

PM asymmetrically curved.

Range

Santonian to Campanian II, but only common in Campanian I-Il.

Cytherella Sp. 4

Fig. 2F

This morphotype resembles C. Sp. 2, but differs in having a moderately

inflated posterior area and a slightly less arched DM.

Range

Santonian to Campanian II.

Genus Cytherelloidea Alexander, 1929

Cytherelloidea umzambaensis Dingle, 1969

Figs 3A, 4A-B

?Cytherella williamsoniana Jones 1849, Chapman, 1904: 236.

Cytherelloidea umzambaensis Dingle, 1969: 351-353, fig. 3.

This species occurs in small numbers (maximum 9 %) throughout the bore-

hole, but is slightly more abundant and consistently present in the Campanian

part. It is clearly an environmentally-tolerant species, with a preference for

moderate water depths (assemblage 4). In its type section (the Santonian of

Umzamba) it is moderately abundant (5-12 %).

Range

Santonian to Campanian II.

Cytherelloidea cf. C. gardeni Dingle, 1971

Cytherelloidea delicata sp. nov. Dingle, 1969: 353-354, fig. 4.

Cytherelloidea gardeni nom. nov. Dingle, 1971a: 353.

One specimen, possibly referable to this delicately reticulate species, was

found in the Santonian part of the borehole (sample at 139,8 m).

Range

Santonian.

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

umzambaensis riesbachi

A Bo 3

Cc newtoni D contorta

CY THERELLOIDEA

Fig. 3. Sketches of Cytherelloidea species right values with positive features shaded.

A. C. umzambaensis Dingle, 1969, BH9 120,22 m. B. C. griesbachi sp. nov., holotype, SAM—

K5575, BH9 88,39 m. C. C. newtoni sp. nov., holotype, SAM-—K5574, BH9 120,22 m.

D. C. contorta sp. nov., holotype SAM—K 5576, BH9 110,0 m.

Scale bars all 100p.

Cytherelloidea newtoni sp. nov.

Figs 3C, 4C

Derivation of name

In honour of Dr R. Bullen-Newton for his contributions to the geological

knowledge of Zululand.

Holotype

SAM-K5574. RV, BH-9, 120,22 m, Richards Bay, Santonian.

Dimensions

Length 0,46 mm, height 0,28 mm.

Diagnosis

Species with a right-angle-shaped ridge in the posteroventral part of the

valve.

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 9

Fig. 4. Cytherelloidea. A. C. umzambaensis Dingle, 1969, SAM—K5572, Umzamba cliff, LV.

B. C. umzambaensis Dingle, 1969, SAM—K5573, BH9 106,0 m, LV. C. C. newtoni sp. nov.,

holotype, SAM-K5574, BH9 120,22 m, RV. D. C. griesbachi sp. nov., holotype, SAM—K5575,

BH9 88,39 m, RV. E. C. contorta sp. nov., holotype, SAM—K5576, BH9 110,0 m, RV.

Scale bars all 100z.

10 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

External features. In lateral view AM symmetrically rounded, PM

asymmetrically rounded with a gently sloping posterodorsal part. Greatest

height lies over the posterodorsal angle, giving the valve a hump-backed outline.

VM nearly straight. Surface bears several short, rounded ridges; an anterior

ridge with a scalloped posterior edge, a short, high ventrolateral ridge, a right-

angle-shaped ridge in a posteroventral/posteromedian location, and a sinuous

dorsolateral ridge which terminates anteriorly in a median expansion. All these

ridges are separated, giving the surface a rather knobbly appearance.

No internal features seen.

Remarks

The humped dorsal margin and distinctive rib disposition give this species

easily recognizable characters. It is restricted to the lower (Santonian) part of

the borehole, suggesting that it was environmentally bound to shallow-water

environments. C. newtoni has not been found at Umzamba.

Range

Santonian.

Cytherelloidea griesbachi sp. nov.

Figs 3B, 4D

Derivation of name

In honour of Dr C. L. Griesbach for his contribution to the geological

knowledge of the Cretaceous rocks of south-east Africa.

Holotype

SAM-K5575, RV, BH-9, 88,39 m, Richards Bay, Campanian II.

Dimensions

Length 0,51 mm, height 0,30 mm.

Diagnosis

Species with three longitudinal ridges, the ventromedian one being

upturned posteriorly.

Description

External features. In lateral view, AM symmetrically rounded, PM

asymmetrically and bluntly pointed, being ventrally truncated and gently

sloping dorsally. Highest part of the valve at about mid-length. Surface bears

three longitudinal ridges. The dorsal ridge is slightly sinuous, and joins a wide,

raised anterior area which does not really deserve the description of a ridge.

The median ridge is short and confined to the posterior third, whilst the ventral

ridge is upturned posteriorly, and is separate from the anterior raised area. The

posterior part of the valve is compressed, and there is a conspicuous, flat

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 11

posteroventral area. Several small pustules occur in the posterodorsal part of

the valve surface. No internal features seen.

Remarks

The rib disposition of C. griesbachi is similar to that of C. umzambaensis,

but the two differ significantly in general outline (C. umzambaensis is more

elongate) in the shape of the ridges in the posteroventral area, and in their

posterodorsal outlines (Fig. 3). C. griesbachi is rare in the borehole and first

appears just below the base of the Campanian. One specimen, possibly referable

to this species, has been found at Umzamba.

Range

Uppermost Santonian to Campanian II.

Cytherelloidea contorta sp. nov.

Figs 3D, 4E

Derivation of name

Reference to its rather unusual shape and rib disposition.

Holotype

SAM-K5576, RV, BH-9, 110 m, Richards Bay, Campanian I.

Dimensions

Length 0,50 mm, height 0,28 mm

Diagnosis

Species with a strongly asymmetric and narrow posterolateral outline, a

marginal rim, and subdued ventrolateral and dorsolateral ridges. There is no

ridge.

Description

External features. In lateral view, rounded AM, but a curiously ‘contorted’

posterior outline. The dorsal part slopes gently down, whilst the ventral part is

truncated. Highest part of the valve lies in the anterior third. Ventral outline

concave, DM short and straight. Surface smooth, and ornamented with short,

subdued ridges. There is a narrow marginal rim that extends from the anterior

cardinal angle, to the midpoint of the PM. A short, rounded and rather indistinct

ventrolateral ridge, highest at its posterior end, lies at about midlength, and

there is a sinuous dorsal ridge which has swellings at its posterior and anterior

ends. There is no median ridge.

Internal features not seen.

Remarks

This unusually shaped species occurs in only one sample (110 m,

Campanian I), but is very distinctive in its shape and rib disposition by which

it is easily distinguished from the other species in the borehole. No representa-

12 ANNALS OF THE SOUTH AFRICAN MUSEUM

tives of C. contorta have been recorded from Umzamba. Its closest relative is

C. westaustraliensis Bate, 1972, from the Santonian and Campanian of western

Australia. The two species have similar ornamentation, but differ in their

posterodorsal outlines and size (the Australian species is more evenly rounded

posteriorly, and much larger, typically about 0,90 mm in length).

Range

Campanian I.

Suborder PODOCOPINA Sars, 1866

Superfamily BAIRDIACEA Sars, 1888

Family Bairdiidae Sars, 1888

Genus Bairdoppilata Coryell, Sample & Jennings, 1935

Bairdoppilata andersoni sp. nov.

Fig SA-F

Derivation of name

In honour of Dr W. Anderson for his contribution to Zululand stratigraphy.

Holotype

SAM-K5577, RV, BH-9, 88,39 m, Richards Bay, Campanian II

Paratypes

SAM-K5578, LV, BH-9, 88,39 m, Richards Bay, Campanian II

SAM-K5579, RV, BH-9, 88,76 m, Richards Bay, Campanian II

SAM-K5580, LV, BH-9, 88,76 m, Richards Bay, Campanian II

SAM-K5581, Carapace, BH-9, 88,39 m, Richards Bay, Campanian II

Dimensions

length mm heightmm width mm

K-5577 1,45 0,80

K-5578 1,50 0,90

K-5579 1,48 0,80

K-5580 1,49 0,95

K-5581 1,10 0,59

Diagnosis

Species in which LV and RV differ in shape especially in the lateral outlines

of the AM, PM and DM.

Description

External features. In lateral view, LV and RV differ in shape. The RV has

the classical ‘Bairdia’ outline with upturned posterior and anterior outlines. The

VM is slightly concave about midlength. The LV has a more ‘almond’ shape

with no noticeable upturn of the PM and a more rounded DM. In dorsal view

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 13

Fig. 5. Bairdoppilata andersoni sp. nov. A. Holotype, SAM—K5577, BH9 88,39 m, RV.

B. SAM-K5578, BH9 88,39 m, LV. C. SAM-K5579, BH9 88,76 m, internal RV. D. SAM-—

K5580, BH9 88,76 m, internal LV. E. SAM—K5581, BH9 88,39 m, dorsal view, carapace.

F. SAM-K5580, BH9 88,76 m, muscle scars, LV.

Scale bars: F = 100p, others = 300p.

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

the LV is more inflated. The valve surface is finely pitted all over, and possesses

numerous large normal pore canal openings.

Internal features. Hinge typical for the genus, with very fine serration on

the bars at the anterior and posterior ends of the RV flange groove. MS in LV

consist of an anteriorly opening semi-circlet of 8 spots enclosing two large and

two small spots, with a further two small spots anterodorsally. All spots are

rounded. MA moderately wide with small anterior vestibules.

Remarks

Bairdoppilata andersoni shows considerable intra-specific variation in

lateral outline and with further study it may be possible to effect further sub-

division. Its distribution in the Richards Bay borehole is remarkably clear cut—

throughout the Santonian section four specimens were recovered (assemblages

1-2), whilst in the Campanian rocks it is the dominant ostracod—up to 40% of

the total fauna. Its appearance in large numbers is sudden and coincides with

the Santonian/Campanian boundary where it marks the establishment of open

ocean low energy conditions (assemblage 4). Towards the top of the section, as

the water deepens further, its total numbers continue to increase although with

the influx of a large population of Platycopina it still maintains about a 40 per

cent level of the total population.

Bairdoppilata sp. (probably andersoni) has been recorded from Umzamba

as rare, single valves.

Range

Santonian to Campanian II.

Genus Bythocypris Brady, 1880

Bythocypris richardsbayensis sp. nov.

Fig. 6A-E

Derivation of name

Locality of type.

Holotype

SAM-K5582, LV, BH-9, 88,39 m, Richards Bay, Campanian II.

Paratypes

SAM-K5583, RV, BH-9, 106,00 m, Richards Bay, Campanian I

SAM-K5584, Carapace, BH-9, 88,5 m, Richards Bay, Campanian II

Dimensions

length mm height mm

K5582 0,55 0,28

K5583 0,48 0,26

K5584 RV 0,47 0,24

K5584 LV 0,48 0,28

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 15

Fig. 6. Bythocypris richardsbayensis sp. nov. A. Holotype, SAM-—K5582, BH9 88,39 m,

internal LV. B. SAM-K5583, BH9 106,0 m, internal RV. C. SAM-K5584, BH9 88,5 m,

carapace, RV. D. SAM—K5583, BH9 106,0 m, muscle scars, RV. E. SAM-K5583, BH9

106,0 m, hinge, RV.

Scale bars: D-E = 30y, others = 100p.

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

Diagnosis

An elliptical species with reticulate ornamentation on the inside surface of

the outer lamella.

Description

External features. In lateral view, LV larger than RV, overlaps around

entire margin, but especially along VM. RV : AM asymmetrically rounded,

truncated dorsally, PM rounded, greatest height just in front of midlength.

LV : AM symmetrically rounded, and PM asymmetrically rounded, truncated

dorsally, greatest height at midlength. Surface smooth.

Internal features. Hinge: a smooth bar in RV fits into a shallow groove in

the LV. The bar is downward projecting in lateral view, and is thickest at its

anterior end. MS consist of rosette of four petal-like ventral scars and an

E-—W-elongated dorsal scar. MA: anterior is wide with a prominent, anteriorly

‘tapering vestibule; posterior is narrow. MPC: 12 fine, straight canals radiate

from the apex of the anterior vestibule; 6 very short fine posterior canals. The

inside surface of valve is ornamented with reticulation of polygonal fossae

(mostly hexagonal) and narrow muri.

Remarks

In shape and hinge, B. richardsbayensis is very similar to B. howchiniana

Chapman, 1917, as illustrated by Neale (1975) from the Santonian of Australia.

The MS of the latter are reported to consist of a cluster of 4 large scars, whereas

the illustration (plate 5 fig. 2b of Neale 1975) suggests that 5 are present—in

which case the similarity with B. richardsbayensis would be even closer. The

main outstanding difference is the presence of reticulation in the inner surface

of the outer lamella in the South African species.

B. richardsbayensis occurs sporadically throughout the Richards Bay

borehole but it is most abundant (12%) and consistent in the Campanian (I-II)

and lower Santonian parts, and is absent from the uppermost Santonian rocks

(assemblage 3).

Range

Santonian to Campanian II.

Superfamily CYPRIDACEA Baird, 1845

Family Paracyprididae Sars, 1923

Genus Paracypris Sars, 1866

Of the non-Cytheracean types, this genus is the most consistently present

throughout the borehole. It never occurs in large numbers, however (maximum

of about 11°), and is most abundant in the Santonian section (i.e. prefers the

shallower water, higher energy environments of assemblages 1-3).

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 17

Paracypris umzambaensis Dingle, 1969

Figs 7A-C, 9A

Macrocypris simplex Chapman 1898, Chapman, 1904: 233, pl. 29 (fig. 22).

Paracypris ? umzambaensis Dingle, 1969: 354-356, fig. 5.

This species was provisionally placed in Paracypris by Dingle (1969)

because the MS had not been seen at that time. SEM pictures obtained during

the present study confirm the original assignment. Two varieties of the species

occur—a slender variety with a markedly pointed PM, and a fatter variety with

a less acuminate posterior outline. Both varieties occur in the Richards Bay

borehole and at Umzamba. The species occurs throughout the borehole section

but is most abundant in the Santonian (up to 11 %) and rare (up to 3 %) in the

Campanian. The fatter variety is similar in outline to Paracypris sp. nov.

described by Neale (1975) from the Santonian of western Australia.

Range

Santonian to Campanian II.

Paracypris zululandensis sp. nov.

Figs 7D-G, 9B

Derivation of name

Location of borehole.

Holotype

SAM-K5588, RV, BH-9, 106,00 m, Richards Bay, Campanian I

Paratypes

SAM-K5589, LV, BH-9, 110,00 m, Richards Bay, Campanian I

SAM-K5590, LV, BH-9, 110,00 m, Richards Bay, Campanian I

SAM-K5591, RV, BH-9, 110,00 m, Richards Bay, Campanian I

Dimensions

length mm height mm

K5588 0,73 0,37

K5589 0,75 0,38

K5590 0,78 0,42

K5591 0,73 0,35

Diagnosis

A plump species with a tapering, rounded posterior outline.

Description

External features. In lateral view AM is rounded, slightly extended in the

RV, PM tapers but is distinctly round and curves upwards at the posteroventral

corner. DM strongly arched and VM convex in LV and slightly concave in RV.

Surface smooth.

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 7. Paracypris. A. P. umzambaensis Dingle, 1969, SAM—K5585, Umzamba cliff, slim

variety, LV. B. P. umzambaensis Dingle, 1969, SAM—K 5586, BH9 88,39 m, plump variety, LV.

C. P. umzambaensis Dingle, 1969, SAM-K5587, BH9 118,22 m, plump variety, RV.

D. P. zululandensis sp. nov., holotype, SAM-K5588, BH9 106,0 m, internal RV. E. P. zulu-

landensis, sp. nov., SAM—K5589, BH9 110,0 m, internal LV. F. P. zululandensis, sp. Nov.,

SAM-K5590, BH9 110,0 m, LV. G. P. zululandensis sp. nov., SAM—K5591, BH9 110,0 m, RV.

Scale bars all 100z.

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 19

Internal features. Hinge consists of a thick bar (RV) and a corresponding

groove and dorsal lip (LV). MS: cluster of five scars and a large, elliptical dorsal

scar, MA wide, with a wide anterior vestibule that extends almost to the anterior

margin. Seven anterior and five posterior thin, indistinct MPC.

Remarks

P. zululandensis is difficult to distinguish from the plumper varieties of

P. umzambaensis. The two differ on: degree of acumination of PM, ventral

outline (straighter in umzambaensis), more acutely arched DM of umzambaensis,

and details of MS pattern (Fig. 9).

P. zululandensis occurs in small numbers (up to 5%) sporadically through-

out the Richards Bay borehole. It is most consistently present just above the

Santonian/Campanian boundary (assemblage 4).

Range

Santonian to Campanian II.

Superfamily CyTHERACEA Baird, 1850

Except in the uppermost part of the borehole (above 88 m, assemblage 5)

Cytheracea are numerically the largest part of the ostracod population. They

are the dominant component in the Santonian section (up to 90% of the fauna),

but steadily decrease in importance upwards (between 50% and 38% in the

Campanian section).

Family Xestoleberididae Sars, 1928

Xestoleberis sp. A

Fig. 8

This species occurs in minor amounts (up to 7%) in the upper part of the

borehole (i.e. Campanian II and the upper part of Campanian I). It has a

Fig. 8. Xestoleberis sp. A. A. SAM-K5592, BH9 88,5 m, LV. B. SAM-K5593, BH9 88,5 m,

dorsal view, carapace.

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

slightly convex ventral outline in lateral view and a weakly developed antero-

ventral marginal rim. In dorsal view the LV overlaps the RV prominently at the

anterior end, and the carapace is inflated medianly. Until the borehole specimens

can be compared with other material from Zululand (the genus is well repre-

sented in Campanian and Maastrichtian samples) it is not advisable to attempt

to formally designate the new species.

Illustrated specimens

SAM-K5592, LV, BH-9, 88,50 m, Richards Bay, Campanian IT

SAM-K5593, Carapace, BH—9, 88,50 m, Richards Bay, Campanian II

Dimensions

length mm height mm width mm

K5592 0,37 0,23

K5593 0,38 0,24

Range

Campanian I-II.

Family Cytherideidae Sars, 1925

Genus Pondoina Dingle, 1969

Pondoina sulcata Dingle, 1969

Fig. 9C

Pondoina sulcata Dingle, 1969: 356-358, fig. 6.

This species occurs as a rare component in the middle part of the Santonian

section of BH-9, where the specimens are poorly preserved. In contrast it

occurs moderately abundantly (10-15 %) throughout the Santonian sequence at

Umzamba. A sketch of MS is included here to supplement the original

descriptions.

Range

Santonian.

Family Schizocytheridae Mandelstam, 1960

Genus Amphicytherura Butler & Jones, 1957

Amphicytherura tumida Dingle, 1969

Fig. 1OA-F

Amphicytherura (A.) tumida Dingle, 1969: 368-370, fig. 13.

This species was first recorded in the Santonian rocks at Umzamba, where

it forms a minor element of the fauna (2-10 %). In the Richards Bay borehole it

occurs in small numbers (up to 10%) and its range extends from the lower part

of the sequence into Campanian I. It is, however, absent from the uppermost

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 21

eee

“o

umzambaensis zululandensis

Pondoina sulcata

Fig. 9. Muscle scars. A. Paracypris umzambaensis Dingle, 1969, Umzamba cliff, LV.

B. Paracypris zululandensis sp. nov., SAM—K5589, BH9 110,0 m, LV. C. Pondoina sulcata

Dingle, 1969, Umzamba cliff, LV.

Scale bars all 30y.

Santonian, and across the Santonian/Campanian boundary. Some of the

specimens in BH—9 are very well preserved and are illustrated to supplement the

original descriptions given by Dingle (1969).

Range

Santonian to Campanian I.

Amphicytherura zululandensis sp. nov.

Fig. 11A-G

Derivation of name

Locality of borehole.

Holotype

SAM-K5598, RV, BH-9, 97,5 m, Richards Bay, Campanian I

22 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pave

a rans CePA

Fig. 10. Amphicytherura tumida Dingle, 1969. A. SAM-K5594, BH9 125,0 m, RV. B. SAM—

K5595, BH9 139,8 m, LV. C. SAM-K5596, BH9 128,0 m, internal RV. D. SAM—-K5597,

BH9 128,0 m, internal LV. E. SAM-K5597, BH9 128,0 m, hinge LV. F. SAM-K5596, BH9

128,0 m, hinge RV.

Scale bars: E-F = 30p, others = 100z.

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 23

Fig. 11. Amphicytherura zululandensis sp. nov. A. Holotype, SAM—K5598, BH9 97,5 m, RV.

B. SAM-K5599, BH9 97,5 m, LV. C. SAM-K5600, BH9 110,0 m, internal RV. D. SAM-

K5601, BH9 110,0 m, internal LV. E. SAM-K5600, BH9 110,0 m, hinge RV. F. SAM-K5601,

BH9 110,0 m, muscle scars, LV. G. SAM-K5601, BH9 110,0 m, hinge LV.

Scale bars: E-G = 30y, others = 100z.

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

Paratypes

SAM-K5599, LV, BH-9, 97,5 m, Richards Bay, Campanian I

SAM-K5600, RV, BH-9, 110,00 m, Richards Bay, Campanian I

SAM-K5601, LV, BH-9, 110,0 m, Richards Bay, Campanian I

Dimensions

length mm height mm

K5598 0,53 0,30

K5599 0,42 0,26

K5600 0,42 0,23

K5601 0,55 0,30

Diagnosis

Species with prominent posteroventral alae, the under surfaces of which are

coarsely reticulate.

Description

External features. Quadrate in lateral view. AM symmetrically rounded,

PM bluntly pointed. DM nearly straight, VM strongly upturned posteriorly, but

outline is obscured by a small ala which has a pointed, right-angled posterior

corner. The undersides of the alae are coarsely reticulate. Lateral valve surface

is uneven, with three main irregular, raised areas and a north-south median

depression. A prominent eye spot lies on a short, curved ridge, and there is a

prominent hump below the posterior cardinal angle. Overall, the surface is

smooth, finely punctate (especially in the posterior half), locally faintly reticulate

and bears a few, scattered pustules.

Internal features. Hinge strongly schizodont with the PTE of the RV

undercut posteriorly. MS consist of a curved posterior row of four rounded

scars, and two rounded frontal scars. Internal openings of the NPC are promi-

nent. MA wide with 7-9 anterior and 5 posterior MPC.

Remarks

A. zululandensis is closely allied to A. tumida, but differs in lacking a median

lateral ridge, a SCT, and in having more prominent posteroventral alae. There

are also subtle differences in the hinge structure (e.g. the anterior part of the ME

in LV is elongate and sloping in A. tumida, whilst it is shorter and not inclined in

A. zululandensis).

A. zululandensis occurs in small numbers in the Campanian I section of the

borehole, where its range overlaps with that of A. tumida over a short section

(4-5 m).

Range

Campanian I.

oT eee ooo ———— —

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 25

Family Brachycytheridae Puri, 1954

Genus Brachycythere Alexander, 1933

Numerically this is the most important genus in the borehole. In the lower

(Santonian) half, it commonly forms more than 30 per cent of the total ostracod

population, and near the base reaches 56 per cent at one horizon (Fig. 32).

A rapid decline takes place in the upper part of the Santonian (between 120

and 124 m), above which it reaches >20 per cent at one horizon only (at the

Campanian J-II boundary). A similar distribution was recorded from the

Santonian rocks at Umzamba where the genus made up >30 per cent of the

population in all four samples studied by Dingle (1969).

Brachycythere longicaudata (Chapman, 1904)

Figs 12A—C, 13A—D

Cytheridea longicaudata Chapman, 1904: 234, pl. 29 (fig. 21). Howe & Laurencich, 1958: 279.

Cythere ?drupacea Jones 1884, Chapman, 1904: 234.

Brachycythere longicaudata (Chapman), Dingle, 1969: 358-361, fig. 7.

Dingle (1969) erected a neotype on topotypic material from the Santonian

section at Umzamba, where B. Jongicaudata is the dominant element in the fauna

(about 30%). It is found in two morphotypes, elongate and squat, in addition

to which there is considerable small-scale variation within the population;

S. l-_ fet ae

72 YE is

Fig. 12. Muscle scars of Brachycythere longicaudata (Chapman, 1904). A. SAM-—K5605, BH9

88,39 m, RV. B. Umzamba cliff, RV. C. Umzamba cliff, LV.

Scale bars all 30z.

ornamentation, shape of the ventrolateral overhang, outline of the AM and PM,

and even in the MS pattern.

B. longicaudata occurs throughout BH-9 (mid Santonian to Campanian II),

and, as at Umzamba, it is the dominant element in the lower half of the

Santonian section (up to 50%) and an important element in the rest of the

succession (up 12% in Campanian II and 22% in the Upper Santonian). This

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

large, robust species was obviously tolerant of varying environmental conditions,

as indicated by its re-establishment after the faunal break that marks the local

Santonian/Campanian boundary.

Range

Santonian to Campanian II.

Fig. 13. Brachycythere. A. B. longicaudata (Chapman, 1904), SAM—K5602, BH9 88,39 m, LV.

B. B. longicaudata (Chapman, 1904), SAM-K5603, BH9 106,2 m, RV. C. B. longicaudata

(Chapman, 1904), SAM—K5604, BH9 92,27 m, LV. D. B. longicaudata (Chapman, 1904),

SAM-K5605, BH9 88,39 m, internal RV. E. B. sicarius sp. nov., SAM—K5608, BH9 97,5 m,

muscle scars, RV.

Scale bars: E = 30u, B-C = 100n, A, D = 300z.

MARINE SANTONIAN AND CAMPANIAN OSTRACODS Di

Brachycythere pondolandensis Dingle, 1969

Brachycythere pondolandensis Dingle, 1969: 361-362, fig. 8.

This small, rounded, and relatively ornate species is rare in the Richards Bay

borehole, where it reaches a maximum of about 8 per cent at the top of its range.

It is confined to rocks of Santonian age. It is also rare in the Santonian of

Umzamba, its type locality, and may prove to be a good marker fossil for

Santonian rocks in south-east Africa. Its preservation is never good, suggesting

that it may have readily been destroyed in sections that are not completely fresh.

Range

Santonian.

Brachycythere sicarius sp. nov.

Figs 13F, 14A—F

Derivation of name

From the Latin sica (a dagger or dirk), reference to dagger-like alae spines.

Holotype

SAM-K 5606, RV, BH-9, 89,0 m, Richards Bay, Campanian II

Paratypes

SAM-K 5607, LV, BH-9, 88,39 m, Richards Bay, Campanian II

SAM-K 5608, RV, BH-9, 97,5 m, Richards Bay, Campanian I

SAM-K5609, LV, BH-9, 97,5 m, Richards Bay, Campanian I

Dimensions

length mm height mm

K5606 0,85 0,40

K5607 0,82 0,42

K5608 0,79 0,40

K5609 0,80 0,42

Diagnosis

Species with pronounced, ventrolateral, keeled alae, coarse circular fossae

on the median lateral surface, and coarse anterior and posterior marginal spines.

Description

External features. Triangular in lateral view. Rounded AM and pointed PM.

DM and VM straight, converging posteriorly, but ventral outline is convex

because of ventrolateral overhang by an ala with a keel-like outer edge. The AM

carries several coarse, stubby spines, whilst the posteroventral area typically

carries two or three posteriorly pointing dagger-like spines. A further large spine

occurs at the posterior end of the ala, although this is retained only on well-

preserved specimens. The lateral surface is covered all over with rounded fossae

which are largest along the inner edge of the alae. There is a prominent anterior

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

Nvoes Og

Nees

Fig. 14. Brachycythere sicarius sp. nov. A. Holotype, SAM—K5606, BH9 89,0 m, RV.

B. SAM-K5607, BH9 88,39 m, LV. C. SAM—-K5608, BH9 97,5 m, internal RV. D. SAM-

K5609, BH9 97,5 m, internal LV. E. SAM-K5608, BH9 97,5 m, hinge RV. F. SAM—K5609,

BH9 97,5 m, hinge LV.

Scale bars: E-F = 30p, others = 100y.

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 29

cardinal angle, on which is situated a rounded, subdued eye spot, with a distinct

post-ocular sulcus (especially in the LV).

Internal features. Hemiamphidont hinge. In RV, ATE is a dentate peg, with

a denticulate dorsal surface, and PTE is an elongate dentate bar with six teeth.

The ME consists of a crenulate bar with an anterior, inclined rectangular peg.

MS consist of four elongate posterior scars and a hooked anterior scar. MA wide

with 20-23 long, slender, irregular anterior and 12 posterior MPC. The latter are

concentrated on the ventral side.

Remarks

Although typical, well-preserved specimens of B. sicarius are easily

separated from examples of B. longicaudata by their spinose, alate outline and

punctate ornamentation, poorly-preserved specimens may be confused with the

more angular varieties of the latter species. A somewhat similar species,

B. angulata Grekoff has been recorded from the Coniacian—Campanian of

West Africa (e.g. Apostolescu 1961). It possesses short keeled alae, but lacks the

coarse reticulation and spines of B. sicarius.

B. sicarius occurs in small numbers (maximum of 11%) throughout BH-9,

but is more abundant in the Campanian section. Rare specimens occur in the

Santonian of Umzamba, although they were not recorded by Dingle (1969) as a

separate species in his original description of the fauna.

A length/height scattergram (Fig. 15) effectively separates the three impor-

tant species of Brachycythere found in the Richards Bay/Umzamba areas.

B. sicarius, whilst having approximately the same length/height ratio as

B. longicaudata, its closest relative, is a smaller species.

Range

Santonian to Campanian II.

se X<

0,8 ee Ne

7 N

EE Ss

BRACHYCY THERE Be x

0,7 ae 2 Se

Ci e \

4 e \

~ Bea e e \

= A NY

a=) 3G \

206 $e }

a 4

. : Tag

PEsh. OE Cee ee een OnE ae

‘ e ee dd

0,5 (Gre) ee an” Dy ak ei ye eee at longicaudata

e e al

. . ‘s Ba aa eaat

sicarius _._. a caer,

nee was ondolandensis

mm wes SA ‘, P

0,4 Ye > I

"0,6 ice +09 1,0 1,1 1,2 1,3 1,4 1,5

4 are length

Fig. 15. Scattergram of length v. height for adults of species of Brachycythere

in BH9 Richards Bay.

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Trachyleberididae Sylvester-Bradley, 1948

Unicapella gen. nov.

Type species

Unicapella sacsi sp. nov.

Derivation of name

To commemorate the 150th anniversary of the founding of the University

of Cape Town in 1829: Uni... Cap... + ella.

Diagnosis

A blind, reticulate Trachyleberid with a prominent SCT and spinose/nodose

dorsal and ventral margins.

Remarks

Unicapella belongs to a group of recently-described deep water (>1 000 m)

genera which includes Abyssocythere, Paleoabyssocythere and Atlanticythere

(Benson 1971, 1977). Apparently living in water depths of no more than 500 m,

Unicapella possibly forms a link between the deep-water forms and their

shallow-water ancestors. The group seems to have arisen in the Upper

Cretaceous, with its main centre of development in the South Atlantic/South-

western Indian Ocean areas.

Unicapella is closest to Paleoabyssocythere, which is first recorded from the

Campanian of the Brazilian continental margin (Benson 1977), but it differs in

ornamentation (Paleoabyssocythere is foveolate, not reticulate, and has a

massive anterior marginal rim), in size (Unicapella is typically less than three-

quarters the length of Paleoabyssocythere), and hingement (Unicapella is

holamphidont, Paleoabyssocythere is lobodont).

At the type locality (Richards Bay borehole) the new genus (2 spp) is

restricted to Campanian II strata, but at least one other species occurs in

Maastrichtian sediments farther north (personal unpublished data).

Unicapella sacsi sp. nov.

Fig. 16A-G

Derivation of name

To commemorate the 150th anniversary of the founding of the South

African College Schools (SACS) in 1829, from which developed the University

of Cape Town.

Holotype

SAM-K5610, RV, BH-9, 88,39 m, Richards Bay, Campanian II

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 31

Fig. 16. Unicapella sacsi gen. et sp. nov. A. Holotype, SAM-K5610, BH9 88,39 m, RV.

B. SAM-K5611, BH9 87,0 m, LV. C. SAM-K5612, BH9 87,0 m, internal RV. D. Holotype,

SAM-K5610, BH9 88,39 m, detail ornamentation anterior part RV. E. SAM-K5612, BH9

87,0 m hinge RV. F. SAM-K5613, BH9 87,0 m, dorsal view, carapace. G. Holotype,

SAM-K5610, BH9 88,39 m, detail anterior part RV.

Scale bars: D = 10yu, E = 30p, others = 100p.

32 ANNALS OF THE SOUTH AFRICAN MUSEUM

Paratypes

SAM-K5611, LV, BH-9, 87,00 m, Richards Bay, Campanian II

SAM-K5612, RV, BH-9, 87,00 m, Richards Bay, Campanian II

SAM-K5613, Carapace, BH-9, 87,00 m, Richards Bay, Campanian II

Dimensions

length mm heightmm width mm

K5610 0,60 0,33

K5611 0,59 0,31

K5612 0,52 0,27

K5613 0,55 0,27

Diagnosis

Species with large, domed SCT and fine surface reticulation.

Description

External features. In lateral view, spinose, symmetrically rounded AM, and

asymmetrically acuminate PM. DM straight, with prominent anterior cardinal

angles that are notched in the RV, and have a projecting ‘ear’ in the LV.

Surface reticulate and spinose. Several short, large, perforate spines lie along the

DM, and an ill-defined ventrolateral clava. Large bullae occur in posterodorsal

and posteroventral positions. The SCT is large, bulbous hemispherical and

smooth, and forms the widest part of the valve in dorsal view. Valve surface

finely reticulate with a few conjunctive pustules, particularly behind the SCT.

The depressed anterior area is coarsely reticulate with delicate secondary retic-

ulation. Secondary reticulation also developed in the posterior part of the LV.

Fossa muri are finely crenulate, both in primary and secondary reticulation.

“Internal features. Hinge amphidont with a narrow terminally widening

groove above the ME in RV. In RV, ATE subdivided, PTE smooth or weakly

subdivided. MS not seen. MA wide with twelve long, slender, straight anterior,

and eight posterior MPC.

Remarks

U. sacsi is close to Paleoabyssocythere cretacea Benson, 1977. In addition

to differing in aspects related to generic status, Unicapella sacsi has less massive

spines and bullae and a less symmetric anterior outline. The known range of

P. cretacea is Santonian/Campanian.

Range

Campanian II.

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 33

Unicapella reticulata sp. nov.

Fig. 17A-B

Derivation of name

Reference to coarsely reticulate ornamentation.

Holotype

SAM-K5614, RV, BH-9, 88,76 m, Richards Bay, Campanian II.

Dimensions

length mm height mm

K5614 0,65 0,32

Diagnosis

Coarsely reticulate species with spinose muri.

Description

External features. In lateral view, spinose, rounded AM and acuminate PM.

DM straight, sloping downwards posteriorly. VM slightly convex. Stout spines

Fig. 17. Unicapella reticulata gen. et sp. nov.

A. Holotype, SAM—K 5614, BH9 88,76 m, RV.

B. Holotype, SAM-K5614, BH9 88,76 m,

detail anterior part RV.

Scale bars: B = 30u, A = 100p.

34 ANNALS OF THE SOUTH AFRICAN MUSEUM

occur along the DM and on a well-defined ventrolateral clava. Large spinose

bullae occur in posterodorsal and posteroventral positions. Large, reticulate

SCT. Surface coarsely reticulate, with polygonal fossae with small disjunctive

and conjunctive spines and pores, and inward pointing mural spines. Depressed

anterior area contains very large fossae. There is a marked step at the anterior

cardinal angle. Only RV were available for study and no internal features were

seen.

Remarks

Despite the limited amount of material available, the distinction between

U. reticulata and U. sacsi can be confidently made: the SCT are reticulate and

smooth, respectively; and surface reticulation is coarse and spinose in the former,

and fine with crenulated muri in the latter. U. reticulata shows some similarities

to species allocated to the family Pennyellidae by Neale (1975), with species of

both Agulhasina and Pennyella having spinose muri and reticulate sub-central

areas.

U. reticulata is very rare in the Richards Bay borehole (three valves).

Range

Campanian II.

Genus Cythereis Jones, 1849

Cythereis transkeiensis Dingle, 1969

Fig. 18A

Cythereis ?ornatissima Reuss, 1846 var. reticulata Jones & Hinde, 1890, Chapman, 1904:

234 (no illustration).

Cythereis transkeiensis Dingle, 1969: 377-378, fig. 8.

Three poorly-preserved specimens of C. transkeiensis were found in the

basal sample (159 m) of the Richards Bay borehole. In contrast, this species

occurs in small numbers (c. 5°) throughout the Santonian part of the sequence

at Umzamba that was examined by Dingle (1969). MS are recorded for the first

time and show a typical Cythereis arrangement of four posterior, and one

U-shape frontal scar.

Range

Santonian.

Cythereis klingeri sp. nov.

Figs 18B-F, 19A-F

Derivation of name

In honour of Dr H. C. Klinger for his recent contributions to knowledge of

the stratigraphy and palaeontology of the Cretaceous rocks of Zululand.

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 35

Fig. 18. Cythereis. A. C. transkeiensis Dingle, 1969, SAM-—K5615, BH9 158,0 m, LV.

B. C. klingeri sp. nov. SAM—K 5622, BH9 157,0 m, LV. C. C. klingeri sp. nov. SAM—KS5623,

BH9 124,0 m, LV. D. C. klingeri sp. nov. SAM—K5624, BH9 118,22 m, LV. E. C. klingeri

sp. nov. SAM-K5621, BH9 115,9 m, LV. F. C. klingeri sp. nov. SAM—K5618, BH9 82,03 m,

muscle scars, RV.

Scale bars: F = 30u, others = 100p.

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

Holotype

SAM-K5616, RV, BH-9, 82,03 m, Richards Bay, Campanian II

Paratypes

SAM-K5617, LV, BH-9, 82,03 m, Richards Bay, Campanian II

SAM-K5618, RV, BH-9, 82,03 m, Richards Bay, Campanian II

SAM-K5619, LV, BH-9, 157,0 m, Richards Bay, Santonian

SAM-K5620, RV, BH-9, 157,0 m, Richards Bay, Santonian

SAM-K5621, LV, BH-9, 115,9 m, Richards Bay, Santonian

SAM-K5622, LV, BH-9, 157,0 m, Richards Bay, Santonian

SAM-K5623, LV, BH-9, 124,0 m, Richards Bay, Santonian

SAM-K5624, LV, BH-9, 118,22 m, Richards Bay, Santonian

Dimensions

length mm height mm

K5616 0,78 0,40

K5617 0,79 0,44

K5618 0,80 0,42

K5619 0,74 0,41

K5620 0,75 0,38

K5621 0,58 0,33

K5622 0,83 0,46

K5623 0,75 0,45

K5624 0,58 0,32

Diagnosis

Reticulate species with a prominent, flared, dorsolateral ridge.

Description

External features. In lateral view, AM rounded, DM and VM nearly

Straight, outline tapering posteriorly. Anterior cardinal angle is prominent,

particularly in the LV where it projects well above the dorsal margin. Lateral

surface bears three longitudinal ridges which are not interconnected and do not

project to the AM. The dorsal ridge is deflected anteromedianly, is flared, and

is subdivided about midlength, particularly in the LV. Weakly developed SCT,

across which median ridge is deflected anteroventrally. Whole surface strongly

reticulate, with ovate to rounded fossae and numerous mural pore cones.

Internal features. Holamphidont hinge. In RV, ATE consists of a dentate,

weakly subdivided peg with a thumb-like posterior projection, PTE is a massive,

smooth tooth, with an anterior depression to accommodate a projection of the

LV ME. Anterior socket of LV is flanked by two large teeth: one at the end of

the ME and the other anterior of it. The posterior part of the LV ME is stepped.

MS set in a shallow depression and consist of four posterior scars and a hook-

shaped frontal scar. There is a prominent fulchral point. MA moderately wide

with twelve anterior and six posterior MPC.

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 37

Fig. 19. Cythereis klingeri sp. nov. A. Holotype, SAM-—K5616, BH9 82,03 m, RV. B. SAM-—

K5617, BH9 82,03 m, LV. C. SAM-K5618, BH9 82,03 m, internal RV. D. SAM-K5619, BH9

157,0 m, internal LV. E. SAM-K5620, BH9 157,0 m, hinge RV. F. SAM-K5619, BH9

157,0 m, hinge LV.

Scale bars: E-F = 30p, others = 100p.

38 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

Exterior architecture of C. klingeri resembles that of C. transkeiensis, but

differs in being more strongly reticulate, in having a flared dorsal ridge, a

tapering posterior outline, and in details of hinge and MS. In the upper part of

the Santonian C. klingeri is the dominant ostracod (up to 46 % of the population)

in what is thought to have been a quiet, shallow-water environment. A good

deal of intra-specific variation is displayed in strength of reticulation and

prominence of the lateral ridges.

Range

Santonian to Campanian II.

Genus Haughtonileberis Dingle, 1969

This genus was created to accommodate local forms that are closely

related to the Dumontina/Curfsina/Hazelina Complex, but which differ con-

sistently in details of outline, ornamentation, hingement and marginal areas.

HAUGHTONILEBERIS gy a a

radiatu

haughtoni seas cu Paissilie

i Oo :

Sele

height

length we bs

Fig. 20. Scattergram of length v. height for adults of species of Haughtonileberis in southern

Africa.

Four species have been assigned to Haughtonileberis to date, all from

southern Africa (a fifth remains to be described): three from the Santonian/

Campanian at Umzamba/Richards Bay (H. haughtoni, H. fissilis, H. vanhoepeni),

and one from the Eocene of the JC—1 borehole offshore Natal (H. radiatus).

H. haughtoni and H. fissilis are well represented in the Santonian to Campanian

rocks of the Richards Bay borehole, but both species show a good deal of intra-

specific variation, possibly environmentally influenced. H. vanhoepeni is rarer

(up to 15%) and is confined to Campanian II rocks.

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 39

The opportunity is taken here to supplement the original descriptions of

H. haughtoni and H. fissilis with SEM pictures.

Figure 20 shows a scattergram of length/height ratios of the four species of

Haughtonileberis so far described.

Haughtonileberis haughtoni Dingle, 1969

Fig. 21A—E.

Haughtonileberis haughtoni Dingle, 1969: 372-373, fig. 15.

Two morphotypes of this species have been recognized both at Umzamba

and Richards Bay: an elongate form which ranges across the Santonian/

Campanian boundary and occurs throughout most of the borehole, and a

squatter variety which is confined to the Santonian (shallower water) strata.

In the squat variety the dorsolateral ridge tends to be foreshortened, and the

median ridge wide throughout its length. In contrast, in some of the more

elongate forms the posterior part of the median ridge is reduced to a narrow rib.

The holotype appears to be an intermediate form.

H. haughtoni is one of the most important species in the Richards Bay

borehole, and it is particularly abundant in the Santonian sequence (assem-

blage 2) where it reaches 55 per cent and is typically >30 per cent. Its numbers

decline sharply in the more open water environments of assemblages 3 and 4

(10-15%) and the species disappears just after the establishment of the deep-

water assemblage 5.

Range

Santonian to Campanian II.

Haughtonileberis fissilis Dingle, 1969

Fig. 22A-B

Haughtonileberis fissilis Dingle, 1969: 374-375, fig. 16. Dingle, 1976: 59, fig. 3(48).

H. fissilis occurs sporadically throughout the Richards Bay borehole, and

is most abundant (up to 19%) in the Campanian I, moderate-water depth

environments. It is recorded from only four horizons in the Santonian section

(top assemblage 1, and assemblage 2), where it reaches 14 per cent. On this

evidence it would appear that H. fissilis favoured somewhat deeper water than

its close relative H. haughtoni.

As with H. haughtoni, H. fissilis is known to occur in two morphotypes:

a slim finely reticulate type, which is very rare in the Richards Bay borehole, and

a squatter, more coarsely reticulate variety. The latter variety has not been

found in the Umzamba rocks, where H. fissilis is a minor component of the

ostracod fauna (<10°%).

Range

Santonian to Campanian II.

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

ase. .

. * “

ae ”"

Cre ry

Fig. 21. Haughtonileberis haughtoni Dingle, 1969. A. SAM—K5625, BH9 157,0 m, elongate

variety, LV. B. SAM—K5626, BH9 157,0 m, elongate variety, RV. C. SAM-—K5627, BH9

92,27 m, elongate variety, LV. D. SAM-—K5628, BH9 120,22 m, squat variety, RV. E. SAM-—

K5629, BH9 120,22 m, squat variety, LV.

Scale bars all 100p.

| MARINE SANTONIAN AND CAMPANIAN OSTRACODS 41

Fig. 22. A. Haughtonileberis fissilis Dingle, 1969, SAM-—K5630, BH9 115,19 m, LV.

B. Haughtonileberis fissilis Dingle, 1969, SAM—K5631, BH9 100,0 m, RV. C. Hermanites

kennedyi sp. nov. SAM-K5638, BH9 88,39 m, papillate solum, RV. D. Hermanites kennedyi

sp. nov. SAM-K5642, BH9 88,5 m, ATE dorsal view, LV. E. Hermanites kennedyi sp. nov.

SAM-K 5643, BH9 88,39 m, ATE dorsal view, RV. F. Hermanites kennedyi sp. nov. SAM-—

K5639, BH9 88,39 m, muscle scars, LV. G. Hermanites kennedyi sp. nov. SAM-K5641, BH9

88,76 m, dorsal view, carapace. H. Haughtonileberis vanhoepeni sp. nov. SAM—K5636, BH9

106,0 m, dorsal view, carapace.

Scale bars: C = 3n, F = 10n, D-E = 30p, others = 100z.

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

Haughtonileberis vanhoepeni sp. nov.

Figs 22H, 23A-F

Derivation of name

In honour of Dr E. C. N. van Hoepen for his studies of Zululand Cretaceous

ammonites.

Holotype

SAM-K 5632, RV, BH-9, 100,00 m, Richards Bay, Campanian I.

Paratypes

SAM-K 5633, RV, BH-9, 92,27 m, Richards Bay, Campanian I

SAM-K5634, RV, BH-9, 106,00 m, Richards Bay, Campanian I

SAM-K 5635, LV, BH-9, 106,00 m, Richards Bay, Campanian I

SAM-K 5636, Carapace, BH-9, 106,00 m, Richards Bay, Campanian I

Dimensions

length mm height mm width mm

K5632 0,46 0,22

K5633 0,46 0,26

K5634 0,41 0,21

K5635 0,41 0,23

K5636 0,41 0,19

Diagnosis

Small species with an inclined median rib which is joined to the ventro-

lateral rib by a loop.

Description

External features. Small species. In lateral view, AM broadly and asym-

metrically rounded, PM bluntly pointed. VM straight, DM slopes posteriorly.

Surface bears three narrow longitudinal ribs: dorsal rib is slightly curved,

projects beyond the DM and is strongly deflected medianly at its posterior end

where it forms a prominent posterior cardinal angle; median rib inclined,

projects anterior of the SCT, but does not join the anterior rim, curves ventrally

at its posterior end to join the strongly inclined ventrolateral rib which runs

from the anteroventral corner of the valve when it joins the anterior rim.

Small, prominent eye spots lie on the anterior rim at the cardinal angles in both

valves. Intercostal areas smooth, with a few small ridges and minor reticulation.

Several prominent pustules. In dorsal view the posterior ends of the dorsal ribs

appear as small ears.

Internal features. Hinge holamphidont. In RV ATE is a stepped tooth,

and PTE a small, undivided smooth tooth. In LV ME is a smooth bar which

thickens terminally with a downward inclined tooth at its anterior end. MA

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 43

Fig. 23. Haughtonileberis vanhoepeni sp. nov. A. Holotype, SAM—K5632, BH9 100,0 m, RV.

B. SAM-K5633, BH9 92,22 m, RV. C. SAM-K5634, BH9 106,0 m, internal RV. D. SAM-—

K5635, BH9 106,0 m, internal LV. E. SAM-K5634, BH9 106,0 m, hinge RV. F. SAM-K5635,

BH9 106,0 m, hinge LV.

Scale bars: E-F = 30y, others = 100p.

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

moderately wide with small vestibules. MPC short and straight: 20-25 anteriorly,

c. 10 posteriorly. MS not clearly seen.

Remarks

Differs from the two other Upper Cretaceous species of the genus in ridge

disposition, intercostal ornamentation, and in being smaller. In ornamentation

and rib arrangement, H. vanhoepeni is intermediate between the more elongate

varieties of H. fissilis and an unnamed species from the Campanian at False Bay.

Preservation of this species is typically very good—specimens are clean and

semi-translucent to translucent, and mostly in the form of carapaces.

H. vanhoepeni is restricted to assemblage 4 where it locally reaches 15 per cent

of the population.

Range

Uppermost Santonian to Campanian I.

Genus Hermanites Puri, 1955

Hermanites kennedyi sp. nov.

Figs 22C-G, 24A—F

Derivation of name

In honour of Dr W. J. Kennedy for his contribution to the palaeontology

and stratigraphy of the Cretaceous rocks of south-east Africa.

Holotype

SAM-K 5637, LV, BH-9, 82,03 m, Richards Bay, Campanian II

Paratypes

SAM-K 5638, RV, BH-9, 88,39 m, Richards Bay, Campanian II

SAM-K5639, LV, BH-9, 88,39 m, Richards Bay, Campanian II

SAM-K5640, RV, BH-9, 88,76 m, Richards Bay, Campanian II

SAM-K5641, Carapace, BH-9, 88,76 m, Richards Bay, Campanian II

SAM-K5642, LV, BH-9, 88,5 m, Richards Bay, Campanian II

SAM-K 5643, RV, BH-9, 88,39 m, Richards Bay, Campanian II

Dimensions

length mm height mm width mm

K5637 0,71 0,38

K5638 0,72 0,37

K5639 0,71 0,36

K5640 0,72 0,38 ;

K5641 0,74 0,43

K5642 0,75 0,20

K5643 0,74 0,20

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 45

Fig. 24. Hermanites kennedyi sp. nov. A. Holotype, SAM-K5637, BH9 82,03 m, LV.

B. SAM-K5638, BH9 88,39 m, RV. C. SAM-K5639, BH9 88,39 m, internal LV. D. SAM-—

K5640, BH9 88,76 m, internal RV. E. SAM-K5639, BH9 88,39 m, hinge LV. F. SAM—K5640,

BH9 88,76 m, hinge RV.

Scale bars: E-F = 30p, others = 100p.

46 ANNALS OF THE SOUTH AFRICAN MUSEUM

Diagnosis

Species with prominent alae and a ventrally inclined posterior outline in

the RV lateral view.

Description

External features. In lateral view, AM weakly spinose and symmetrically

rounded in RV, slightly asymmetric in LV, DM and VM converge slightly

posteriorly. PM coarsely spinose, bluntly acuminate, ventrally inclined in RV.

There are three longitudinal lateral ribs, the dorsal and ventral of which flare

posteriorly, the latter prominently. The median ridge is short and lies posterior

to, and detached from the prominent SCT. There is a prominent eye spot which

is not attached to the strong AM rim. Surface coarsely reticulate with occasional

conjunctive pustules. Fossae are polygonal, with rounded corners and papillate

sola. In dorsal view there is a prominent projection of the LV valve margin

across the hingeline anterior of the eye spot.

Internal features. Hinge holamphidont, with smooth peg-like terminal

elements in the RV, the PTE being weakly subdivided. Posterior groove of the

RV ME is seen to be weakly crenulate in well-preserved specimens. MS consist

of four posterior spots, the dorsal one of which is a deformed V-shape, and two

anterior scars—an asymmetric U and a rounded dorsal scar. MA very narrow

with twenty-three anterior and eight posterior MPC.

Remarks

H. kennedyi resembles H. sagitta from the Santonian-Campanian of

Australia (Bate 1972). They both possess an arrow-head outline in dorsal view,

and flared ventral and dorsal ribs, but A. kennedyi differs in possessing a short

median lateral rib, is more spinose posteriorly, has a different posterior outline

in RV, has a finely crenulate ME in the hinge, and possesses a more complex MS

pattern. H. volans from the Santonian of Australia (Neale 1975) possesses

prominent alae, but differs from the new species in not having a prominent

dorsal rib, nor a median rib behind the SCT.

In the Richards Bay borehole, H. kennedyi is restricted to the Campanian

section where it locally forms an important element in the fauna (c. 16%).

It appears to have a preference for the deeper-water environment.

Range

Campanian II.

Genus Oertliella Pokorny, 1964

Oertliella pennata sp. nov.

Fig. 26A—E

Acanthocythere ? aff. A. horridula (Bosquet, 1854), Dingle, 1969: 378-380, fig. 19.

Derivation of name

From the Latin penna (feather), reference to feather-like posterodorsal

spine in LV.

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 47

Holotype

SAM-K 5644, LV, BH-9, 115,9 m, Richards Bay, Santonian.

Paratypes

SAM-K 5645, RV, BH-9, 115,9 m, Richards Bay, Santonian.

SAM-K 5646, LV, BH-9, 120,22 m, Richards Bay, Santonian.

Dimensions

length mm height mm

K5644 0,68 0,36

K5645 0,71 0,34

K5646 0,75 0,38

OERTLIELLA

exquisita /¥Y~"—--~.,

Voy)

0,4

ay bsiatelnne

_- africana

0,6 length 0,8

Fig. 25. Scattergram of length v. height for adults of Oertliella

pennata, O. africana, O. sp. A and O. exquisita Bate.

Diagnosis

Elongate species with a large, serrated spine at the posterior cardinal angle

of the LV.

Description

External features. In lateral view, elongate quadrate outline with spinose,

rounded AM and bluntly acuminate PM. DM and VM straight, tapering

slightly posteriorly. Surface coarsely reticulate with numerous conjunctive and

disjunctive spines. Fossae are polygonal (3 and 4 sided) and the main muri

radiate from the prominent SCT, which has a large verrucose spine. The most

48 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 26. Oertliella. A. O. pennata sp. nov., holotype, SAM—K5644, BH9 115,9 m, LV.

B. O. pennata sp. nov. SAM-K5645, BH9 115,9 m, RV. C. O. pennata, sp. nov. SAM-K5646,

BH9 120,22 m, internal LV. D. O. pennata sp. nov. SAM—K5646, BH9 120,22 m, muscle

scars, LV. E. O. pennata sp. nov. SAM-K5646, BH9. 120,22 m, hinge, LV. F. O. sp. A,

SAM-K 5647, BH9 115,9 m, LV. G. O. africana sp. nov. SAM—K5650, BH9 82,03 m, muscle

scars, LV.

Scale bars: D-E, G = 30p, others = 100p.

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 49

prominent spines lie along the DM, a posteriorly upswinging ventrolateral clava,

and at the antero- and posteroventral corners of the valve. The spine at the

posterodorsal angle of the LV is particularly prominent—it is flattened and

serrated at its distal end, giving it a feather-like appearance. Eye spot is large,

hemispherical, situated at the anterior cardinal angle.

Internal features. Hinge holamphidont. MS situated in and on the outer

edge of a prominent pit. Four posterior scars lie in a line, the lower two are

wedge-shaped and lie close together, the 3rd scar is rounded, and lies in the

centre of the pit, whilst the dorsal scar is small and rounded. A further small

rounded impression lies posterior of this group. Anterior scar is V-shaped, but

partially subdivided. MA narrow with twenty paired, short, straight, anterior

and twelve posterior MPC.

Remarks

O. pennata differs from its relation O. africana, also from the Richards Bay

borehole, in details of ornamentation, hinge, and in possessing a prominent eye

spot. It is closest to O. exquisita from the Campanian of western Australia (Bate

1972) but the two can be separated on subtle differences in shape, reticulation

of the posterior parts of the valves, the ‘feather-like’ spine of pennata, the

generally smaller size of the African species, as well as its greater length/height

ratio:

Average values: RV—1,81 exquisita 2,05 pennata

LV—1,80 exquisita 1,98 pennata

O. pennata occurs in small numbers (up to 15%) throughout assemblages 3

and 4, and it is found at one horizon in assemblage 2.

Range

Santonian to Campanian I].

Oertliella africana sp. nov.

Figs 26G, 27A-E

Derivation of name

Location of type.

Holotype

SAM-K5648, LV, BH-9, 82,03 m, Richards Bay, Campanian II

Paratypes

SAM-K 5649, RV, BH-9, 87,0 m, Richards Bay, Campanian II

SAM-K 5650, LV, BH-9, 82,03 m, Richards Bay, Campanian II

SAM-K5651, RV, BH-9, 97,5 m, Richards Bay, Campanian I

50 ANNALS OF THE SOUTH AFRICAN MUSEUM

Dimensions

length mm_ height mm

K5648 0,76 0,37

K5649 0,63 0,27

K5650 0,88 0,38

K5651 0,65 0,32

Diagnosis

An elongate species with prominent posteroventral and dorsal spines, and

a weak eye spot.

Description

External features. In lateral view, elongate quadrate. AM rounded, PM

bluntly acuminate—both coarsely spinose, particularly the posteroventral area,

which carries a long pointed spine. DM straight, VM slightly convex in the LV.

Surface reticulate with small, rounded fossae. Muri not symmetrically arranged.

SCT not well developed, often only represented by a smooth, slightly raised area.

DM carries a few large, flared, blade-like spines, particularly in the LV, where

the anterior cardinal angle is surmounted by a very prominent spine. There is an

upswinging ventrolateral clava with several verrucose spines. The eye spot is

weakly developed: in the LV it is usually present below the anterior cardinal

angle, and in the RV it is frequently not distinguishable.

Internal features. Hinge paramphidont, with a large projection in front of

the ATE in the LV. MS consists of four elongate posterior scars and a V-shaped

anterior scar set in a shallow depression. The anterior scar has three 1aised

portions, suggesting incipient subdivision. MA narrow, with fifteen short

straight anterior and seven posterior MPC.

Remarks

Its closest relative is O. pennata from which it differs in possessing a weak

eye spot, rounded fossae, lacking a radiating pattern to its reticulation, and

having only a weak SCT. The two species also have different hinge structures and

slightly different length/height ratios (Fig. 25).

O. africana is most abundant (up to 4%) in assemblage 5 (quiet deep water).

Range

Campanian I and II.

Oertliella sp. A

Fig. 26F

Remarks

Three specimens resembling O. africana, but lacking the coarse spines.

Fossae are more quadrate and the valve outline somewhat plumper.

Range

Campanian I and II.

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 51

gt =. ye Are

“a

of or te

Fig. 27. Oertliella africana sp. nov. A. Holotype, SAM-K5648, BH9 82,03 m, LV. B. SAM-—

K5649, BH9 87,0 m, RV. C. SAM-K5650, BH9 82,03 m, internal LV. D. SAM-K5651, BH9

97,5 m, internal RV. E. SAM-K5651, BH9 97,5 m, hinge RV. F. SAM-K5650, BH9 82,03 m,

hinge LV.

Scale bars: E-F = 30p, others = 100y.

$2 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Trachyleberis Brady, 1898

Two species are herein assigned to the genus on the basis of their shape,

hinge, and ornamentation. Whilst both are somewhat less spinose than the

genotype, their features are commensurate with some recent interpretations of

the genus (e.g. Neale 1975). The lack of distinct posteromedian spine alignment

suggests that neither belongs in the genus Actinocythereis, and whilst both

(7. minima sp. nov. in particular) are very similar exteriorly to the genotype of

Matronella Damotte, 1974, the apparently smooth ATE and PTE of the

Zululand specimens, and the fact that MS were not seen, precludes positive

assignment to this genus.

Trachyleberis zululandensis sp. nov.

Fig. 28A—C

Derivation of name

Locality of type.

Holotype

SAM-K5652, LV, BH-9, 82,03 m, Richards Bay, Campanian II

Paratype

SAM-K5653, RV, BH-9, 82,03 m, Richards Bay, Campanian II

Dimensions

length mm height mm

K5652 0,82 0,40

K5653 0,78 0,36

Diagnosis

A blind species with a triangular lateral outline, and a prominent SCT.

Description

External features. Valve laterally compressed. In lateral view, spinose,

rounded AM, triangular PM. DM and VM straight, converging posteriorly to

produce a triangular lateral outline. Surface ornamented with stout spines, many

of which are perforate. Castellated spines occur along the DM, and in the

median region. There is a large spinose SCT. Under low magnification the valve

surface appears smooth, but under high magnification is seen to be finely

reticulated with hair-like muri.

Internal features. Hinge holamphidont. MS not seen. MA narrow, with

at least 20 anterior MPC.

Remarks

T. zululandensis is similar to T. schizospinosa Dingle from the Maastrichtian

of the Agulhas Bank (Dingle 19715) and T. raynerae Neale from the Santonian

of Australia (Neale 1975). It differs from the latter in not possessing an eye spot,

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 53

Fig. 28. A. Trachyleberis zululandensis sp. nov., holotype, SAM—K5652, BH9 82,03 m, RV.

B. Trachyleberis zululandensis sp. nov. SAM—K5653, BH9 82,03 m, internal RV. C. Trachy-

leberis zululandensis sp. nov. SAM—K5653, BH9 82,03 m, hinge RV. D. Trachyleberis minima

sp. nov., holotype, SAM—K 5654, BH9 110,0 m, RV. E. Rayneria nealei sp. nov. SAM-—K5656,

BH9 115,9 m, dorsal view, carapace. F. Rayneria nealei sp. nov. SAM—K5656, BH9 115,9 m,

detail anterior end, dorsal view, carapace.

Scale bars: C, F = 30pu, others = 100p.

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

in having less elongate spines, but in possessing a more prominent SCT. Its

closest relative is 7. schizospinosa, but this species is plumper, more quadrate in

outline, and possesses small eye spots.

T. zululandensis is rare in the Richards Bay borehole (only two samples

contain it, up to 3%) and it is confined to assemblages 4 and S. Its slightly

greater abundance in sample 82,03 and its blindness suggest that it preferred

deep-water environments.

Range

Campanian I and II.

Trachyleberis minima sp. nov.

Fig. 28D

Derivation of name

Its small size.

Holotype

SAM-K5654, RV, BH-9, 110,00 m, Richards Bay

Dimensions

length mm height mm

K5654 0,54 0,25

Diagnosis

Quadrate species with a concave ventral outline in lateral view.

Description

External features. In lateral view quadrate with rounded AM and a bluntly

rounded PM, which is directed slightly posteroventrally. Both end margins bear

numerous small, short spines. DM is straight, VM is concave—posterior to

line of maximum height. The lateral surface has several short, small spines which

are confined to the DM, a short curved ventrolateral clava, and the median area

(on the prominent SCT, and in a cluster behind the SCT). Interspinal areas are

smooth, with numerous small, low, circular pustules. There is a large globular

eye spot at the anterior cardinal angle. No internal features seen.

Remarks

Placement of this small spiney species is provisional because no internal

features were seen. The VM concavity and downturned PM make it easily

recognized. T. minima is rare in the Richards Bay borehole in one sample only

(at 3%) in Campanian I.

Range

Campanian I.

MARINE SANTONIAN AND CAMPANIAN OSTRACODS B))

Genus Rayneria Neale, 1975

Rayneria nealei sp. nov.

Figs 28E-F, 29A-E, 30G

Cythereis ?quadrilatera (R6mer) Chapman, 1923: 5, pl.1(5).

Derivation of name

In honour of Prof. J. W. Neale who erected the genus.

Holotype

SAM-K 5652, RV, BH-9, 139,8 m, Richards Bay, Santonian

Paratypes

SAM-K5653, LV, BH-9, 106,00 m, Richards Bay, Campanian I

SAM-K5654, RV, BH-9, 115,19 m, Richards Bay, Campanian I

SAM-K5655, LV, BH-9, 139,8 m, Richards Bay, Santonian

SAM-K5656, Carapace, BH-9, 115,9 m, Richards Bay, Santonian

SAM-K 5657, RV, BH-9, 118,22 m, Richards Bay, Santonian

Dimensions

length mm heightmm width mm

K5652 0,53 0,27

K5653 0,50 0,24

K5654 0,54 0,28

K5655 0,53 0,28

K5656 0,57 0,32

K5657 0,51 0,12

Diagnosis

Small, blind species with small SCT.

Description

External features. In lateral view, small, subquadrate ostracod. AM bears

short spines, symmetrically rounded in RV, extended ventrally in LV. PM

rounded in LV, asymmetric in RV, bears several stout spines posteroventrally.

DM hidden, outline is convex. VM concave. Surface bears three short longi-

tudinal ridges: the dorsal ridge is curved and slightly flared; the median ridge is

short, commences just anterior of the small, elongate SCT, and has a postero-

ventral deflection; the ventral ridge is convex. Intercostal areas heavily calcified,

with small raised pustules and scattered rounded fossae. Cardinal angles rounded

in LV, but in the RV the anterior cardinal angle is weakly developed. No eye

spots. There is a prominent ear-like overlap by the LV just in front of anterior

cardinal angle in dorsal view.

Internal features. Hinge holamphidont but with a stepped and posteriorly-

elongated ATE in RV. There is a small additional tooth in front of ATE in LV.

56 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 29. Rayneria nealei sp. nov. A. Holotype, SAM-K5652, BH9 139,8 m, RV. B. SAM-

K5653, BH9 106,0 m, LV. C. SAM-K5654, BH9 115,19 m, internal RV. D. SAM-K5655,

BH9 139,8 m, internal LV. E. SAM-K5654, BH9 115, 19 m, hinge RV. F. SAM-K5655, BH9

139,8 m, hinge LV.

Scale bars: E-F = 30, others = 100z.

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 57

MS not clearly seen, but they lie in a deep central pit. MA narrow, no vestibules,

up to 18 anterior and 8 posterior MPC.

Remarks

Rayneria nealei is similar to Rayneria ginginensis Neale from the Santonian

of Australia in its surface ornamentation, but differs by not having eye spots

and by possessing a small SCT.

The possession of eyespot was considered by Neale (1975: 59) to be a

generic characteristic, but it is not thought to be desirable to create a new genus

on this criterion alone, where Neale’s is otherwise mono-specific.

R. nealei is a sporadic component of assemblages 1, 2 and 3, and occurs in

small numbers in the early range of assemblage 4. It is never common and only

locally accounts for >10 per cent of the total fauna.

Range

Santonian to lower Campanian I.

Genus Gibberleberis Dingle, 1969

Gibberleberis africanus Dingle, 1969

Figs 30A-D, 31A

Gibberleberis africanus Dingle, 1969: 370-377, fig. 17.

This species was first described from the Santonian of Umzamba cliff

section, where it occurs sparingly (up to 3%). In the Richards Bay borehole it is

also restricted to Santonian rocks and again occurs in small numbers (up to

15%). It is most common in the uppermost Santonian rocks (assemblage 3).

Some of the new material is very well preserved and SEM pictures are

included to supplement the original description. MS (previously not recorded)

are rather complex, with 5 posterior scars and two anterior scars. The edges of

individual scars are serrated. Topotypic specimens are more heavily calcified

than individuals from Richards Bay.

Range

Santonian.

Gibberleberis elongata sp. nov.

Figs 30, 31B

Derivation of name

Elongate shape in lateral view.

Holotype

SAM-K5660, RV, BH-9, 124,0 m, Richards Bay, Santonian

Paratype

SAM-K5661, RV, BH-9, 124,0 m, Richards Bay, Santonian

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pe ie: :

ee ns i

5 ed, Shane

Fig. 30. A. Gibberleberis africanus Dingle, 1969, SAM-K5658, BH9 118,22 m, LV.

B. Gibberleberis africanus Dingle, 1969, SAM-K5659, BH9 118,22 m, internal LV.

C. Gibberleberis africanus Dingle, 1969, SAM-K5659, BH9 118,22 m, hinge LV.

D. Gibberleberis africanus Dingle 1969, SAM-K5659, BH9 118,22 m, muscle scars, LV.

E. Gibberleberis elongata sp. nov. Holotype, SAM—K5660, BH9 124,0 m, RV. F. Gibberleberis

elongata sp. nov. SAM-K5661, BH9 124,0 m, RV. G. Rayneria nealei sp. nov. SAM-K5657,

BH9 118,22 m, ATE and PTE of hinge, dorsal view, RV.

Scale bars: C-D, G = 30p, others = 100p.

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 59

Dimensions

length mm height mm

K5660 0,6 0,25

K5661 0,6 0,30

Diagnosis

Elongate, finely reticulate species, with straight ventral outline in lateral

view.

Description

External features. In lateral view, AM symmetrically rounded, PM asym-

metrically acuminate. Dorsal outline broadly humped, ventral outline straight,

producing a posteroventral elongation. Surface finely reticulate with, in the

upper part of the valve, main muri running longitudinally, parallel to an arched,

low, dorsal rib. This accentuates the humped appearance. Valves slightly

swollen ventrally, particularly posterior of mid-length. SCT represented by a

smooth area in the reticulate pattern. Eyespot elongate and low.

No internal features seen.

Remarks

G. elongata and G. africanus are very closely related. In addition to

differences in lateral outline, the most obvious point of reference is the difference

in disposition of the reticulation ribs (Fig. 31). In G. elongata the muri on the

GIBBERLEBERIS

EM) GES

elongata africanus

Fig. 31. Sketches of Gibberleberis species showing main rib patterns. A. G. elongata

sp. nov., holotype, SAM—K 5660, BH9 124,0 m, RV. B. G. africanus Dingle, 1969,

BH9 140,0 m, RV.

Scale bars 100p.

dorsolateral area are more numerous, concentrically arranged, and less curved,

and in the anterior area are larger.

G. elongata is rare in the Richards Bay borehole (six valves only recorded

from three horizons), and occurs in both Santonian and Campanian rocks.

Range

Santonian to Campanian II.

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

DISCUSSION

The ostracods in BH-9 are all benthic types, and thirty-six species belonging

to seventeen genera have been recognized. There are several distinctive changes

in the composition of the ostracod populations along the length of the borehole,

and it is convenient to examine these from two viewpoints: palaeoecological and

biostratigraphical.

PALAEOECOLOGY

The regional Cretaceous geology in south Zululand, in combination with

ev dence to be presented below, indicates that the sedimentary history of the

borehole is of a marine transgression across a granitic surface followed by a

progressive increase in water depth. Temporary returns to shallower water and/or

higher energy environments at intervals up the borehole may represent minor

regressive episodes. Changes in the geometry of the depositional area caused by

the progressive increase in water depth would have constantly modified

ecological factors, such as current strengths (e.g. tidal, wave-driven, and

oceanographic), salinity, food supply, and water temperature. Because such

factors would have determined the make-up of the ostracod faunas, it ought to

be possible, by their study, to monitor the changes in the sedimentary environ-

ment. Whilst, at this stage, the influences of the more subtle factors cannot be

recognized, it is possible to express the overall sedimentary history in terms of

alterations in such generalized factors as water depth, overall energy of the

environment, and access to open water (i.e. oceanic) regimes. This can be done

by the recognition of ostracod assemblages, which can be assigned to particular

sedimentary environments on the basis of species distribution and overall

population trends.

Ostracod assemblages 1-5

The vertical distribution of each ostracod species is shown in Table 1, and

various statistical data on species and population trends are shown in Figure 32.

Five ostracod types numerically dominate the faunas (>20% of total popula-

tion), either singly or in combinations, and in successive zones up the borehole

these are (maximum % based on three point means in brackets): Brachycythere

longicaudata (51%), B. longicaudata (40%) with Haughtonileberis haughtoni

(56%), Cythereis klingeri (45°), Bairdoppilata andersoni (36%), and B. andersoni

(41%), with various species of Cytherella (32%). The level at which the

dominance of these types or combinations changes is usually clear cut (1-2 m

wide) and coincides with important changes in numbers and presence/absence of

secondary and minor taxa, as well as alterations in diversity, similarity and

overall size of the entire ostracod populations (Fig. 32, Tables 2-3). Only the

change over from the B. longicaudata/H. haughtoni to the C. klingeri-dominated

faunas is transitional and takes place with a good deal of fluctuation in all

parameters over a distance of about 4 m in the borehole. The distinctive

ostracod populations in these zones have been designated ‘ostracod assemblages’

1 to 5 and are shown in Figure 32 and Tables 2-3. Most of the secondary and

turnover &

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 61

minor taxa occur in more than one assemblage (although several are confined to

only two), but six species are restricted to only one. It is possible therefore to

characterize each assemblage in terms of its dominant types (>20%) with, in

three cases, additional specialized, but relatively rare, taxa. These associations

are summarized in Table 3. At a more general level, reference to Table 1 shows

Campanian Santonian |

11 I upper middle age

(ee hance ae gs (naa l(a assemblage

population

30 stability

% 20 4 increase

¥ decrease

25 z =

> sue’ < bah Hee gh

= 1 7 - ‘“+-- a rends

: ‘ 5) oe turnover <

> o

mostly Cytheracea ;

60 Bairdoppilata higher

taxa

Bairdiacea

= Te mostly

ae al Cytherellidae Bythocypris

Roan eee

60 <

Cytherella spp. C. klingeri dominant

Z = P

ao = “Qn, = ¥ \

20 =

| — ff Saewtont

— B.pondolandensis

@ Psulcata

A.tumida

O.pennata Baws Bea other

C.griesbachi species

H.vanhoepeni

-zujulandensis

A.zululandensis __

O.africana

X.sp

H.kennedyi

U.spp

Fig. 32. Variations in the ostracod faunas of BH9, Richards Bay. Discontinuities in the various

curves have been used to delimit ostracod assemblages 1 to 5 which are described in detail in

the text and Tables 2-3. Population stability has been measured by variations in the similarity

index between sample points up the borehole (ticks on the column marked borehole depth).

Population turnover is a measure of the number of new appearances and extinctions in each

sample, and the population diversity is calculated as number of species per hundred specimens.

Note that for most of the borehole there is an inverse relationship between population

diversity and the total number of valves per sample i.e. largest populations tend to be the least

diverse. The only large deviation from this trend is in the lower part of ostracod assemblage I.

These relationships may in part, however, be influenced by the statistical bias of dealing with

small numbers of specimens. Note that in all cases, percentage of species and higher taxa

quoted in this figure and in the text are based on three-point running mean values.

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

that assemblages 1, 2 and 3 are closely related both in terms of individual minor

and secondary species (e.g. Cytherelloidea newtoni, Brachycythere pondo-

landensis, and Gibberleberis africanus) and higher taxa (Cytheracean dominated,

> 60%), and that similarly, assemblages 4 and 5 have many characteristics in

common (nine species do not extend below 115,9 m, and the faunas are mainly

Bairdiacean/Cytherellid types).

It is clear that the overriding driving force behind these population altera-

tions was environmental change because such alterations were sudden but yet

did not lead to the extinction of the dominant species. Some of the appearances

and extinctions in the minor taxa may be phylogenetic, but the close corre-

TABLE 2

Make-up of faunal assemblages.

Dominant

Secondary

Minor

rare and sporadic

B. andersoni

H. haughtoni

B. longicaudata

Cytherella spp.

B. richards-

bayensis

1 2 3

aa B. longicaudata B. longicaudata

oN H. haughtoni

a C. klingeri

is)

H. haughtoni H. haughtoni

C. klingeri C. klingeri

= B. longicaudata

oN P. umzambaensis

C. newtoni C. newtoni C. newtoni

= B. richards- B. richards-

2 bayensis bayensis

€ P. umzambaensis —_P. umzambaensis

5 P. zululandensis P. zululandensis

n R. nealei R. nealei R. nealei

iz B. sicarius B. sicarius

7) G. africanus G. africanus G. africanus

3 B. pondolandensis __B. pondolandensis

° H. fissilis

=] A. tumida

zi P. sulcata*

x O. pennata

<<]

x C. umzambaensis C. umzambaensis

3 C. transkeiensis

= H. fissilis

E G. elongata

8 O. pennata

= O. sp. A

3 C. griesbachi

4 B. sicarius

* restricted to one ostracod assemblage

T —transitional

P. umzambaensis

P. zululandensis

R. nealei

B. sicarius

H. fissilis

A. tumida

O. pennata

C. umzambaensis

C. griesbachi

O. africana

A. zululandensis*

H. vanhoepeni*

O. sp. A

T. zululandensis

T. minima*

X, sp.

C. contorta*

B. andersoni

Cytherella spp.

H. kennedyi*

P. umzambaensis

P. zululandensis

B. sicarius

H. fissilis

O. pennata

C. umzambaensis

C. griesbachi

O. africana

Unicapella spp.

X. sp.

G. elongata

O. sp. A

T. zululandensis

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 63

spondence between maxima in population turnover and assemblage boundaries,

where the assemblages are defined by dominant taxa present (Fig. 32) suggests

that most are local, ecologically-related events.

Sedimentary environments

Various lines of evidence can be used to deduce the palaeosedimentary

environments from the character of the ostracod assemblages 1 to 5. Overall

conclusions are summarized in Table 3. The two most clear-cut cases are

assemblages 1 and 5. In the former, the dominant species is B. longicaudata,

which appears in the earliest sediments immediately above the basement surface

(sample 159 m). There is a rapid build-up in total size of population from an

estimated 3 600 adults/m? at the base to a maximum of about 10 000 adults/m?

higher up, coeval with a steady influx of new species colonizing the newly-

created marine environment. The latter trend is seen on Figure 32 as a high

turnover rate (dominantly appearances). Once established, the fauna has a high

diversity (>20%) and an average similarity index value of >25% (which

indicates a moderately stable population). Corroded and blackened valves near

the base of the borehole, together with the absence of Bairdoppilata and low

numbers of Cytherella suggest that ostracod assemblage 1 inhabited a shallow

water, high energy environment with restricted access to the open ocean.

Sedimentation rates were probably fairly high, but cannot be quantified. On this

evidence, B. Jongicaudata can be regarded as a ‘hardy’ species and its temporary

resurgence at intervals high up in the borehole (especially in the upper part of

assemblage 4 and between 123 and 125,5 m) may indicate temporary returns to

somewhat higher energy environments.

Assemblage 5 consists of a mixed Bairdiacea/Cytherellidae population with

relatively rare (<20°%) Cytheracean types. The combination of Bairdoppilata

with abundant Cytherella in modern environments appears to indicate

moderately deep-water (outer continental shelf) environments (e.g. Rosenfeld &

Bein 1978). A similar conclusion for assemblage 5 is supported by the presence

of four blind Cytheracean species (Unicapella (2 spp), Oertliella africana, and

Trachyleberis zululandensis), as well as Xestoleberis sp. An interesting and

significant member of this assemblage is the. unusual genus Unicapella, whose

closest relative is Paleoabyssocythere from the Campanian of the northern Rio

Grande Rise (DSDP site 21) (Benson 1977). The latter genus was the forerunner

of the psychrospheric Cainozoic genus Abyssocythere. Benson (1977) reckons

that Paleoabyssocythere cretacea lived at depths of at least 1 000 m. Assemblage 5

certainly does not suggest depths as great as this, but does indicate that the

environment was ‘oceanic’ in the sense that there were free connections to the

palaeo South Atlantic. The relative abundance of the architecturally complex

Hermanites kennedyi, as well as the low faunal diversity (<20 species/100 valves),

but high population densities (10 400 adults/m?) and stabilities (> 30%) indicate

low energy, deep (perhaps between 200 and 500 m) conditions. One surprising

absentee is the typically deep-water ‘marker’ Krithe.

ANNALS OF THE SOUTH AFRICAN MUSEUM

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MARINE SANTONIAN AND CAMPANIAN OSTRACODS 65

Assemblages 2, 3 and 4 represent environments intermediate to 1 and 5.

Abundant Bairdoppilata, the increasing importance of Cytherella, and the

appearance of blind forms suggest that assemblage 4 marks the onset of

moderately deep-water conditions and probably represents a midshelf (100-200 m

depth) environment. The increasing specialization of the fauna in assemblage 4

is indicated by the low population diversity (<20%) coupled with a large

population density (12 000 adults/m?).

Through assemblages 2 and 3, the importance of B. longicaudata is in

progressive, if erratic decline, and in assemblage 3 the Cytheracean component

as a whole declines sharply (about 90% at the base to 60% at the top). Deep-

water elements are largely absent and the population stability is only low to

medium as the faunas constantly adjusted to deepening water and lower energy

environments. Both assemblages 2 and 3 suggest shallow water, low energy

environments, and the low population diversity in assemblage 2 (~20%)

probably indicates somewhat restricted conditions. Population stability (as

measured by averaging the similarity indices) is especially low at about 128 m

in the borehole, above which, for a short distance (about 3 m), rapid fluctuations

in C. klingeri/H. haughtoni/B. longicaudata numbers indicate a period of

population instability probably related to fluctuating current strengths caused

by sea-level movements (? minor regressions). Similar, but less pronounced

events, occur at the boundaries between assemblages 3 and 4, and 4 and 5.

Table 3 summarizes the conclusions on palaeoenvironments up the borehole.

Finally, the trends exhibited in the ostracod population as it adjusted to

changing environmental conditions over a period of about 5 m.y. can be

graphically expressed in Figure 33A. Here, total Cytheracea—Cytherellidae—

Bairdiacea/Cypridacea components are plotted on a triangular diagram. Each

of the ostracod assemblages occupies a well-defined field although areas of

assemblages 1, 2 and 3 overlap. An interesting feature of this diagram is that it

emphasizes the trend followed by the initial, colonizing fauna (assemblage 1) as

it rapidly develops towards Cytheracean dominance, which persists, and is more

firmly established, through the duration of assemblage 2. The field occupied by

assemblage 3 partially overlaps with that of assemblage 1, but can be dis-

tinguished from it by the presence of several moderate water depth elements in

the Cytheracean component (e.g. O. pennata, A. tumida). Fields for assem-

blages 4 and 5 are well separated, with the latter showing a progressive trend

towards the Bairdiacea/Cypridacea—Cytherellidae base line as the water depth

increases. Despite the closeness of parts of fields 4 and 5 to field 1, distinction

can easily be made by the make-up of their respective Bairdiacea/Cypridacea

components: to the ‘north’ of the divide this component is a Paracypris plus

subordinate Bythocypris population, whereas to the ‘south’ it is a Bairdoppilata

plus subordinate Paracypris and Bythocypris population.

Translating these population fields into palaeoenvironments Figure 33B is

generated. At present, its applicability to other areas has not been tested and it

can only be considered relevant to the BH—9 borehole.

66 ANNALS OF THE SOUTH AFRICAN MUSEUM

CYTHERACEA

BAIRDIACEA & CYTHERELLIDAE

CYPRIDACEA

@ OAC

Evie

wo t

FNP ae

~ 77) 8

s ef! f evcroy

& Salsa decreasing

¥ oosD Fs

“on ero

B oe axet

~ oe

& 7 Oe

B&C

Fig. 33. A. Triangular plot of ostracod populations of all samples in the BH9 borehole in

terms of percentage of Cytheracea, Bairdiacea plus Cypridacea, and Cytherellidae. The fields

of ostracod assemblage 1 to 5 are outlined. B. Variations in the ostracod populations (expressed

in terms of Cytheracea, Bairdiacea plus Cypridacea, and Cytherellidae) with changing sedi-

mentary environments in the BH9 borehole. Arrows indicate direction of change up the bore-

hole based on movement within the fields of the various assemblages shown in Figure 33A.

In the elongate fields of assemblages 1 and 5 these trends are actually followed in successive

samples up the borehole. The square dot line separates high and low energy environments,

whilst the open circle line separates shallow and moderate/deep-water environments.

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 67

BIOSTRATIGRAPHY

The main stratigraphic horizons recognized by Klinger & Kennedy (1977,

fig. 12) in BH—-9 are the Santonian/Campanian, and Campanian I/II boundaries:

the former is marked by the appearance of Vendegiesiella cf. V. spinosa and

V. trituberculata, and the latter by the appearance of Baculites vanhoepeni.

These horizons, together with a suggested subdivision of the Santonian based

on a comparison with Madagascar ammonite faunas, are shown in Figure 32.

Both the Santonian/Campanian, and Campanian I/II boundaries fall very close

to the boundaries between ostracod assemblages 3 and 4, and 4 and 5, respec-

tively, which raises the possibility that the appearance of the different ammonite

species might also be facies-controlled and not strictly phylogenetic. Further

work on the Santonian and Campanian ostracods of Zululand will clarify this

issue.

Notwithstanding this possibility, it is desirable to see if any phylogenetic

trends can be recognized in the ostracod faunas against the dominating patterns

of environmental control. Certainly at this stage it is not possible to say, for

instance, that the appearance of Cytherelloidea griesbachi, Haughtonileberis

vanhoepeni, Trachyleberis zululandensis or Amphicytherura zululandensis is

diagnostic of the basal Campanian strata throughout south-east Africa. They

may turn out to be so, but their apparent environmentally controlled ranges in

the borehole leads one to suspect that this is not the case, unless of course they

are endemic to the Zululand area and they evolved solely because of environ-

mental stress—in which case they will be restricted to a particular facies. On the

other hand, some of the obviously less environmentally-bound species may show

phylogenetic relationships that will turn out to be biostratigraphically useful.

Amongst them may be Brachycythere pondolandensis, Gibberleberis africanus

and Cytherelloidea newtoni (mid-upper Santonian), Rayneria nealei (mid-

Santonian to Campanian I), Amphicytherura tumida (upper Santonian to

Campanian II), and Oertliella pennata (upper Santonian to Campanian I).

As might be anticipated, ostracods from the lower part of the borehole show

very close affinities with the Santonian assemblages from the lower part of the

Umzamba cliff section. According to Klinger & Kennedy (1977), the lowest 7 m

at Umzamba are a condensed equivalent of the section 159 m—115 m in the

Richards Bay borehole, but whilst the ostracod assemblages in the borehole

show considerable temporal variations, the four samples studied by Dingle

(1969, fig. 21) show little or no significant variation from the base to the top of

the Umzamba Santonian sequence. At the latter, Brachycythere longicaudata

dominates the fauna throughout (>30°%) with Cytherelloidea umzambaensis,

Haughtonileberis haughtoni and Pondoina sulcata consistently present up to about

10% each. On this evidence, the restriction of Pondoina sulcata and Cythereis

transkeiensis to assemblages 3 and 1, respectively, in the borehole can only be

explained by environmental control, whilst a similar restriction of Oertliella

pennata (originally identified as Acanthocythereis aff. A. horridula by Dingle

(1969)) to the Santonian at Umzamba has no biostratigraphic significance.

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 4

Santonian ostracods at Richards Bay and Umzamba.

Richards Bay Umzamba

Cythereis transkeiensis *

C. klingeri

Cytherelloidea newtoni

C. umzambaensis

C. gardeni

C. griesbachi

Brachycythere longicaudata

B. pondolandensis

B. rotunda

B. sicarius

Paracypris umzambaensis

P. zululandensis

Haughtonileberis haughtoni

A. fissilis

H. vanhoepeni

Cytherella spp.

Rayneria nealei

Gibberleberis africanus

G. elongata

Bairdoppilata andersoni

Bythocypris richardsbayensis

Pondoina sulcata

Amphicytherura tumida

Oertliella pennata

O. sp. A.

Paraphysocythere thompsoni

Veenia obesa

?Cnestocythere

28 types (Cytherella spp counted as one), 17 common. Similarity 61 %.

eee | eee | x | eee | ee | ee ee we | ee | we

Table 4 shows the species identified at both localities and indicates a 61 per cent

similarity at the species level. Significant differences include the restriction of

several species to either area: to Umzamba Paraphysocythere thompsoni,

Brachycythere rotunda and Veenia obesa; to Richards Bay Gibberleberis elongata,

Cytherelloidea newtoni, and the virtual substitution of Cythereis transkeiensis by

C. klingeri in the borehole. In terms of Figure 33B the Umzamba faunas fall

within the shallow water high energy environments, with one sample on the

border of the field adjacent to a somewhat lower energy environment.

Commentary on the similarity of the Richards Bay ostracods with extra-

southern African Santonian/Campanian faunas will be postponed until the

ostracods from other outcrops of a similar age in Zululand have been studied.

ACKNOWLEDGEMENTS

The author thanks Dr H. Klinger of the South African Museum for making

the borehole samples available, and for helpful discussion on the stratigraphy of

Zululand. The borehole material was originally the property of the Geological

Survey of South Africa, but has recently been donated to the South African

MARINE SANTONIAN AND CAMPANIAN OSTRACODS 69

Museum by the Survey Director Dr W. L. van Wyk. The study was made

possible by a University of Cape Town staff research grant for which the author

is grateful. Mr G. Lowcock and Dr D. Crawford of the Electron Microscopy

Unit at the University of Cape Town are thanked for their help in the SEM

photography which was undertaken on the Unit’s instruments.

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ALEXANDER, C. I. 1933. Shell structure of the ostracode genus Cytheropteron and fossil species

from the Cretaceous of Texas. J. Paleont. 7: 181-214.

APOSTOLESCU, V. 1961. Contribution a l’étude paléontologique (Ostracodes) et stratigraphique

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779-867.

BairD, W. 1845. Arrangement of British Entomostraca, with a list of species, particularly

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Bairp, W. 1850. The natural history of the British Entomostraca: 1-364, London, Ray Society.

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BENSON, R. H. 1977. The Cenozoic ostracode faunas of the Sao Paulo Plateau and the Rio

Grande Rise (DSDP Leg 39, Sites 356 and 357). Initial Reports of the Deep Sea Drilling

Project 39: 869-883. Washington: United States Government Printing Office.

BosquET, J. 1854. Monographie des crustacés fossiles du terrain Crétacé du Duché Limbourg.

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2: 53-126.

Brapy, G. S. 1880. Report on ostracoda dredged by H.M.S. Challenger during the years

1873-76. Reports on the scientific results of the voyage of H.M.S. Challenger, Zoology 1(3):

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Brapy, G. S. 1898. On new or imperfectly-known species of ostracoda, chiefly from New

Zealand. Trans. zool. Soc. Lond. 14: 429-452.

BuT er, E. A. & Jones, D. E. 1957. Cretaceous ostracoda of Prothro and Rayburn salt domes

Bienville Parish, Louisiana. Bull. geol. Surv. La. 32: 1-65.

CHAPMAN, F. 1898. On ostracoda from the ‘““Cambridge Greensand”. Ann. Mag. nat. Hist.

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Bull. geol. Sury. West. Aust. 72: 1-87.

CHAPMAN, F. 1923. On some foraminifera and ostracoda from the Cretaceous of the Umzamba

River, Pondoland. Trans. geol. Soc. S. Afr. 26: 107-118.

CorYELL, H. N., SAMPLE, C. H. & JENNINGS, P. H. 1935. Bairdoppilata, a new genus of ostra-

coda, with two new species. Am. Mus. Novit. 777: 1-5.

DAMmoTrTE, R. 1974. Un noveau genre d’ostracode de |’Albien du Bassin de Paris: Matronella n.g.

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DINGLE, R. V. 1969. Upper Senonian ostracods from the coast of Pondoland, South Africa.

Trans. R. Soc. S. Afr. 38: 347-385.

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39: 353.

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African continental margin). Trans. R. Soc. S. Afr. 39: 393-418.

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DINGLE, R. V. 1976. Palaeogene ostracods from the continental shelf off Natal, South Africa.

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geology of the world II, The Mesozoic A: 401-434. Amsterdam: Elsevier.

Forster, H. D. 1975. Die geologische Entwicklung von Stid-Mozambique seit der Unterkreide

und die Ammoniten-Fauna von Unterkreide und Cenoman. Geol. Jb. 12: 3-324.

Howe, H. V. & LaurencicH, L. 1958. Introduction to the study of Cretaceous Ostracoda.

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Jones, T. R. 1849. A monograph of the Entomostraca of the Cretaceous Formations of England.

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Jones, T. R. 1884. Notes on the Foraminifera and Ostracoda from the deep boring at Richmond

Q. Jl geol. Soc. Lond. 40: 766-777.

Jones, T. R. & HINDE, G. J. 1890. A supplementary monograph of the Cretaceous Entomostraca

of England and Ireland. London: Palaeontographical Society.

KeEnneby, W. J. & KLINGER, H. C. 1975. Cretaceous faunas from Zululand and Natal, South

Africa. Introduction, stratigraphy. Bull. Br. Mus. nat. Hist. (Geol.) 25: 263-315.

Kuincer, H. C. & KENNEDY, W. J. 1977. Upper Cretaceous ammonites from a borehole near

Richards Bay, South Africa. Ann. S. Afr. Mus. 72: 69-107.

LATREILLE, P. A. 1806. Genera Crustaceorum et Insectorum 1: 1-303. Paris.

MANDELSTAM, M. I. 1960. In: OrnLov, Y. A. ed. Elements of palaeontology. Vol. VIII.

Arthropods: 1-515, Moscow, Geological and subsurface prospecting: Scientific and

Technical Edition. (In Russian.)

Maup, R. & Orr, W. 1975. Aspects of post-Karroo geology in the Richards Bay area. Trans.

geol. Soc. S. Afr. 78: 101-109.

Moore, R. C. ed. 1961. Treatise on invertebrate paleontology. Part Q, Arthropoda 3. Lawrence:

University of Kansas Press.

MULLER, G. W. 1894. Die Ostracoden des Golfes von Neapel und der angrenzenden Meeresab-

schnitte. Fauna Flora Golf. Neapel 31: 1-404.

NEALE, J. W. 1975. The ostracod fauna from the Santonian Chalk (Upper Cretaceous) of

Gingin, Western Australia. Special Papers in Palaeontology 16: 1-81.

Puri, H. S. 1954. Contribution to the study of the Miocene of the Florida Panhandle. Part III

(Ostracoda). Bull. geol. Surv. Fla. 3: 217-309.

Puri, H. S. 1955. Hermanites, a new name for Hermania Puri 1954. J. Paleont. 29: 558.

Poxorny, V. 1964. Oertliella and Spinicythereis, new ostracod genera from the Upper

Cretaceous. Vest. ustred. Ust. geol. 39: 283-284.

Reuss, A. E. 1846. Die Versteinerungen der bohmischen Kreide Formation: 2: 59-148, Stuttgart;

Schweizerbart.

Sars, G. O. 1866. Oversigt af Norges marine ostracoden. Forh. VidenskSelsk. Krist. 7: 1-130.

Sars, G. O. 1888. Nye Bidrag til Kundskaken om Midlehavets Invertebrat fauna 4. Ostracoda

Mediterranea. Arch. Math. Natury. 12: 173-324.

Sars, G. O. 1922-1928. An account of the Crustacea of Norway 9. Ostracoda, Parts 1-16:

1-277. Bergen: Bergen Museum.

ROSENFELD, A. & BEIN, A. 1978. A preliminary note on recent ostracodes from shelf to rise

sediments off Northwest Africa. Meteor ForschErgebn. 29: 14-20.

SYLVESTER-BRADLEY, P. C. 1948. The ostracode genus Cythereis. J. Paleont. 22: 792-797.

SYLVESTER-BRADLEY, P. C. & BENSON, R. H. 1971. Terminology for surface features in ornate

ostracods. Lethaia 4: 249-286.

6. SYSTEMATIC papers must conform to the International code of zoological nomenclature

(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be

followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.

nov., syn. nov., etc.

‘An author’s name when cited must follow the name of the taxon without intervening

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Synonymy arrangement should be according to chronology of names, i.e. all published

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order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

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Holotype

SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach

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Note standard form of writing South African Museum registration numbers and date.

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Biological Abstracts.

R. V. DINGLE

MARINE SANTONIAN AND CAMPANIAN

OSTRACODS FROM A BOREHOLE AT

RICHARDS BAY, ZULULAND

MAY 1980 ISSN 0303-2515

OF THE SOUTH AFRICAN

MUSEUM

CAPE TOWN

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Examples (note capitalization and punctuation)

BuLLouGu, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

Fiscuer, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

Fiscuer, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs

Zool. exp. gén. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.

Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.

Bull, Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910, Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische

und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.

Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 82 Band

May 1980 Mei

Part 2 Deel

RECORDS OF THE SUBANTARCTIC FUR SEAL

(ARCTOCEPHALUS TROPICALIS) FROM

SOUTH AFRICA WITH NOTES ON ITS BIOLOGY

AND SOME OBSERVATIONS OF

CAPTIVE ANIMALS

PETER D. SHAUGHNESSY

GRAHAM J. B. ROSS

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na gelang van die

beskikbaarheid van stof

Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad

OUT OF PRINT/UIT DRUK

1, 2(1-3, 5-8), 3(1-2, 4-5, 8, t.—p.i.), 5(1-3, 5, 7-9),

6(1, t.—p.i.), 711-4), 8, 9(1—2, 7), 10(1-3),

11(1-2, 5, 7, t.-p.i.), 15(4—5), 24(2), 27, 31(1-3), 32(S), 33

Kopieregnayvrae aan die Suid-Afrikaanse Museum

ISBN 0 908407 92 0

Printed in South Africa by In Suid-Afrika gedruk deur

The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk.,

Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

RECORDS OF THE SUBANTARCTIC FUR SEAL (ARCTOCEPHALUS

TROPICALIS) FROM SOUTH AFRICA WITH NOTES ON ITS

BIOLOGY AND SOME OBSERVATIONS OF CAPTIVE ANIMALS

PETER D. SHAUGHNESSY

Sea Fisheries Branch, Cape Town

GRAHAM J. B. Ross

Port Elizabeth Museum, Port Elizabeth

(With 2 figures and 7 tables)

LMS. accepted 7 February 1980]

ABSTRACT

Information is provided on 22 records of the subantarctic fur seal from South Africa

from June 1966 to June 1979. They occurred between Cape Town and Richards Bay, Natal,

principally between May and September. In the three years 1976 to 1978 an average of about

four seals per year was recorded. Suggested sources of these seals are Gough Island in the

South Atlantic Ocean and the Prince Edward Islands in the Southern Indian Ocean. External

measurements, organ masses, stomach contents and parasites are described for some of the

seals. Observations on the behaviour and feeding of captive animals are included.

CONTENTS

PAGE

Introduction . : : j : : ae ol

Methods ; i : 2 ; ; x2

Records ‘ ; ; ; : : ee A

Stomach contents and feeding . a en Ot

Parasites : : ‘ i ‘ : ess

External body measurements . : A 85:

Discussion . ; : 3 : 5 = 85

Acknowledgements . 3 ; 3 . 88

References . ; , ; é j Erss

INTRODUCTION

The subantarctic or Amsterdam Island fur seal, Arctocephalus tropicalis,

breeds on islands in the subantarctic region: Gough and the Tristan da Cunha

Islands in the South Atlantic Ocean; the Prince Edward Islands, New Amster-

dam and St Paul Islands in the Southern Indian Ocean (Shaughnessy 1980).

One vagrant was recorded at Macquarie Island (54°S 159°E) in 1959 (Csordas

1962), five at South Georgia (54°S 38°W) between 1972 and 1976 (Payne 19795),

and one from Tramandal, southern Brazil in 1976 (Castello & Pinedo 1977);

eight in South Africa have been referred to briefly by Nel (1971) or Shaughnessy

(1980). Details of these last eight records and another fourteen in South African

Ann. S. Afr. Mus. 82 (2), 1980: 71-89, 2 figs, 7 tables.

72 ANNALS OF THE SOUTH AFRICAN MUSEUM

waters up to June 1979 are reported here. Localities where they were observed

or captured are shown in Figure 1. Catalogue numbers of specimens used in this

study that are stored in the South African and Port Elizabeth Museums are

provided in Table 1.

Nomenclature of fur seals in this paper follows that of Rice (1977).

METHODS

Live subantarctic fur seals in South African waters have been distinguished

from resident Cape fur seals, A. pusillus, primarily by the colour of their pelage,

especially by their cream-coloured chest, throat and face. In addition, the

presence of a tuft of black guard hairs on the top of the head (a crest) has been

taken as uniquely indicative of A. tropicalis bulls (Bonner 1968; Repenning et al.

1971; Condy 1978). External features of stranded fur seals have also been com-

pared with the identification guide for A. tropicalis and A. gazella prepared

by Condy (1978). The antarctic or Kerguelen fur seal, A. gazella, breeds on

islands south of the Antarctic Convergence, e.g. South Georgia (Payne 1978),

and also on Marion Island (Condy 1978).

Specimens for which skeletal material was available have been identified

by means of the following characters. A. tropicalis has simple, highly pointed

postcanine teeth which rarely have secondary cusps, whereas A. pusillus has

prominent anterior and posterior cusps (Repenning et al. 1971). The postcanines

of A. gazella also lack cusps, but their postcanines are very much reduced,

particularly the rear two (King 1959; Repenning et al. 1971; Condy 1978).

In addition, skulls of A. tropicalis can be distinguished from those of A. gazella

by their narrow arched palate. The width of the palate at the sixth postcanine

relative to the condylobasal length has been compared with measurements

provided in Table 1 of King (1959), namely 9,4 to 14,8 per cent for A. tropicalis

(the northern form) and 14,9 to 17,9 per cent for A. gazella (the southern form).

Most whole animals which were inspected were measured following the

recommendations of the American Society of Mammalogists (1967). In addition,

straight-line point-to-point measurements were taken from the snout to the

anterior insertion of fore-flipper, centre of ear, gape and eye, while circumference

of head was measured at eyes and at canine teeth. The mass of most of the seals

was measured with a spring balance; that of three vagrants at Durban Aquarium

was estimated by comparison with captive Cape fur seals of known mass. Mass

determinations of testes were made after epididymides were removed. Seals

were judged emaciated if the thickness of the blubber and skin above the

sternum was less than 20 mm or if they appeared thin with prominent scapulae

and lumbar vertebrae.

Males were classified as adult if they had a black crest on the top of their

head, and subadult if a crest could not be seen. Females were classified as

immature if they had narrow uterine cornua (each less than 10 mm) or a small

body (mass less than 15 kg).

PiDMpy 4404

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74 ANNALS OF THE SOUTH AFRICAN MUSEUM

The ages of those specimens for which skulls were available were estimated

by three methods. Ridges were counted on the external surface of the root of an

upper canine tooth after it had been removed by boiling, and ages assigned on

the assumption that each ridge represented one year of life. This technique has

been used to age northern fur seals, Callorhinus ursinus, and antarctic fur seals

(Scheffer 1950, Payne 1978), but is only useful for animals up to about 4 years

of age. In addition, a longitudinal section of one of the upper canines was

prepared by hand-grinding on wet carborundum paper and the number of

growth layers counted in the dentine or (when the pulp cavity was closed) in

the cement. It has been assumed that each layer corresponds to one year of life

in this species, as shown for A. gazella by Payne (1978). Of the three methods,

the ridge count is considered to be most accurate for young animals and the

dentine count for older animals. As pups are born from mid-November to mid-

December at New Amsterdam (Tollu 1974) and mainly during mid-December

at Gough (Bester 1977), the age of vagrants has been calculated on the

assumption that they were born at the beginning of December.

RECORDS

Information on each of the twenty-two A. tropicalis vagrants in South Africa

is provided in chronological order. The location and date of stranding of each

animal are provided in Table 1, together with the date on which it died or

returned to the sea, the methods used to identify it as A. tropicalis and the

material and other data collected from it. Skulls were collected from twelve of

the vagrants. Data on their condylobasal length and the width of their palate

at the sixth postcanine are provided in Table 2, together with information on

the determination of their age from canine teeth. External measurements and

mass determinations of organs are provided for some of the seals in Tables 3

and 4. A typical A. tropicalis bull (seal number 19) is shown in Figure 2.

1. The first subantarctic fur seal ashore in South Africa of which we are

aware was an extremely emaciated, immature female near Port Elizabeth in

June 1966. It was taken to the Port Elizabeth Oceanarium, and was referred to

by Nel (1971). Initially it had a mass of 2,3 kg and a standard length of 57 cm.

In September 1967 it had attained a mass of 15 kg and a length of 68 cm.

It died in April 1969 after 2 years 10 months in captivity. As the number of

growth layers on a canine tooth was estimated at 44, the seal must have been

18 months old when caught.

This seal had rudimentary anterior cusps on postcanines 2 to 5; such cusps

are unusual in A. tropicalis, but have been noted by King (1959). They were also

apparent on the postcanines of seal number 21. Its fur was noticeably denser

and its eyes larger than those of Cape fur seals in the Oceanarium. A strong,

musk-like odour associated with this seal was readily distinguishable from the

odour of Cape fur seals.

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76 ANNALS OF THE SOUTH AFRICAN MUSEUM

Unlike the goat-like bleat of Cape fur seal pups in the Oceanarium, this seal

uttered a high-pitched wail or cry when requesting food or attention. Similarly,

its typical bark and shorter staccato sounds were also higher pitched than calls

of Cape fur seal pups. When approached closely it sometimes gave a deep

growl before lunging.

2. An adult male which came ashore near Port Elizabeth in September 1970

was also taken to the Oceanarium. It was treated for internal haemorrhaging

but died a few days later. This seal appeared to sip sea-water from its pool on

several occasions.

3. An emaciated, immature female which came ashore near Port Elizabeth in

July 1971 was kept at the Oceanarium for 2 years 5 months. Its age at death is

calculated to have been 3 or 4 years, from which it is deduced that it was

7 or 19 months old when it was captured.

4. An adult male came ashore on the beach near the Durban Aquarium in

May 1973 and remained there for two days before swimming away. Colour

transparencies of the seal were provided by J. Bass (Oceanographic Research

Institute, Durban).

5. An emaciated, adult male which came ashore at Sunset Beach; Cape Town

in December 1974 was put to death 2 days later.

TABLE 2

Skull measurements and estimated age of subantarctic fur seals that

stranded on the coast of South Africa.

Width of palate

at pe 6° Canine teeth

Record CBL*

no. mm No. No.

mm % of CBL | external dentinal

ridges layers

1 Seal no. 1 was in captivity for 2 years 10 months, no. 3 for 2 years

5 months and no. 8 for 2 years 10 months.

2 Pulp closed.

3 Seventeen cement layers counted.

4 Condylobasal length.

5 Sixth postcanine tooth.

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6. An emaciated, subadult male which came ashore at Strandfontein in False

Bay in August 1975 was put to death 3 days later.

7. An adult male captured at Durban in May 1976 was taken to the Durban

Aquarium. As it had a deep laceration on the flank, it was put to death soon

after capture. No specimens were retained from this animal but it was seen by

one of the authors (GJBR).

8. Anemaciated, adult male was caught in Port Elizabeth harbour in June 1976

and taken to the Oceanarium where its mass was determined as 59 kg. After

recovering in isolation for 13 days, it was placed with six Cape fur seals in a

pool (30 m x 10 m), where it immediately fought with the largest male (mass

250 kg) and gained control of the only island. Over the following two months,

however, the Cape fur seal bull regained control of the pool and the island.

In July 1978 this seal was transferred to another aquarium where it died on

2 April 1979, 34 months after capture. Its age at death is calculated as 18 years

and, therefore, its age at capture as 15 years.

9. A subadult male that came ashore at Richards Bay in July 1977 was taken

to the Durban Aquarium. Its mass was estimated at 20 kg. It was released

41 months later on a secluded beach at Port Edward (31°03’S 30°14’E). Colour

80 ANNALS OF THE SOUTH AFRICAN MUSEUM

transparencies of the seal provided by M. Smale (Oceanographic Research

Institute, Durban) indicated that the seal was not A. pusillus and was most

likely A. tropicalis.

10. In July 1977 an emaciated, adult male came ashore at Durban. It was

taken to the Aquarium where it was put to death the next day. Its mass was

estimated at 80 kg and its standard length measured 1,37 m.

11. In August 1977 an immature female came ashore at Port Elizabeth and was

taken to the Oceanarium. Like previous young subantarctic fur seals kept there

its high-pitched vocalizations, large prominent eyes and dense fur were notice-

ably different from those of resident Cape fur seals. Five and a half months after

capture it had a mass of 15,5 kg. It was still alive at the time of writing.

This seal and number 8 were in the Oceanarium together for 11 months.

However, they had no opportunities of breeding as they were kept in different

pools, number 11 with immature Cape fur seals, and number 8 with larger ones.

12. In September 1977 an emaciated, adult male came ashore at Sheffield

Beach. It was taken to the Durban Aquarium where its mass was estimated at

60 kg. It died 4 days later. One of the authors (GJBR) inspected the skin of this

seal.

13. In September 1977 an emaciated, adult male came ashore 2 km north-west

of the Cape of Good Hope where it died on the same day.

An unusual feature of the skull of this seal is the presence of seven post-

canines on each side of the upper dental array.

14. In October 1977 an adult male came ashore at Summerstrand, Port

Elizabeth and returned to the sea on the same day. It was identified from a news-

paper photograph (Eastern Province Herald, 27 October 1977).

15. In July 1978 an adult maie came ashore at Reunion Rocks near Durban.

It was taken to the Aquarium and put to death. It was recognized by the

Aquarium staff as being similar to previous subantarctic fur seals.

16. In August 1978 an adult male came ashore in the harbour at Port Elizabeth.

Monel metal tags (Hasco size 49L) were applied to its fore-flippers (numbers

B 4006 and 4007) and it was released near by at Cape Recife on the same day.

Its mass was 73 kg.

17. In August 1978 an emaciated, immature female came ashore at Durban.

It was taken to the Aquarium where it died 4 days later. Its flippers were injured

and its left eye was damaged. The width of each uterine cornu was 7 mm.

18. In August 1978 an emaciated, immature female came ashore at Three

Anchor Bay, Cape Town. It was blind in the left eye. Four days later it died in

captivity. Inspection of its skull revealed that the left supraorbital process had

been broken. Its uterine cornua were 4 mm (left) and 3 mm wide.

19. In September 1978 an emaciated, adult male came ashore at Summerstrand,

Port Elizabeth (Fig. 2). Tags were applied to its fore-flippers (numbers B 4008

RECORDS OF THE SUBANTARCTIC FUR SEAL 81

and 4009) and it was released the same day at Cape Recife. It was still there on

the next day, but had left 2 days later.

20. In March 1979 an emaciated, immature female came ashore at Strand-

fontein, False Bay, and was put to death 3 days later. It was lacerated on the

ventral surfaces of the left fore-flipper and on the left shoulder, and had three

bite marks in the centre of the dorsum. Its uterine cornua were both 4 mm wide.

21. In May 1979 a subadult male came ashore at Strandfontein, False Bay and

was put to death the same day. Rudimentary cusps on the anterior edge of its

postcanines were better developed than those of seal number 1.

22. In June 1979 an adult male in good condition was seen on a bank of the

Swartkops River 6 km from the sea. It was tagged on the fore-flippers (num-

bers B 4012 and 4013) before being released at Cape Recife.

STOMACH CONTENTS AND FEEDING

Stomachs of eight of the seals were examined. All died within 5 days of

stranding. One was fed finely ground fish, but no fish remains were found in its

stomach. Only one of them (from number 20) contained no food items. Squid

beaks were found in the remaining seven stomachs (Table 5). They have been

identified to family level by means of the features outlined by Clarke (1962).

Their designations below family level follow the names used by Ross (1979).

Other food items were also found in two of the stomachs: seal number 6

contained two unidentified crustacean fragments, and seal number 13 contained

1,8 g of sand and 6,8 g of partly digested gladii from squid.

The squid most commonly taken were members of the family Histio-

teuthidae and particularly the neritic genus Loligo. At New Amsterdam Island,

Paulian (1964) and Tollu (1974) noted that squid were very important food of

TABLE 5

Numbers of squid identified from beaks in stomachs of seven sub-

antarctic fur seals stranded on the coast of South Africa.

Seal record no.

Squid species

Loligo reynaudi 1 1 1 8 34

Sepia sp. (large) 1

Sepia sp. (small) 4

Ommastrephes caroli 1

Onychoteuthidae:

type 2? 1 1

Histioteuthidae:

type 22 2

type 8* 1 1 3 1

1 Following the designations used by Ross (1979).

82 ANNALS OF THE SOUTH AFRICAN MUSEUM

A. tropicalis, whereas at Marion Island, Rand (1956) noted that fish were more

important than squid. No fish remains were found in the stomachs of

A. tropicalis on the South African coast.

Information is available on the food fed to five of the vagrants that were

kept in captivity.

Seal number 1, an immature animal, was fed the freshwater fish Sarotherodon

mossambicus. Seal number 2, an adult, was fed fish, mainly pilchard, Sardinops

ocellata.

Most information is available for seal number 8, an adult male. It was fed

fish, mainly Pagellus natalensis, maasbanker, Trachurus trachurus, and pilchard,

as well as squid, Loligo reynaudi. For the first 6 months it consumed an average

of 5,7 kg per day, and its body mass increased from 59 kg to over 90 kg. Thus

the feeding rate was between six and ten per cent of the body mass per day. Its

food intake during summer (from November to January) averaged 2,2 and

0,7 kg per day for 1976-7 and 1977-8, respectively. This was considerably lower

than for the remainder of the year at 3,6 kg per day (Table 6) and is possibly

associated with the breeding season when bulls fast while ashore holding

territories for an average of 45 days (Bester 1977). A similar decrease in the

food intake of captive northern elephant seals Mirounga angustirostris during

breeding and moulting seasons has been reported by Bryden (1969).

Seal number 9, a subadult, was fed hake, Merluccius spp., herring, Clupea

harengus, pilchard, and Pomadasys olivaceum. When captured its mass was

estimated at 20 kg; this had decreased to 15 kg when it was released 135 days

later. It ate on 80 days and consumed fish at the rate of 0,67 kg per day over its

total time in captivity. This is equivalent to between three and four per cent of

its body mass daily. Captive seals require six to ten per cent of their body mass

TABLE 6

Average daily mass (kg) of food consumed by

seal no. 8 (PEM 1521/67) during captivity at

Port Elizabeth Oceanarium from 16 June 1976

to 26 July 1978.

1976 1977 1978

January 2,6 0,8

February 2,6 1,6

March Dei 2,0

April 2,9 4,1

May 3,6 6,0

June 5,8 Sif 5,9

July 5,4 3,6 6,0

August 5,9 3,6

September 5,6 3,6

October 5,9 2,4

November 5,7 0,8

December 2 0,4

{

(

RECORDS OF THE SUBANTARCTIC FUR SEAL 83

per day for maintenance purposes according to Keyes (1968), although adult

harp seals, Pagophilus groenlandicus, have been maintained on a daily intake of

three to four per cent of their body mass (Ronald et al. 1975). In addition to

fish, it was provided with liquid paraffin (to prevent constipation), vitamin

supplements and antibiotics.

Seal number 11, an immature animal, was fed for 2. months on an artificial

milk formula based on that recommended by Marine Mammal Biological

Laboratory (1969). Later it took solid fish (pilchards and maasbankers). For the

7 months December 1977 to June 1978 its average daily food intake was 0,72 kg

per day (range 0 to 1,0). In mid-February 1978 its mass was 15,5 kg and it was

much larger than when captured. Thus its daily feeding rate of about five per

cent of its body mass was sufficient for its mass to increase.

PARASITES

External parasites were found only on seal number 13. In its pelage were

several hundred lice identified as Proechinophthirus sp. by J. A. Ledger (South

African Institute for Medical Research). The density of these parasites was

greatest on the seal’s head and neck, and decreased posteriorly.

Nasal passages and sinuses of two seals were examined for parasites. Two

mites were found in seal number 21 and twenty-seven mites in seal number 8.

Representatives were identified as Orthohalarachne diminuata by R. Domrow

of the Queensland Institute of Medical Research. Paulian (1964) found nasal

mites, O. chabaudi (= O. diminuata) in Arctocephalus tropicalis from New

Amsterdam Island.

The blubber of nine seals was inspected for cestode cysts, Phyllobothrium sp.,

and their number counted on one side of a mid-ventral incision from chin to

anus of seals number 6 (1 cyst), 8 (2 cysts), 13 (12 cysts), 17 (none), 18 (6 cysts),

20 (4 cysts) and 21 (1 cyst). Small numbers of cestode cysts were also seen in the

blubber of numbers 3 and 5. The cyst infestation of adult males (seals number 5,

8 and 13) was less than for A. pusillus bulls (PDS, pers. obs.).

The viscera of nine of the seals were examined for parasites. None was seen

in seals number 3, 6 and 21, nor in the internal organs examined in seal number 8

(heart, lungs, trachea and oesophagus). Seal number 5 contained nematodes,

Contracaecum osculatum, in the oesophagus (sixteen individuals) and in the

stomach (75 g), and a few acanthocephalans, Corynosoma australe, in the small

intestine. Seal number 13 contained a few unidentified cestodes in its small

intestine. Its stomach contained 7 nematodes: 6 Anisakis simplex and | tenta-

tively identified as C. osculatum. These parasites were identified by D. I. Gibson

of the British Museum (Natural History). One acanthocephalan was found in the

small intestine of the seal number 17. 3 nematodes were found in the oesophagus

and 4 in the stomach of seal number 18; its small intestine contained a small

number of unidentified cestodes. The stomach of seal number 20 contained

109 nematodes.

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RECORDS OF THE SUBANTARCTIC FUR SEAL 85

Paulian (1964) found cestode cysts in the blubber of 11 of the 13 Arcto-

cephalus tropicalis he examined at New Amsterdam. Of the above-mentioned

parasites, he found only Contracaecum ogmorhini (= C. osculatum) in the

stomach. In addition, he observed Terranova (= Porrocaecum) decipiens in

stomachs.

The nematode Anisakis simplex and the acanthocephalan Corynosoma

australe are not listed as parasites of A. tropicalis in the comprehensive checklist

of Dailey & Brownell (1972).

EXTERNAL BODY MEASUREMENTS

An attempt has been made to distinguish between Arctocephalus tropicalis

and A. pusillus by external body measurements, so that less reliance is placed on

body colour for identifying vagrants of the former in South Africa. Five

measurements concerning the flippers and the head, expressed as proportions of

the standard body length, were compared (Table 7). Two size groups were

considered, following the availability of A. tropicalis vagrants, namely: large

males of standard length 129-165 cm, and small animals of both sexes of standard

length 85-100 cm. The A. pusillus used for the comparisons were collected at sea

between March 1977 and March 1979.

For both the large males and the small seals fore- and hind flippers are

proportionally longer in A. pusillus. Furthermore, among the small seals, the

fore-flippers are attached to the body at a more posterior position in A. pusillus

than in A. tropicalis and the head is longer. However, these measurements are of

limited value in distinguishing vagrant A. tropicalis from A. pusillus because of

overlap in their ranges.

Condy (1978) pointed out that at Marion Island flippers of A. tropicalis are

short and broad while those of A. gazella are distinctly longer. Payne (1979a)

graphed anterior lengths of flippers and tabulated mean standard lengths of

A. gazella at South Georgia from birth to 11 years of age. The fore-flipper length

(expressed as a proportion of standard length) ranged from 0,28 to 0,33. This is

considerably larger than (and does not overlap with) that of the small sample of

eleven vagrant A. tropicalis with range 0,19 to 0,26 (see Table 7). The hind

flipper length of A. gazella at South Georgia ranged from 0,22 to 0,28, which is

also larger than that of the vagrant A. tropicalis (with range 0,12 to 0,16). These

comparisons, therefore, support Condy’s statement that flippers of A. tropicalis

are shorter than those of A. gazella.

DISCUSSION

The occurrence of A. tropicalis in South Africa is decidedly seasonal, with

19 of the 22 records occurring from May through September, i.e. predominantly

in the winter and early spring. At Gough Island (Bester 1977) and Marion Island

(Condy 1978) the number of seals ashore increased during September and

86 ANNALS OF THE SOUTH AFRICAN MUSEUM

October prior to the breeding season and then declined during April and May.

Thus the months when A. tropicalis most commonly occurs in South Africa

coincide with the period when they are scarce on Gough and Marion Islands.

Records of crabeater seals, Lobodon carcinophagus, and southern elephant seals,

Mirounga leonina, in South Africa are also seasonal, but occur mostly in the

summer months (Ross et al. 1978a; Ross 1969; Best 1971).

Records of A. tropicalis ashore have increased from 6 in the 10 years

between 1966 and 1975 (with no more than one per year) to 2 in 1976, 6 in 1977

and 5 in 1978. This increase most likely reflects the increased awareness of

marine mammals by the public in recent years and follows the establishment of

facilities for displaying them in Port Elizabeth and Durban. It may also be

related to population increases at breeding islands (Condy 1978).

7 of the 22 A. tropicalis hauled out on the coast of Natal, 9 in the vicinity of

Port Elizabeth and 6 near Cape Town. The areas in which these records are

grouped occur near interested biologists, which suggests that animals elsewhere

go unrecognized or unrecorded. A similar phenomenon has been noted for

crabeater seals on the South African coast (Ross et al. 1978a) and is also

apparent for records of elephant seals (Ross 1969; Best 1971) and rockhopper

penguins, Eudyptes chrysocome (Cooper et al. 1978). The distribution of

A. tropicalis records shows a greater concentration on the eastern coast than for

the other vagrant seal species. Since fur seals in general are rare east of Port

Elizabeth, the presence of a seal there would attract more attention than one on

the west coast where A. pusillus is common.

The ages of twelve of the vagrant seals can be deduced from information in

Table 2. The number of dentinal layers in the canine teeth indicates that 5 of

them (all males) were older than 10 years and one was 6 years of age. The

number of external ridges indicates that 6 of them (5 females and 1 male) were

younger than 5 years.

The last group seem small for their estimated age of 2 to 44 years. The

average mass and standard length of four of the females (omitting number |

which was extraordinarily small) was 8,5 kg and 86,8 cm, and that of the single

male was 8,0 kg and 94 cm. According to Tollu (1974), at age 10 months females

at New Amsterdam Island have an average mass of 14,4 kg and length of

85,8 cm, and males have averages of 18,0 kg and 92,7 cm, respectively. The mass

of these vagrants cannot be compared with that of seals from a colony because

they were emaciated. However, as they appear to be only slightly longer on

average than 10-month-old seals, they were either well below average size for

their age or their age estimates were inflated.

While handling the canine teeth of the six young seals it was noticed that

that of the male (number 6) was considerably larger in cross-section than those

of the five females. This suggests that there is a sexual dimorphism for size of

canine teeth in A. tropicalis (at least in juveniles) similar to that described for

northern elephant seals by Briggs & Morejohn (1975).

Palate width of the 12 vagrants measured ranged from 10,2 to 13,6 per cent

RECORDS OF THE SUBANTARCTIC FUR SEAL 87

of the skull condylobasal length. This is well within the range proposed by King

(1959) for A. tropicalis. Furthermore, none of the skulls had particularly

reduced postcanine teeth, supporting the conclusion that they were A. tropicalis.

Black crests were visible on the heads of 5 males aged 10 years or more

(numbers 2, 5, 8, 10 and 13), and on another 4 males judged to be adult on

account of their length (numbers 15 and 19) or mass (numbers 12 and 16). On

the other hand, crests could not be seen on 3 of the males. 2 of these (numbers 6

and 21) were judged to be 2 and 6 years old, respectively, and the third

(number 9), with a mass of 20 kg, was judged to be a subadult. However, small

crests were visible on the tanned skins of seals numbers 6 and 21. Thus it seems

that crest development in the male occurs between 6 and 10 years of age, but that

rudimentary crests (not usually visible in the dead, intact animal) may be present

in younger males.

The uterine cornua of three seals were measured (numbers 17, 18 and 20).

The maximum width was 7 mm, indicating that they were not pregnant. The age

of these seals was estimated at less than 4 years.

Of the 22 vagrants, 13 (59 per cent) were adult males and 3 (14 per cent)

were subadult males. Because adult females nurse their pups for 10 to 11 months

(Tollu 1974; Bester 1977) when they visit the colonies regularly, it is not

surprising that most of the vagrants were males.

The strong, musk-like odour noted for seal number 1 at the Port Elizabeth

Oceanarium was also noted by GJBR when handling other subantarctic fur seals

at Port Elizabeth (numbers 2, 3, 8, 11, 14, 16, 19 and 22), and was noticeably

different from the odour of Cape fur seals.

The source of these seals is unknown. The largest population of this species

occurs at Gough Island (Bester 1977), 3 000 km west-south-west of Cape Town.

This is a likely source of emigrants as its population is expanding rapidly.

Vagrants from there would be assisted by the West Wind Drift until they

reached the complex system of currents in the vicinity of the Cape Peninsula.

A closer breeding area is the Prince Edward Islands, 1 800 km south-east of

Port Elizabeth, where the population size is much smaller (Condy 1978).

To move north from there seals must first cross the West Wind Drift, which

flows between north-east and north in that vicinity (Hydrographic Department

1961). They could then enter the northerly flowing Agulhas Return Current and

reach the Natal coast in the Agulhas Current (Harris 1970). Such current-

assisted access between the Prince Edward Islands and the east coast of South

Africa has been suggested by Ross et al. (19786) for a macaroni penguin,

E. chrysolophus.

A local origin for the seals (from an undetected colony) is discounted

because of the marked seasonality of the records, which coincides with an

off-shore movement at their breeding islands. This conclusion is reinforced by

the fact that about half of the vagrants were in an emaciated condition when

they were reported. 5 of them died and 7 had to be killed within 4 days. 4 of these

12 were injured or damaged when caught. 5 of the vagrants were kept in captivity

88 ANNALS OF THE SOUTH AFRICAN MUSEUM

for periods of between 44} and 34 months, and 1 of these is still alive. 6 of the

seals returned to the sea (one of them after 4$ months in captivity).

ACKNOWLEDGEMENTS

We are grateful to the following for assistance with the collection of

specimens: Drs E. A. Fernhead, M. Smale and J. Bass of the Oceanographic

Research Institute, Durban; Mr H. C. V. Bell of the Cape of Good Hope

Society for the Prevention of Cruelty to Animals; Mr J. Seal of the Snake and

Animal Park, Hartebeespoort; the South African National Foundation for the

Conservation of Coastal Birds; Mr G. Wright and Mr N. Clark of the Cape of

Good Hope Nature Reserve. We thank the Directors of the Port Elizabeth and

South African Museums and of the Oceanographic Research Institute for per-

mission to examine and publish data on specimens in their collections. Thanks

are also extended to Dr D. I. Gibson, Dr R. Domrow and Dr J. A. Ledger for

identifying parasites, and to Dr P. B. Best of the Sea Fisheries Branch and Dr

P. R. Condy of the Mammal Research Institute, University of Pretoria, for

comments on the manuscript.

REFERENCES

AMERICAN SOCIETY OF MAMMALOGISTS 1967. Standard measurements of seals. J. Mammal. 48:

459-462.

Best, P. B. 1971. Stalked barnacles Conchoderma auritum on an elephant seal: occurrence of

elephant seals on South African coast. Zool. afr. 6: 181-185.

Bester, M. N. 1977. Habitat selection, seasonal population changes, and behaviour of the

Amsterdam Island fur seal Arctocephalus tropicalis on Gough Island. Unpublished

D.Sc. thesis, University of Pretoria, Pretoria.

Bonner, W. N. 1968. The fur seal of South Georgia. Br. Antarct. Surv. Sci. Repts 56: 1-81.

Briccs, K. & MOREJOHN, G. V. 1975. Sexual dimorphism in the mandibles and canine teeth

of the northern elephant seal. J. Mammal. 56: 224-231.

BrRYDEN, M. M. 1969. Growth of the southern elephant seal, Mirounga leonina (Linn.).

Growth 33: 69-82.

CASTELLO, H. P. & Pinepo, M. C. 1977. Arctocephalus tropicalis, first record for Rio Grande

do Sul coast (Pinnipedia, Otariidae). Atlantica, Rio Grande 2: 111-119.

CLARKE, M. R. 1962. The identification of cephalopod ‘beaks’ and the relationship between

beak size and total body weight. Bull. Brit. Mus. (nat. Hist.), Zool. 8: 419-480.

Conpy, P. R. 1978. Distribution, abundance, and annual cycle of fur seals (Arctocephalus spp.)

on the Prince Edward Islands. Sth. Afr. J. Wildl. Res. 8: 159-168.

Cooper, J., Ross, G. J. B. & SHAUGHNESSY, P. D. 1978. Seasonal and spatial distribution of

rockhopper penguins ashore in South Africa. Ostrich 49: 40-44.

CsorDAS, S. E. 1962. The Kerguelen fur seal on Macquarie Island. Vict. Nat. 79: 226-229.

DaiLey, M. D. & BROWNELL, R. L. Jr. 1972. A checklist of marine mammal parasites. In:

RipGway, S. H. ed. Mammals of the sea: biology and medicine: 528-589. Springfield,

Illinois: Thomas.

Harris, T. F. W. 1970. Features of the surface currents in the south west Indian Ocean.

SANCOR symposium: Oceanography in South Africa, Durban, August 1970. Paper C3.

HYDROGRAPHIC DEPARTMENT 1961. The Antarctic pilot. 3rd ed. London: Her Majesty’s

Stationery Office.

Keyes, M. C. 1968. The nutrition of pinnipeds. Jn; HARRISON, R. J., HUBBARD, R. C.,

PETERSON, R. S., Rice, C. E. & SCHUSTERMAN, R. J. eds. The behavior and physiology of

pinnipeds: 359-395. New York: Appleton-Century-Crofts.

—————

RECORDS OF THE SUBANTARCTIC FUR SEAL 89

Kina, J. E. 1959. The northern and southern populations of Arctocephalus gazella. Mammalia

23: 19-40.

MARINE MAMMAL BIOLOGICAL LABORATORY 1969. Fur seal investigations, 1966. U.S. Fish

Wildl. Serv. Spec. Sci. Rep. Fish. 584: 1-123.

NEL, J. A. J. 1971. Order Pinnipedia. Jn: MEESTER, J. & SETZER, H. W. eds. The mammals of

Africa: an identification manual. Washington, D.C.: Smithsonian Institution Press.

PAULIAN, P. 1964. Contribution a l’étude de l’otarie de Vile Amsterdam. Mammalia 28;

Supp! 1: 3-146.

PAYNE, M. R. 1978. Population size and age determination in the Antarctic fur seal Arcto-

cephalus gazella. Mammal. Rey. 8: 67-73.

Payne, M. R. 1979a. Growth in the Antarctic fur seal Arctocephalus gazella. J. Zool., (Lond.)

187: 1-20.

Payne, M. R. 19795. Fur seals Arctocephalus tropicalis and A. gazella crossing the Antarctic

Convergence at South Georgia. Mammalia 43: 93-98.

RAnpb, R. W. 1956. Notes on the Marion Island fur seal. Proc. zool. Soc., Lond. 126: 65-82.

REPENNING, C. A., PETERSON, R. S. & Huss, C. L. 1971. Contributions to the systematics of

the southern fur seals, with particular reference to the Juan Fernandez and Guadalupe

species. In: BuRT, W. H. ed. Antarctic Pinnipedia. Antarctic Research Series 18: 1-34.

Washington, D.C.: American Geophysical Union.

Rice, D. W. 1977. A list of the marine mammals of the world. NOAA Technical Report

NMEFS SSRF 711: 1-15.

RONALD, K., UTHE, J. F. & FREEMAN, H. 1975. Effects of methyl mercury on the harp seal.

ICES CM 1975/N:9 15 pp. (mimeo).

Ross, G. J. B. 1969. The southern elephant seal, Mirounga leonina, on South African coasts.

Ann. Cape Prov. Mus. (nat. Hist.) 6: 137-139.

Ross, G. J. B. 1979. The smaller cetaceans of the south east coast of southern Africa.

Unpublished Ph.D. thesis, University of Port Elizabeth, Port Elizabeth.

Ross, G. J. B., SHAUGHNESSY, P. D. & Best, P. B. 1978a. New records of crabeater seals

(Lobodon carcinophagus) from South Africa. Ann. S. Afr. Mus. 75: 153-158.

Ross, G. J. B., VAN DER Est, R. & De Vituiers, A. F. 1978b. First record of the macaroni

penguin in South Africa. Ostrich 49: 47.

SCHEFFER, V. B. 1950. Growth layers on the teeth of Pinnipedia as an indication of age.

Science 112: 309-311.

SHAUGHNESSY, P. D. 1980. The status of the Amsterdam Island fur seal. In: Mammals in the seas.

FAO Fisheries Series 5(3).

ToLLu, B. 1974. L’otarie de l’ile Amsterdam Arctocephalus tropicalis tropicalis (Gray 1872).

Thése de doctorat de troisiéme cycle, |’Université Paris— VII, Paris.

6. SYSTEMATIC papers must conform to the International code of zoological nomenclature

(particularly Articles 22 and 51). ‘

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be

followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.

nov., syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening

punctuation and not be abbreviated; if the year is added, a comma must separate author’s

name and year. The author’s name (and date, if cited) must be placed in parentheses if a

species or subspecies is transferred from its original genus. The name of a subsequent user of

a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published

scientific names by which the species previously has been designated are listed in chronological

order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

comma separates author’s name and year

“semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby

the year is placed in front of each entry, and the synonym repeated in full for each entry, is

not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-

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PETER D. SHAUGHNESSY

GRAHAM J. B. ROSS

RECORDS OF THE SUBANTARCTIC FUR SEAL

(ARCTOCEPHALUS TROPICALIS) FROM

SOUTH AFRICA WITH NOTES ON ITS BIOLOGY

AND SOME OBSERVATIONS OF

CAPTIVE ANIMALS

_ VOLUME 82 PART 3 JUNE 1980 ISSN 0303-2515

| CAPE TOWN

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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FiscHer, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

FiscHer, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs

Zool. exp. gén. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.

Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.

Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische

und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.

Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 82 Band

June 1980 Junie

Part 3 Deel

VATE: TERTIARY SEALS

OF THE SOUTH ATLANTIC OCEAN

C. DE MUIZON

Q. B. HENDEY

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

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Court Road, Wynberg, Cape Courtweg, Wynberg,

LATE TERTIARY SEALS OF THE SOUTH ATLANTIC OCEAN

C. DE MUIZON

Institut de Paléontologie, Paris

Q. B. HENDEY

South African Museum, Cape Town

(With 15 figures and 2 tables)

LMS. accepted 7 February 1980]

ABSTRACT

A lectotype is designated for Prionodelphis rovereti Frenguelli, 1922 from Argentina and

it is assigned to the Cetacea, while the monachine seal ‘Prionodelphis’ capensis Hendey &

Repenning, 1972, from South Africa is assigned to the new genus, Homiphoca. A nearly

complete skull and mandible is designated as a paratype of H. capensis and this, and other,

additional material is described. H. capensis is morphologically intermediate between living

monk seals of the genus Monachus (Phocidae, Monachinae, Monachini) and the seals of

Antarctica (Phocidae, Monachinae, Lobodontini). It is here assigned to the latter group and,

contrary to earlier opinion, is suggested to be more closely related to the crabeater, Lobodon

carcinophagus, than any other living seal.

CONTENTS

PAGE

Introduction : ; P z : : : 91

South Atlantic late Tertiary Phocidae . é 92

The Argentinian material , : : 92

The South African species : : : 94

Discussion . : ; ; ; : ‘ : 120

Summary/Résumé . : : : : DS:

Acknowledgements . : : ; : 2 126

References . ‘ E ‘ ¢ ‘ , ~. 126

INTRODUCTION

The true seals (family Phocidae), which are today widely distributed in the

oceans of the world, have a very poor fossil record, and there are many

uncertainties concerning their origin, evolution and dispersal (Ray 1976a).

Living Phocidae are generally divided into two subfamilies, the Phocinae

and the Monachinae. The Phocinae are a diverse group comprised of the seals

of northern middle and high latitudes. They include such species as the common

or harbour seal (Phoca vitulina), the ringed seal (Pusa hispida) and the grey

seal (Halichoerus grypus). The Monachinae are the subfamily to which the

fossil seals discussed in this paper belong, and they are here divided into

three groups. The first is the Monachini, which includes the monk seals of the

Mediterranean (Monachus monachus), the Caribbean (M. tropicalis), and

Ann. S. Afr. Mus. 82 (3), 1980: 91-128, 15 figs, 2 tables.

92 ANNALS OF THE SOUTH AFRICAN MUSEUM

Hawaii (M. schauinslandi) (King 1956). The second group is the Lobodontini,

which comprises the seals of Antarctica, namely, the crabeater (Lobodon

carcinophagus), leopard (Hydrurga leptonyx), Weddell (Leptonychotes weddelli),

and Ross seals (Ommatophoca rossi) (Scheffer 1958). Thirdly, there are the

elephant seals (Mirounga leonina, M. angustirostris), whose relationships to other

monachines are discussed elsewhere (De Muizon 1979).

The most significant fossil Phocidae are from Miocene/Pliocene deposits in

five widely separated regions. Important material has been found in the

Paratethyan region, which extends from Austria to the southern Soviet Union.

It has been discussed by several authors since first described by Eichwald (1853),

and Grigorescu (1976) has commenced a much-needed revision. Secondly, the

Antwerp Basin in Belgium has yielded a diverse assemblage of skeletal elements

described by Van Beneden (1877). This material was recently revised by Ray

(1976a, in press), and has also been studied by the senior author. The Calvert

and Yorktown Formations in eastern North America have been the source of a

wealth of material studied by Ray (1976a, in press).

The remaining two regions are in the Southern hemisphere. The Pisco

Formation on the southern coast of Peru is the source of the most complete late

Tertiary phocids ever discovered. This material represents five new species

belonging to four new genera, and has been studied by the senior author

(De Muizon 1979). The last region is the south-western coast of South Africa,

with almost all the known material having come from a single locality, namely,

‘E’ Quarry at Langebaanweg near Cape Town. The ‘E’ Quarry material

represents a single species, which is now the best represented fossil phocid in the

world (Hendey & Repenning 1972; Hendey 1976).

Less significant material has been discovered elsewhere, including

Argentina. The Argentinian material, which consists of a few isolated teeth and

a mandible fragment, is identified as Prionodelphis rovereti (Frenguelli 1922;

Cabrera 1926). It was to the genus Prionodelphis that the South African species

(P. capensis) was referred.

It is the purpose of this paper to re-examine the status of the South African

and Argentinian material.

SOUTH ATLANTIC LATE TERTIARY PHOCIDAE

THE ARGENTINIAN MATERIAL

Frenguelli (1922) based the identification of the new genus and species,

Prionodelphis rovereti, from the late Miocene/early Pliocene of Argentina on two

isolated teeth, one of which is obviously that of a delphinoid cetacean, while the

other is a cheek tooth of a monachine seal. These two teeth are syntypes and the

first illustrated specimen, the cetacean tooth (Frenguelli 1922: 492, fig. 1a), is

here designated the lectotype of the species. P. rovereti is thus a cetacean, which

is the group to which Frenguelli believed both teeth belonged, although it should

LATE TERTIARY SEALS OF THE SOUTH ATLANTIC OCEAN 93

perhaps be regarded as a nomen vanum (Simpson 1945: 27), since a single tooth

is an inadequate basis on which to identify a cetacean. The monachine tooth

(Frenguelli 1922: 492, fig. 1b-c) is also of low diagnostic value and is here

informally identified as Monachinae A.

In the second part of his paper Frenguelli (1922) assigned to P. rovereti

three teeth which had previously been described and figured by Ameghino (1889)

as those of a ‘creodont’, Apera sanguinaria. This species was identified by

Ameghino (1886) on the basis of two teeth previously identified by Burmeister

(1885) as ‘Ferae’ close to the ‘felid’, Eutemnodus americanus. These two teeth

are not included amongst the three figured by Ameghino (1889) and Frenguelli

(1922), which undoubtedly belong to a monachine, although not necessarily

Monachinae A. They cannot be referred to either Eutemnodus or Apera. The

latter was listed as a junior synonym of the ‘creodont’ Eutemnodus by Trouessart

(1898), a view supported by Marshall (1978), although he included Eutemnodus

in the Borhyaenidae. The three additional teeth referred to P. rovereti by

Frenguelli (1922) are here identified as Monachinae B.

Cabrera (1926) realized that the P. rovereti hypodigm included phocid

material, and noted the existence of a phocid mandible with one tooth which

came from the same deposits as the two teeth described in the first part of

Frenguelli’s (1922) paper. Although Frenguelli (1926) doubted that P. rovereti

was a seal, and although the matter was never clarified by the designation of a

lectotype, this species became established in later literature as a phocid

(e.g. Kraglievich 1934; Kellogg 1942; King 1964).

Hendey & Repenning (1972: 92) referred the far superior material from

Langebaanweg, South Africa, to Prionodelphis, with the identification being

justified by the following statements: ‘In assigning the Langebaanweg phocid

to the genus Prionodelphis, it is recognized that reassessment may be required

when more material of P. rovereti is found. Generic identity is based upon the

remarkable similarity of the few fragments from Argentina to the South African

material and on the belief that the lack of greater knowledge is a stronger

argument against the establishment of a new genus than it is against tentative

assignment to the same genus.’

It is the three teeth of the Argentinian Monachinae B which are most like

their counterparts in the South African species. This applies particularly to the

M! (Frenguelli 1922: 497, fig. 2A), which resembles that of ‘Prionodelphis’

capensis in having a strongly recurved and sharp-pointed principal cusp,

although it is distinct in having a small, anterior accessory cusp closer to the

apex of the principal cusp than any of the South African specimens. The two

lower cheek teeth (Frenguelli 1922: 497, figs 2B-C) resemble their ‘P.’ capensis

homologues in being relatively narrow and with an inflation of the cingulum

posterolingually. They are, however, distinct in having less prominent accessory

cusps which are situated closer to the apices of the principal cusps.

Judging from a cast of the Monachinae A tooth, probably a P*, it differs

from its counterparts in ‘P.’ capensis by being broader, and in having the lingual

94 ANNALS OF THE SOUTH AFRICAN MUSEUM

margin semicircular in occlusal view, rather than having a posterolingual

expansion of the cingulum. In both these respects the Argentinian tooth is

similar to the P* of Monachus monachus. In addition, the teeth of Monachinae A

and M. monachus are relatively high crowned, more so than those of ‘P.’ capensis

and, apparently, Monachinae B.

The situation in respect of material previously assigned to Prionodelphis

rovereti may be summarized as follows:

1. Prionodelphis rovereti is a cetacean, with the first figured specimen (Frenguelli

1922: 492, fig. 1a) here designated as a lectotype.

2. The second tooth figured by Frenguelli (1922: 492, fig. 1b-c) belongs to a

monachine, here designated Monachinae A. It apparently represents a species

closer to Monachus monachus than any other monachine. It is not conspecific,

and may not even be congeneric with ‘Prionodelphis’ capensis.

3. The three teeth of ‘Apera sanguinaria’ described and figured by Ameghino

(1889) and referred to P. rovereti by Frenguelli (1922: 497, fig. 2) also represent

a monachine, which is here designated Monachinae B, since it is not necessarily

conspecific, or even congeneric, with Monachinae A. Monachinae B is, however,

close to ‘P.” capensis, and both probably represent the same genus.

It follows that the South African seal hitherto referred to Prionodelphis

must now be assigned to a new genus.

THE SOUTH AFRICAN SPECIES

Family Phocidae

Subfamily Monachinae

Tribe Lobodontini

Diagnosis

Monachinae characterized by the simultaneous presence of the following

two features:

1. The tympanic bulla covers the petrosal posteriorly.

2. A mastoid lip overlaps the posterior wall of the bulla.

Homiphoca gen. nov.

Type species

Prionodelphis capensis Hendey & Repenning, 1972.

Amended diagnosis

A monachine phocid with a skull superficially similar to that of Monachus.

It differs from Monachus in having a relatively large rostrum, which is wide

posteriorly and narrow anteriorly. As in Monachus, but unlike Lobodontini, the

premaxillae terminate against the nasals, where they are anteroposteriorly

25)

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96 ANNALS OF THE SOUTH AFRICAN MUSEUM

elongated. The premaxillae have prominent tuberosities anteriorly. The

ascending process of the maxilla is relatively high as in Lobodontini and, viewed

anteriorly, is not strongly recurved medially as in Monachus. Dental formula:

2.1.4.1/2.1.4.1. The premolars are morphologically similar to those of Monachus,

and unlike those of Lobodontini. They differ from those of Monachus in being

lower crowned, relatively narrower and in having a pronounced posterolingual

expansion of the cingulum. The accessory cusps on the premolars are small but

distinct, while the M? usually lacks such cusps and is distinct in having a strongly

recurved and sharp, pointed principal cusp. The M, is the largest of the cheek

teeth, with the principal cusp slanted posteriorly, and often with a small

accessory cusp low on the long anterior keel of the principal cusp. The inter-

orbital region is broad and tapers posteriorly as in Lobodon, but unlike all other

monachines. In the auditory region the tympanic bulla covers the petrosal, while

the mastoid forms a lip overlapping the posterior border of the bulla.

The humerus has an entepicondylar foramen, and the tibia and fibula are

fused proximally.

Etymology

From ‘/homi’ and ‘phoca’, which are respectively the Hottentot and Greek

words for ‘seal’. The Hottentots inhabited the Langebaanweg area when

Europeans first settled at the Cape of Good Hope; “/homi’ refers to Arcto-

cephalus pusillus, a member of the family Otariidae (Budack 1977).

Homiphoca capensis (Hendey & Repenning, 1972)

Holotype

SAM-PQ-L15695—an incomplete and partly restored skull with left C and P*,

and right P®.

Paratype

SAM-PQ-L31976—an almost complete skull and mandible, partly restored,

lacking right P?, P4, lower incisors, C, P, and P;, and left lower teeth except

for the lateral incisor and part of P; (Table 1, Figs 1-4).

Referred material

That described by Hendey & Repenning (1972) and many other specimens

in the South African Museum, including:

SAM-PQ-L30080—an almost complete skull, partly restored, with parts of

leit PB toy P=

SAM-PQ-L30568—an almost complete skull and right hemimandible, partly

restored, with right C, P!, P? and Mj, and left P! and P®

SAM-PQ-L32101—an almost complete skull and mandible, partly restored,

lacking left upper lateral incisor, P* and M}, right P, and P,, and left lower

incisors, P,, P, and M, (Fig. 5)

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98 ANNALS OF THE SOUTH AFRICAN MUSEUM

SAM-PQ-L32415—an almost complete skull and left hemimandible, partly

restored, with right upper incisors and P?, left P{ and P?, and parts of

left P,, P3, P, and M,

SAM-PQ-L31278—right hemimandible, partly restored, with P, to M, (Fig. 6)

SAM-PQ-L50304 and others—isolated cheek teeth (Figs 7, 8)

SAM-PQ-L40969—left humerus (Fig. 9)

SAM-PQ-L31957—right ulna lacking distal end (Fig. 10)

SAM-PQ-L40846—right radius (Fig. 11)

SAM-PQ-L30236, L31369—incomplete left innominates (Fig. 12)

SAM-PQ-L30118, L45519—right femora (Fig. 13)

SAM-PQ-L30424—left tibia and fibula, with latter lacking distal epiphysis

(Fig. 14)

Diagnosis

As for genus.

Locality and horizon

The Varswater Formation, ‘E’ Quarry, Langebaanweg, Cape Province.

The material described by Hendey & Repenning (1972) and L40846 and

L40969 are from bed 3aS of the Pelletal Phosphorite Member, whilst most

additional material, including others listed above, are from bed 3aN of the same

member. Some material is also known from the Gravel and Quartzose Sand

Members. (References: Hendey 1976; Dingle et al. 1979; Hendey 1980.)

Age

Langebaanian (latest Miocene/early Pliocene), between 3,5 and 7 Ma.

(References: Hendey 1974, 1976, 1978a.)

Description

The descriptions which follow are confined to a few selected specimens,

mostly collected since 1975, and are intended to supplement the descriptions in

Hendey & Repenning (1972). Most of the material is from bed 3aN of the

Pelletal Phosphorite Member and is, therefore, somewhat younger than that

previously described, which is from bed 3aS. As with some other species common

to these two horizons, there are minor morphological and possibly size

differences between bed 3aS and bed 3aN representatives (Hendey 19785, 1980).

They are, however, too slight to warrant formal nomenclatural recognition, and

simply reflect temporal stages of single species. This matter, as well as a detailed

account of all the Homiphoca capensis material now available, will be the

subjects of future studies.

The skull

Except where otherwise stated the following description is based on the

paratype, L31976. This specimen is essentially similar in all observable respects

to others from bed 3aN.

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100 ANNALS OF THE SOUTH AFRICAN MUSEUM

The skull of Homiphoca is distinct from those of all living Phocidae,

although it is unmistakably monachine and exhibits a combination of characters

found in Monachini (Monachus) and Lobodontini (Lobodon, Hydrurga,

Leptonychotes, Ommatophoca). Superficially it resembles the skulls of Monachus

and Lobodon more than any other monachines.

There are prominent anterior tuberosities on the premaxillae, and in lateral

view the anterior alveolar margin of the premaxillae recedes sharply postero-

ventrally from the apices of the tuberosities towards the crowns of the incisors

and canines. The crowns of these teeth are themselves directed posteroventrally.

A similar situation was observed during this study amongst living monachines

only in M. schauinslandi, although it is much less obvious in this species. In

addition, the premaxillary tuberosities of M. schauinslandi are more widely

separated than in Homiphoca.

The premaxilla/maxilla suture is visible along its entire length in lateral

view, aS in phocines, but unlike the monachines, excluding M. tropicalis, in

which the central part of the premaxillary ascending ramus is within the nasal

aperture. In the phocines the laterally visible part of the ascending ramus is of

constant width, but in Homiphoca that part in contact with the nasals is antero-

posteriorly elongated. The phocine condition is similar to that in all other

carnivores and is apparently the primitive one. Contact between the premaxillae

and nasals is characteristic of Monachus but not Lobodontini.

Homiphoca has two pairs of upper incisors, which is typical of monachines,

but these teeth are small compared with those of living species. They consist of a

principal cusp anteriorly, with an inflated posterior cingulum which is almost in

the form of an accessory cusp. The situation is similar in M. monachus, but not

in M. schauinslandi and living Lobodontini. The lateral incisors are slightly

larger than the medial ones, but the relative size difference is less than in other

monachines, especially the Lobodontini, in which the lateral incisors (and

canines) are enlarged, a specialization for opening breathing holes in sea ice.

The Homiphoca incisors are likely to represent the primitive condition in

monachines.

The roots (and alveoli) of the lateral incisors are oval-shaped in horizontal

cross-section as in living Lobodontini, and unlike those of Monachus, which are

circular. In L31976 the incisors are in a straight line, as in M. tropicalis and

M. schauinslandi, but in other specimens, including the holotype, the incisor row

curves posteriorly, although not as markedly as in M. monachus and the

Lobodontini. In Lobodon, Leptonychotes and Ommatophoca the curvature is less

obvious because of the procumbence of the lateral incisors.

The nasal bones of Homiphoca are elongated as in M. tropicalis and

M. schauinslandi, and, judging from L30568 and L32101, are parallel-sided

anteriorly and sharply tapering posteriorly, terminating above the orbits. The

shape of the nasals in monachines is very variable, but of the Lobodontini those

of Hydrurga are perhaps most like Homiphoca, except that the tapering

commences further anteriorly. In Homiphoca the nasals are relatively wide

LATE TERTIARY SEALS OF THE SOUTH ATLANTIC OCEAN 101

Pa nig

itt

Fig. 4. Lateral view of Homiphoca capensis hemimandible, SAM—PQ-L31976 (paratype).

anteriorly, this being related to the overall width of the snout in this genus (see

below). They are V-shaped anteriorly as in M. tropicalis, and the ethmoid

(osseous nasal septum) is exposed between the branches of the V, unlike the

situation in other monachines.

The ethmoid is a remarkably stout bone, and resembles those in

M. tropicalis, M. schauinslandi and the Lobodontini, although that of Lobodon

is distinct in being recessed within the nasal cavity. In Homiphoca the ethmoid

extends to the posterior end of the nasal cavity. Viewed anteriorly the nasal

aperture of Homiphoca is slightly dorsoventrally elongated, and in this respect

is intermediate between Monachus, in which the aperture is circular or wider

than it is high, and the Lobodontini, in which it is much higher than it is wide.

In Monachus the rostral region is more or less parallel-sided, but in

Homiphoca it is broader posteriorly. The anterior tapering is very marked and

the anterior part of the skull from the premaxillary tuberosities to the posterior

extremity of the jugals is almost V-shaped in dorsal view. In this respect it

differs from all living monachines. This distinctive shape is due largely to

inflation of the maxillae posterolaterally above the three most posterior pairs

of cheek teeth in L31976 and other bed 3aN specimens. This characteristic has

not been observed in living monachines, nor is it found in the H. capensis

holotype. It is evidently due to a greater development of the maxilloturbinals in

the bed 3aN population of Homiphoca. The Lobodontini are also characterized

by well-developed maxilloturbinals, but these are accommodated within the

dorsoventrally expanded nasal cavity, without obvious deformation of the

maxillae externally. The Phocinae also have well-developed maxilloturbinals and

some have a posterolateral expansion of the maxillae similar to the bed 3aN

102 ANNALS OF THE SOUTH AFRICAN MUSEUM

Homiphoca. The possible implications of this development in H. capensis will

be discussed later (see p. 123).

The ascending branch of the Homiphoca maxilla is high and wide, having a

lengthy contact with the nasals. Its anterior margin does not recede posteriorly

as markedly as in other monachines, especially Lobodon. It is, however, more or

less vertical and relatively high as in Lobodontini. By contrast, in Monachus the

ascending branch is low and, viewed anteriorly, is strongly recurved medially,

while in lateral view the snout is flattened anteriorly. These differences are

reflected in the shape of the nasal apertures, and are due to the greater develop-

ment of the turbinals in Lobodontini.

The pre-orbital process is relatively as prominent as that of Hydrurga. In

other monachines it is small or absent. The infraorbital foramen is oval-shaped

TABLE 1

Dimensions of Homiphoca capensis paratype, SAM—PQ-L31976.

Overall length . : 5 é : : : . 270,0

Zygomatic width auenae Woe EY eos, AO SIO)

Mastoid width . ; : 3 x : : . 136,0

Width at supra-orbital processes . : ; . 101,0

Minimum interorbital width . : 5 ‘ - 44,0

Width of premaxillae anteriorly : , . e ssi)

Length of nasals . : Se eae we NN LOSES

Maximum width of nasals , . 4 3 ee0)

Length of incisor row. : PA Mam eI ech on peo 28} 50)

Length of cheek tooth row Abe Paes Belts seal ee ae (S10)

Width of palate between P’’s . 3 z 4 S228

Width of palate between P’s . . . . . 35,3

Width of palate between M?’s . : F ‘ A GBT)

Overall length of mandible wo Be Net SB SO

Height of ascending ramus of mandible. . . 71,0

Lingual

Length Breadth crown

height

Lateral I Sh 7/ 4,4 6,4

Medial I 4,7 3,4 4,8

Cc 9,1 4,4 $2

Ps 8,9 5,9 6,6

Pp? Py) 6,8 7,0

Pp 12,5 6,5 6,6

ips — 6,5 7,2

M? 8,6 Sy9) 7,7

LATE TERTIARY SEALS OF THE SOUTH ATLANTIC OCEAN 103

Fig. 5. Dorsal, lateral and ventral views of Homiphoca capensis skull, SAM—PQ-L32101.

104 ANNALS OF THE SOUTH AFRICAN MUSEUM

as in Lobodon and Hydrurga, but whereas in these genera the long axis is

directed dorsomedially, in Homiphoca it is directed dorsolaterally (Hendey &

Repenning 1972).

In L31976, and other bed 3aN specimens, the palate is concave anteriorly,

but becomes convex on either side of the midline posteriorly. This is also due to

expansion of the maxilloturbinal region of the nasal cavity. Unfortunately the

posterior part of the palate in the H. capensis holotype is lost and it is not known

if it lacked the expansion of this region, as it does the posterolateral expansion

of the maxilla. In all other monachines the palate is either concave or more or

less flat along its entire length. The same apparently applies in the case of

phocines.

The dental arch of Homiphoca is not straight-sided as in Lobodontini, but

diverges posteriorly from the P?’s, making the tooth-rows concave laterally as

in M. tropicalis, and, less so, in M. monachus and M. schauinslandi. Homiphoca

is like Leptonychotes in having a distinct diastema between P* and Mt}, but in

Homiphoca the gap between these teeth is largely filled by M, when the jaws are

closed, whereas in Leptonychotes the small M, comes into contact only with P?.

The diastema between C and P! is variably developed in Homiphoca. For

example, a small diastema is present in L31976 and L32101, but it is absent in

the holotype, L30080 and L30568.

The C of Homiphoca is a relatively small and low-crowned tooth, which is

circular in cross-section and with the crown recurved. Except perhaps for the

recurvature it probably represents the primitive condition in monachines.

In terms of their basic morphology the upper premolars of Homiphoca are

similar to their homologues in Monachus, and are very different from the

specialized teeth of Lobodontini. They are, however, more slender than the

premolars of Monachus. In addition, the P? to P* are expanded posterolingually

and thus taper anteriorly, whereas in Monachus the lingual margin is more or

less semicircular in occlusal view. The P* is Monachus-like, but as with the other

premolars it is lower crowned. The premolars have a prominent principal cusp,

and accessory cusps anteriorly and posteriorly, with P* to P* usually having a

second, cingular cusp posteriorly. There is no cingulum buccally, but a well-

developed one lingually. There is sometimes a small tubercle on the expanded

posterolingual part of the cingulum (Fig. 7B). In this respect Homiphoca

resembles Lobodon and Hydrurga, which may also have a small accessory cusp

in the same position. It is not known in other monachines.

The premolars tend to wear horizontal facets on the principal cusps, which

are later obliterated by sloping facets anteriorly. A similar sloping facet is

sometimes present posteriorly. The wear on the premolars, especially P? to P*,

indicates that these teeth had a crushing function like those of Monachus, but

unlike those of Lobodontini. The premolars of Homiphoca and Monachus

evidently represent the primitive and unspecialized condition in monachines.

There are two reasons for believing that these teeth are ‘primitive’. Their

crushing function suggests that molluscs and crustaceans formed part of the diet,

105,

LATE TERTIARY SEALS OF THE SOUTH ATLANTIC OCEAN

“SLCIE

T-Od-WVS ‘o[qipueunutey sisuadvo vaoyduuopzy JO MIA [eINCT 9 “317

106 ANNALS OF THE SOUTH AFRICAN MUSEUM

and such prey requires a lesser swimming ability on the part of predators than

fast-swimming fishes and planktonic invertebrates. Greater swimming ability is

obviously an advanced characteristic in phocids. Secondly, molluscs and larger

crustacea occur in the littoral environment, the likely habitat of primitive

phocids which had not yet adapted to a more pelagic way of life.

The M! of Homiphoca is distinguished from the premolars by having a

strongly recurved and sharp-pointed principal cusp, and in having accessory

cusps reduced or absent. A small accessory cusp is sometimes developed

anteriorly. This tooth is ovate in occlusal view.

As in other monachines the palatines of Homiphoca become fused to the

maxillae, and are well developed, reaching anteriorly to be in line with the M?’s.

The palatine foramen, through which the maxillary artery passes, is at the

contact between the palatine and maxilla, whereas in all living monachines it is

situated further anteriorly in the maxilla. In Homiphoca there is a groove for the

maxillary artery passing anteriorly from the palatine foramen close to the

lingual alveolar margin.

The intra-orbital part of the palatine is thick and the medial wall of the

orbit is almost complete, as in Hydrurga. In other living Lobodontini this wall

has large lacunae, which are reduced with age. This is apparently an advanced

condition. The posterior border of the palatines, that is, the lower openings of

the secondary choane, are oval in shape and resemble those in Lobodon and

Hydrurga, rather than those of Leptonychotes and Ommatophoca.

The pterygoids of L31976 are poorly preserved, but it is evident that the

pterygoid apophyses were small and that the lateral walls of the choane are

nearly vertical as in Hydrurga.

The inter-orbital region is broad and tapers posteriorly, as in Lobodon, but

in marked contrast to the condition in other monachines, notably Monachus and

Leptonychotes in which the inter- and post-orbital regions are narrow and

parallel-sided. The latter condition is apparently the primitive one.

As noted by Hendey & Repenning (1972), the jugal terminates anteriorly

above the lateral border of the infra-orbital foramen in Homiphoca, above the

centre of this foramen in Monachus, and lateral to this foramen in Lobodontini.

Homiphoca is thus intermediate between the two groups of living monachines in

this respect. The posterior end of the jugal is bifurcated, with a narrow dorsal

branch and a broad ventral one, thus resembling Monachus in this respect, but

differing from the Lobodontini.

The zygomatic process of the squamosal is short, with a pronounced dorsal

inflection, while the glenoid fossa is narrow and deep. In both these respects

Homiphoca is more like Monachus than Lobodontini.

The auditory region of the paratype is virtually identical to the one described

in detail by Hendey & Repenning (1972). Other references to the ear region of

Homiphoca are to be found in Ray (1976b) and Repenning & Ray (1977). A few

additional observations are made here.

As noted by King (1966), the bulla covers the petrosal posteriorly in the

LATE TERTIARY SEALS OF THE SOUTH ATLANTIC OCEAN 107

Fig. 7. Homiphoca capensis premolars. A. Occlusal view of upper left premolar, SAM—PQ-—

L55047B. B. Occlusal and anterior views of upper right premolar, SAM—PQ-I_50304D.

C. Lingual view of upper right premolar, SAM—PQ-L55046. D-—F. Buccal views of upper right

premolars, SAM—PQ-L55047C, 50304C, 50304A. G-—I. Lingual views of lower right premolars,

SAM-—PQ-L55047A, L55047D, L50304B.

ANNALS OF THE SOUTH AFRICAN MUSEUM

108

"VSP0SST-Od-NVS "H “aSp0SsT-Od-WVS “D ‘OSb0SsT-Od-WVS “A

AsSpOssT-Od-WVS “A ‘SOW (H ‘A) Wer pure (D ‘d) Is Jo smola yensuryT “H-A “WrbOSST-Od-NVS ‘A ‘OPr0ssT-Od-WVS ‘OD

arrossT-Od-WVS “& ‘vL60ST-Od-WYS “V

SIN (CG) eT Pur (O-V) SIT JO SMOIA ensury “q-Vy ‘sievjour sisuadva pIOydIMoFT *g “Sty

eo Sea

wd |

LATE TERTIARY SEALS OF THE SOUTH ATLANTIC OCEAN 109

i eit rani Wt ot

unui nnn

nn i

Fig. 9. Anterior, lateral and posterior views of Homiphoca capensis humerus, SAM—PQ-L40969.

110 ANNALS OF THE SOUTH AFRICAN MUSEUM

Lobodontini, whereas in Monachus the posterior part of the petrosal is exposed

in ventral aspect. In Homiphoca the situation is intermediate, with the posterior

extension of the bulla clearly less than in Lobodontini, and not covering the

whole of the petrosal. In addition, the mastoid has a lip overlapping the

posterior wall of the bulla. This condition is typical of the Lobodontini and its

presence in Homiphoca, together with the posterior development of the bulla, is

here regarded as good evidence of its relationship to this group (Hendey &

Repenning 1972).

The bulla in Homiphoca is small and little inflated as in Monachus. The

carotid foramen is located anteriorly as in Lobodon and Ommatophoca, but less

so than in Hydrurga and Leptonychotes. The petrosal has been completely

exposed in several Homiphoca specimens, showing that the promontorium is

better developed than in Monachus, but less so than in Lobodontini. The apical

whorls of the cochlea are visible and are not completely hidden by the consider-

ably expanded basal whorls as in Lobodon and Leptonychotes. Once again the

condition is intermediate between those in Monachus and Lobodontini (see

Repenning & Ray 1977).

The basisphenoid is narrow as in Hydrurga, while the alisphenoid exhibits

a well-developed lateral process as in Monachus. The basioccipital is relatively

narrow and trapezoid in shape. In Monachus, Lobodon, and Leptonychotes the

basioccipital is of similar shape but wider, while in Hydrurga and Ommatophoca

it is narrow and sometimes rectangular.

In L31976, and some other Homiphoca specimens, there is a basioccipital

foramen situated slightly posteriorly to, and medial of, the carotid foramen. On

four of the specimens this foramen is situated at the basioccipital/bulla suture,

while in two it is in the basioccipital itself. In spite of its variable position, it is

apparently always present in Homiphoca, and may thus be characteristic of this

taxon. It was otherwise observed in the present study only in three out of five

Hydrurga skulls. In Leptonychotes there is a partially isolated basioccipital

foramen at the anteromedial corner of the posterior lacerate foramen which may

be homologous. The function of this foramen is not known. It may have accom-

modated a branch of the internal carotid artery, or, perhaps more likely, it may

represent a branch of the ventral venous petrosal sinus. This foramen may be a

primitive characteristic.

The exoccipitals carry triangular paroccipital processes which resemble

those of Monachus and Hydrurga, but differ from the low crests.observed in

Leptonychotes. In Lobodon and Ommatophoca the crest is higher, but the process

is not triangular.

The occipital condyles are similar to those of living monachines, while the

foramen magnum is oval, which is commonly the case in living Lobodontini,

whereas in Monachus it is usually circular.

The occipital crest is V-shaped as in Lobodon, Leptonychotes and Monachus,

and not U-shaped as in Hydrurga and Ommatophoca. The braincase is relatively

smaller than those of living Lobodontini, and similar to that of Monachus.

LATE TERTIARY SEALS OF THE SOUTH ATLANTIC OCEAN 111

nn a EL

| j

“a

Fig. 10. Medial view of Homiphoca capensis ulna, SAM-—PQ-L31957.

ania UU LLL

9 Z

0 2

6 1

nar

(lll

Hill

Fig. 11. Medial view of Homiphoca capensis radius, SAM—PQ-L40846.

112 ANNALS OF THE SOUTH AFRICAN MUSEUM

Because the interorbital region is relatively broad, the anterior limit of the

braincase is not as sharply defined as in Monachus and Leptonychotes. The

anterior curvature of the braincase in dorsal view is gradual as in living

Lobodontini. The sagittal crest is reduced as in Lobodon and Leptonychotes.

The Homiphoca skull is relatively narrow posteriorly, resembling those of

Lobodon and Hydrurga more than other monachines in this respect (Fig. 15,

Table 2).

The mandible of Homiphoca was previously described on the basis of a

specimen lacking the teeth and those parts posterior to the cheek teeth, but

largely complete specimens, many with one or more teeth in position, are now

known. In general, the mandible is similar to that of Monachus and very different

from those of the highly specialized Lobodontini.

The symphysis is short and terminates below the middle of P,. In Lobodon

and Ommatophoca it is much longer, reaching to below the anterior extremity of

P,, while in Monachus, Hydrurga and Leptonychotes it reaches to below the

posterior extremity of P,.

The horizontal ramus is low and narrow, and of constant height beneath

the cheek teeth, much as in Monachus. The ascending ramus is also Monachus-

like, with a very narrow coronoid process and large angular region, which gives

it a rather square shape in lateral view. The condition in Hydrurga and Ommato-

phoca is similar, but these genera differ in having high condyles and much

TABLE 2

Mean dimensions and ratios of Lobodontini skulls.

Lobodon Hydrurga Homiphoca Leptony- Ommato-

carcino- leptonyx capensis chotes phoca

phagus weddelli rossi

N 11 6 3 10 4

1. 292 368 258 271 239

2: 98 116 87 65 47

3h. 157 187 130 177 167

4. 87 105 72,5 86 81

PL I) (3535) 0,315 0,337 0,239 0,196

4:3 0,554 0,561 0,557 0,485 0,485

N—Number of specimens (South African Museum collections).

1.—Overall length of skull.

2.—Length of snout from anterior extremity of premaxilla to anterior end of

jugal.

3.— Height of occiput from basioccipital to top of occipital crest.

4.—Mastoid width.

“69ETET PUL 9EZOET-Od-VS ‘SoveuTUIOUUL sisuadvo voOYdUOFT JO SMOIA [eIOYET *Z] “BIA

113

LATE TERTIARY SEALS OF THE SOUTH ATLANTIC OCEAN

114 ANNALS OF THE SOUTH AFRICAN MUSEUM

smaller coronoid processes. The condyle is Monachus-like in being low and

narrow, while the masseteric fossa is shallow and generally similar to that in

Monachus.

Only three intact teeth remain in the paratype mandible, but other speci-

mens have more complete dentitions, and hundreds of isolated teeth are

available.

The lower incisors are small, nondescript teeth. The medial incisor is the

smaller of the two, and is slightly more posteriorly situated. It lies almost

horizontally in the mandible, whereas the lateral incisor is more vertically

inclined. Both have a small posterior cingulum, as in Monachus. The C is small,

circular in cross-section and with a slightly recurved crown. The alveoli of the

incisors and C merge with one another at the alveolar margin.

The lower premolars are morphologically similar to the uppers, having a

principal cusp, one anterior and two posterior accessory cusps. The lingual

cingula are less pronounced than in Monachus, and in this respect Homiphoca is

intermediate between Monachus and living Lobodontini. The lower premolars

are narrower than the uppers and their homologues in Monachus. The postero-

lingual cingula of P, to P, are slightly inflated.

The M, of Homiphoca is an unusual tooth, being unlike that of any

previously recorded phocid. It has a large crown which is triangular in lateral

view, and which is an elongated oval in occlusal view. The apex of the principal

cusp is directed slightly posteriorly, with the result that the anterior keel is

longer than the posterior one. Small anterior and posterior accessory cusps are

sometimes present, the anterior one being the larger and situated slightly

higher on the crown. A lingual cingulum, which may extend around the anterior

end of the tooth, is present. It is similar to the M, of Lobodon in being larger than

P,. In Monachus and in other Lobodontini the M, is smaller than P,. The

Homiphoca M, is also unlike other double-rooted teeth of this taxon, and of

living monachines, in having the roots converging towards their extremities.

The postcranial skeleton

Most elements of the postcranial skeleton of Homiphoca capensis are now

available for study. Vertebrae, ribs, scapulae and innominates are generally

incomplete, but most, if not all, limb bones are represented by several intact and

well-preserved specimens. A vast number of incomplete limb bones are known.

Curiously, in view of the large number of additional Homiphoca specimens

now available, the incomplete scapula described by Hendey & Repenning (1972)

is still one of the best specimens of this bone. A few supplementary observations

are possible. The lower half of the posterior border of the blade of the scapula is

triangular in cross-section as in Monachus, while in Lobodontini it is always

rounded. The latter is an advanced condition which is discussed in more detail

elsewhere (De Muizon 1979). The depression for the insertion of the triceps

brachii on the posterolateral part of the neck is deep, indicating that this muscle

was more powerfully developed than in living monachines

115

iti

ht iil

145519.

LATE TERTIARY SEALS OF THE SOUTH ATLANTIC OCEAN

Anterior and posterior views of Homiphoca capensis femora, SAM-—PQ-L30118 and

Fig.

116 ANNALS OF THE SOUTH AFRICAN MUSEUM

The previously described humerii were incomplete, but several intact

specimens are now known. In most living Lobodontini the greater trochanter is

lower than the lesser trochanter. In some Monachus specimens they are of equal

height, while in others the greater trochanter is higher, as in most carnivores.

The latter is the primitive condition and is also found in late Miocene Mono-

therium of the North Atlantic and some of the phocids from the Pisco Formation

in Peru (De Muizon 1979). In Leptonychotes and Homiphoca the development

of the trochanters is intermediate between the primitive condition and the

advanced one in other living Lobodontini.

Similarly, the lateral surface of the deltoid crest (from the greater trochanter

to the deltoid tubercle) is more elongated in Homiphoca than in Monachus, and

more like that in Monotherium and living Lobodontini. The prominent deltoid

tubercle and strong relief of the posterolateral side of the deltoid crest indicate

the existence of stronger brachialis and brachioradialis muscles in living

Lobodontini than in Monachini.

The presence of a well-developed supinator ridge and entepicondylar

foramen in the Homiphoca humerus is characteristic of phocines rather than

monachines, and they are evidently primitive features. On the other hand, the

deltoid/pectoral crest reaches the distal epiphysis in Homiphoca, and this is a

typically monachine feature. The bicipital groove appears to be deeper and

narrower than in most living monachines.

A comparison of the general morphology of the humerus of Homiphoca

with those of Monachus and Leptonychotes suggests that it represents a primitive

stage in the development towards the Leptonychotes (i.e. Lobodontini) con-

dition. Particularly significant are the curvature in lateral view (it is straight in

Monachus), size of the lesser trochanter and elongation of the muscle insertion

area on the lateral side of the deltoid crest. In all these respects Homiphoca is

closer to Leptonychotes than to Monachus.

The previously described ulnae represented the entire bone except for the

tubercle for insertion of the anconeus medialis muscle. This tubercle is preserved

in several new specimens, and, as in the Phocinae, it is very prominent. In this

respect the ulna of Homiphoca differs from those of living monachines. The ulna

of Monotherium ? wymani (Ray 19765) is similar to that of Homiphoca, and they

evidently represent the primitive condition.

Complete radii are now known. The radius is very wide distally, as in

Monachus and Lobodon. This represents a primitive condition relative to other

Lobodontini, particularly Leptonychotes (De Muizon 1979). The Homiphoca

radius differs from those of Hydrurga and Ommatophoca in that the area for

insertion of the pronator teres is very pronounced in the latter genera.

No complete innominates are known, and the best available specimen is

probably that described by Hendey & Repenning (1972). This bone is typically

monachine. The pectineal tuberosity was examined in ten specimens, and found

to be reduced in seven, as in living Lobodontini, while in the others it is very

prominent as in Monachus and the Phocinae. The psoas minor inserts on this

—

LATE TERTIARY SEALS OF THE SOUTH ATLANTIC OCEAN

wii

Whi

tyr

ian Il

Fig. 14. Anterior and posterior views of Homiphoca capensis tibia and fibula,

SAM-PQ-L30424.

118 ANNALS OF THE SOUTH AFRICAN MUSEUM

tuberosity and it has the function of bending the back in the caterpillar-like

terrestrial locomotion of seals. The stronger this muscle, the more terrestrial the

species concerned is likely to be, and a prominent pectineal tuberosity may thus

be interpreted as a primitive character. Homiphoca would thus have been more

primitive than living Lobodontini in this respect, but more advanced than

Monachini.

The femur of Homiphoca was previously described on the basis of a single

distal fragment, but complete specimens are now known. This bone is short and

wide as in living Lobodontini, but the head is more spherical and the neck is

more distinct. In the latter respects it is Monachus-like. King (1966) recorded

that the phocine femur was distinct from that of monachines in having a deep

trochanteric fossa, a high trochanter and a pronounced popliteus pit. There are,

however, exceptions amongst both monachines and phocines. For example,

Lobodon has a deep trochanteric fossa, while in some Phocinae (e.g. Erignathus)

it is absent. Homiphoca also has a trochanteric fossa, and, in addition, a

well-developed popliteus pit. The trochanter is variably developed, some-

times being higher than the head as in the Phocinae, and sometimes lower as in

the Lobodontini and M. schauinslandi. Homiphoca is probably most like

M. monachus in this respect. In the Peruvian fossil monachines the trochanter is

higher than the head and the popliteus pit is marked.

The anterior surface of the Homiphoca femur shaft has a marked concavity

mediodistally. A similar concavity is often present in Lobodontini, but it is less

pronounced in Monachus. The patella facet is transversely elongated as in

Lobodontini. In the phocines this facet is dorsoventrally elongated. The area of

insertion of the peroneus longus on the lateral epicondyle is very pronounced

and visible in anterior view as in other Monachinae, whereas in Phocinae

(excluding Erignathus) it is orientated laterally.

Although the phocid femur is more variable and less diagnostic than, for

example, the humerus, the typically monachine Homiphoca femur is in some

respects intermediate between those of Monachus and living Lobodontini. Of

the latter it is perhaps closest to Lobodon because both have a deep trochanteric

fossa.

The tibia and fibula of Homiphoca are fused proximally as in almost all

living and fossil phocids. These bones are known to be articulated proximally

only in M. schauinslandi (Ray 1976a) and a small monachine from the Pisco

Formation in Peru.

The proximal tibial facets are usually markedly concave in the Phocinae and

the tibial spine is high, while in living Monachinae the facets tend to be flat and

the spine is low. The Homiphoca tibia is intermediate in these respects.

One of the most striking features of the Homiphoca tibia is the presence of

very deep tibial fossae (Hendey & Repenning 1972). The posterior one extends

along the proximal two-thirds of the shaft, and the anterior one along the

proximal half of the shaft. A deep posterior fossa in seal tibiae indicates strong

leg musculature. The Homiphoca tibia differs from those of living Lobodontini in

LATE TERTIARY SEALS OF THE SOUTH ATLANTIC OCEAN 119

0,35 06

H. c

L.c

H. | HI

H.c

0,3 L.c.

0,25 0,5

L.w. L.w. & Or

0. r. oe

015 0,4

A B

Fig. 15. Ratios of Lobodontini skull dimensions. A. Snout length:

overall length. B. Occiput height: mastoid width. Data from

Table 2. (H.c.— Homiphoca capensis, L.c.—Lobodon carcinophagus,

H.1.—Hydrurga leptonyx, L.w.—Leptonychotes weddelli, O.1.—

Ommatophoca rossi.)

being relatively short, and in terms of femur-tibia/fibula length proportions,

Homiphoca is closer to Monachus. Possibly the more powerful musculature

inserted on the tibia compensated for its relative shortness.

The anteroposteriorly flattened distal end of the tibia is similar to that of

Pliophoca of the Italian Pliocene (Ugolini 1902; Tavani 1942).

The sharply angled distal fibula facet of the tibia led Hendey & Repenning

(1972) to suggest that the fibula, which was then not known, must have been

markedly bowed. In fact, the fibula is no more bowed than that of Monachus,

although it is more so than in Lobodontini, in which the fibula is almost

straight. The Homiphoca fibula has a small lateral recurved extension to the

astragalus facet which articulates with the calcaneum. This facet is pronounced

120 ANNALS OF THE SOUTH AFRICAN MUSEUM

in the Lobodontini and Mirounga, but is very reduced in Monachus and the

Phocinae.

Most, if not all, elements of the Homiphoca manus and pes are now repre-

sented by complete specimens, but they are not described here.

DISCUSSION

It is abundantly clear that Homiphoca capensis is a member of the subfamily

Monachinae, and in many respects is a morphological intermediate between

living Monachini (Monachus) and Lobodontini (Hendey & Repenning 1972).

Monachus is widely recognized as the least specialized of living Monachinae,

while the Lobodontini are amongst the most highly specialized of all phocids.

Hendey & Repenning (1972: 95) have already suggested that H. capensis is

more specialized than Monachus and that in a ‘broad sense’ its relationships lie

with ‘the Antarctic monachines’ (i.e. Lobodontini), although ‘it is not clearly

ancestral to any of the four living genera’. Subsequently it was suggested that

H. capensis is more closely related to Leptonychotes and Ommatophoca than to

Lobodon and. Hydrurga, although it was probably not directly ancestral to either

of the former (Hendey 1972). The present study has led to a revision of this

opinion.

The informal separation of the Lobodontini into two groups, namely,

Leptonychotes and Ommatophoca on the one hand, and Lobodon and Hydrurga

on the other, is based in part on the following characters:

1. The highly specialized cheek teeth with well-developed accessory cusps of

Lobodon and Hydrurga contrast with the reduced teeth of Leptonychotes and

Ommatophoca, in which accessory cusps are absent in the former, and very small

or absent in Ommatophoca.

2. The general development in Lobodon and Hydrurga of posterolingual cusps

on the upper cheek teeth, which are absent in all other living monachines.

3. The molars (Mj) are well developed in Lobodon and Hydrurga, but are

reduced in Leptonychotes and Ommatophoca.

4. The long snout in Lobodon and Hydrurga contrasts with the shortened one

in Leptonychotes and Ommatophoca (Fig. 15).

5. The relatively high occiput in Lobodon and Hydrurga contrasts with the low

occiput in Leptonychotes and Ommatophoca (Fig. 15).

The earlier opinion that Homiphoca was more closely related to

Leptonychotes/Ommatophoca was based on the belief that while it would be

possible for the teeth of the latter to evolve from those of Homiphoca, the M?

of the latter was already more advanced than those of Lobodon and Hydrurga.

The present study has suggested that this was not necessarily the case. In

addition, there is other evidence which indicates that the relationships of

Homiphoca lie rather with the Lobodon/Hydrurga group. .

The most significant characteristics which Homiphoca shares with Lobodon/

Hydrurga are as follows:

LATE TERTIARY SEALS OF THE SOUTH ATLANTIC OCEAN 121

1. The posterolingual expansion of P? to P*, sometimes with a small accessory

cusp.

2. The well-developed M,.

3. The relatively long snout (Fig. 15).

4. The relatively high occiput (Fig. 15).

At first sight the fact that the Homiphoca M? is relatively smaller than those

of Lobodon and Hydrurga suggests that the former is unlikely to be closely

related to either of these Antarctic genera. It is obvious that the M* of the most

primitive Phocidae must have been large, and that the general trend in phocid

evolution has been towards reduction of this tooth. Amongst living monachines

it is only in Lobodon and Hydrurga that M! is similar in size, or only slightly

smaller than P*.

All the teeth of Lobodon and Hydrurga are larger and more highly specialized

than those of Monachus, which, except for their breadth, may well represent

something approaching the condition typical of late Tertiary monachines. Since

the M! of Monachus is relatively small, it is possible that the large size of this

tooth in Lobodon and Hydrurga is a secondary development accompanying the

general specialization of their dentitions. In other words, the large size of M? in

these genera may be a specialized rather than primitive condition.

Another factor which suggests that the enlargement of the M! in Lobodon

and Hydrurga was secondary is the parallel enlargement of P! in these genera.

Both these teeth thus reflect a development towards homodonty, a condition

which is characteristic of many marine mammals.

If the relatively large size of M! in Lobodon and Hydrurga is indeed a

secondary specialization, then the size of this tooth in Homiphoca does not

necessarily exclude it from being closely related to Lobodon/Hydrurga.

It may also be significant that the crown of the Homiphoca M, is, as in

Lobodon, relatively higher than in any other known monachines. The fact that

the Homiphoca M, was already enlarged relative to those of monachines other

than Lobodon and Hydrurga may foreshadow the enlargement of M? in its

descendants. The M! of Lobodon and Hydrurga is the least functional tooth in

the dentitions in the sense that it alone is.in contact with only one other tooth

(M,). For this reason its enlargement may have lagged behind that of M, and P;

in the Lobodon and Hydrurga lineages.

The most striking aspect of the cheek teeth of Lobodon and Hydrurga is

their highly specialized, comb-like cusps. It is therefore of interest to consider

the manner in which monachine cheek tooth cusps might have evolved.

It was stated earlier that the cheek teeth of Monachus, a genus which is in

almost all respects the least specialized of living monachines, are likely to

resemble those of primitive members of the group. Monachus cheek teeth are

comprised of a principal cusp, and, depending on species and tooth concerned,

either no accessory cusps, one small posterior accessory cusp, or one small

accessory cusp anteriorly and posteriorly. In those genera which are supposedly

close to the origins of the Phocidae, such as Paragale and Potamotherium

122 ANNALS OF THE SOUTH AFRICAN MUSEUM

(Savage 1957; Tedford 1976), the premolars (excluding P*) have a principal cusp

and reduced or absent accessory cusps, with never more than one of the latter

anteriorly and posteriorly. It is, therefore, possible that primitive monachines

were characterized by reduced or absent accessory cusps on their cheek teeth.

It is worth noting in this connection that in the Otariidae accessory cusps are

interpreted as an advanced character (Repenning & Tedford 1977: 66).

Late in the Tertiary there appears to have been a general tendency amongst

phocids to develop accessory cusps, at least on the premolars, especially P? to P4.

Subsequently, amongst the monachines different lineages evolved their cheek

teeth in different ways. In Monachus there was probably little change in the

teeth, just as the rest of the skull and postcranial skeleton remained unspecialized.

In the Leptonychotes, Ommatophoca, and Mirounga lineages the cheek teeth were

reduced and the accessory cusps were often lost. In the case of Ommatophoca it

is known that during the early Pleistocene there was still the basic three-cusped

pattern on P, to P, at least (King 1973). The Lobodon and Hydrurga lineages

retained, and in the case of the former, even supplemented the three-cusped

pattern on P? to Pi, with individual cusps greatly enlarged and morphologically

modified, while P} and Mj evolved to match the characteristics of P3 to P34.

In Homiphoca the cusp number on individual teeth is variable, but the

situation may be summed up as follows:

1. P{ have a well-developed posterior accessory cusp, and sometimes a small

anterior and a second posterior accessory cusp as well. (Fig. 7C)

2. P32 to Pi have well-developed anterior and posterior accessory cusps, and

sometimes a second small posterior accessory cusp. (Figs 7D-I)

3. M, sometimes has a posterior accessory cusp, and less often an anterior

accessory cusp. (Figs 8E-H)

4. M! sometimes has a small anterior accessory cusp. (Figs 8A—D)

In addition, a small posterolingual cusp on P? to P* may be present (Fig. 7B).

Since Homiphoca cheek teeth have better developed, and sometimes also a

greater number of accessory cusps than Monachus, it is possible that it belongs

in that group of Lobodontini in which cheek tooth cusps are enlarged and well

developed, that is, Lobodon/Hydrurga. The fact that it sometimes had one more

cusp than Hydrurga, that is, the second posterior accessory cusp on P} to P34,

which is the same in some Lobodon individuals, suggests that its affinities lie

rather with the latter. However, Lobodon often has a third posterior accessory

cusp on P} to P#, as well as one or two more cusps on its molars than Homiphoca.

Presumably if Homiphoca did belong to a lineage in which cusps were being

developed, it could have achieved the Lobodon condition in the lengthy time

available.

There are other aspects of the accessory cusps in Homiphoca cheek teeth

which suggest a possible connection with Lobodon. In the latter, those accessory

cusps immediately adjacent to the principal cusp have their apices only a little

below the level of the apices of the principal cusps. The homologous cusps in

Homiphoca may also be relatively high on the keels of the principal cusps,

LATE TERTIARY SEALS OF THE SOUTH ATLANTIC OCEAN 123

although they are never separated from the principal cusps as in Lobodon. This

variation in position is particularly noticeable on M}, in which the anterior

accessory cusps, when present, vary from being low on the anterior keel, at or

near the cingulum, to a little way above it (Fig. 8). The significance of this is that

when an accessory cusp is ‘shifted’ up the keel, it is then possible for an additional

accessory cusp to develop from the cingulum anteriorly and posteriorly. It is

apparently always from these positions that the supernumerary cusps of

Lobodon cheek teeth are developed.

In addition, the supernumerary cingular cusps, like other accessory cusps

in Lobodon, are curved in the direction of the principal cusp, a tendency which

is apparent in Homiphoca. The best example of this in Homiphoca is in the

isolated upper premolar, L50304A, in which there is a very pronounced anterior

recurvature of the second posterior accessory cusp (Fig. 7F). This condition is

not known in any other monachine.

Of course, the earlier alternative hypothesis that Homiphoca is more

closely related to those Lobodontini in which teeth and accessory cusps are

reduced or absent (i.e. Leptonychotes|Ommatophoca) cannot be dismissed.

However, this alternative has no other compelling evidence to support it,

whereas there are other characteristics which suggest a close relationship

between Homiphoca and Lobodon (see p. 121). In addition, and perhaps most

significantly, Homiphoca and Lobodon are similar in having a broad inter-

orbital region, a characteristic which distinguishes them from other monachines.

In general the skulls of Homiphoca and Lobodon are similar in morphology

and that of the former is only slightly smaller. The most striking differences are

in the nasal region. In Homiphoca the nasals themselves are long, and the

chambers occupied by the maxilloturbinals are voluminous, particularly in

bed 3aN specimens in which there is deformation of the snout laterally and

ventrally (see p. 101). In Lobodon the nasals are short, and the maxilloturbinals,

although large and complex, are confined to the inter-orbital region. This

contributes at least in part to the relatively broad post-orbital region of Lobodon.

The development of the maxilloturbinals is directly related to ambient air

temperatures, since their mucosa serve to warm and moisten inspired air

(Miller et al. 1964). The apparent differences in maxilloturbinal development

between bed 3aS and bed 3aN Homiphoca populations may be related to the

initiation of, or fluctuations within the major cooling of the late Tertiary. This

cooling led to the cold upwelling within the Benguela Current System and

consequent lowering of ambient air temperatures off the west coast of South

Africa (see Siesser 1978).

It is thus likely that the maxilloturbinal development in bed 3aN Homiphoca

was in an early stage of specialization. Refinement of the arrangement of the

maxilloturbinals, perhaps by greater convolution, may have contributed to the

marked difference in the nature of this region of the Lobodon skull.

The nasal cavity of Lobodon is also distinct in having two pronounced

lateral fossae anterior to the maxilloturbinals. Their purpose is not known, but

124 ANNALS OF THE SOUTH AFRICAN MUSEUM

they are presumably an advanced character related to nasal physiology. They

have not been observed in other monachines, and they too contribute to the

marked differences between the nasal regions of Homiphoca and Lobodon.

This, and other, differences between Homiphoca and Lobodon may all be

interpreted as greater specialization in the latter, with adaptations being for

different feeding habits, better aquatic locomotion, and life in a frigid climate.

There appears to be no character which would preclude Homiphoca from being

closely related to Lobodon. It does not necessarily follow that Homiphoca was

directly ancestral to Lobodon, since it could equally have belonged on a separate

lineage which paralleled that of Lobodon in some respects.

The origins of Homiphoca are obscure because of the extremely poor phocid

record earlier in the Miocene. The best known of early phocids is Monotherium.

Two late Miocene species are recorded in Belgium, namely, M. aberratum and

M. affine (= M. delognii), while a middle Miocene species from North America

is tentatively identified with this genus (M. ? wymani). The auditory region of the

latter is known, and it has the mastoid lip overlapping the posterior wall of the

bulla, a characteristic of Lobodontini. Although available evidence is slender, it

is possible that Monotherium includes an ancestor of Homiphoca.

Other middle to late Miocene phocids are either not well enough known to

be certain of their affinities, or obviously belong to groups other than the

Lobodontini. Prophoca rousseaui from the middle Miocene of Belgium was

referred by Ray (1976a) to the Phocinae, but its humerus appears to be typically

monachine, and its relationships have yet to be firmly established by the

discovery of additional material. Callophoca evidently is a monachine, but it is

not relevant here since it is related to Monachus or Mirounga (Ray 1976a).

The same applies to Pliophoca. The Paratethyan seals are very problematical,

but they, too, are probably irrelevant to the history of the Lobodontini.

In conclusion some observations are made on the past distribution and

dispersal of monachine seals.

It is almost certain that the Monachinae had their origins in the North

Atlantic Ocean, and perhaps that the ancestors of Homiphoca reached the South

Atlantic and South Africa by following the route suggested earlier by Hendey

(1972). This involved the dispersal of European monachines southwards along

the north-west coast of Africa, across the north equatorial region of the Atlantic,

down the east coast of South America, and back across the Atlantic in southern

mid-latitudes, with the oceanic crossings being facilitated by major current

systems. The latter were probably particularly significant in the case of the

southerly dispersal of seals in the South Atlantic. The west to east route from

South America to South Africa by way of subantarctic islands, which follows

the prevailing current system in southern mid-latitudes, was used first by

monachines and later by otariids. The latter must have entered the South

Atlantic from the Pacific round the southern tip of South America, since early

in their history they were confined to the Pacific Ocean.

It is possible that South Atlantic monachines took the same route followed

LATE TERTIARY SEALS OF THE SOUTH ATLANTIC OCEAN 125

later by otariids. The time when monachines first entered the Pacific is not

known, but it could have been as much as 15 m.y. ago (Repenning & Ray 1977).

The older monachines of the Pisco Formation in Peru may be of late Miocene

age and are the earliest known Pacific representatives of their subfamily. They

clearly represent taxa distinct from Homiphoca, and their age relative to the

latter is not certain.

It is also possible that South Atlantic monachines migrated along a more

direct route southwards, either along the east coasts of the Americas, or along

the west coast of Africa. The direction of major current systems does not

necessarily directly influence the movements of seals along coastlines, whereas

they are of paramount importance in oceanic crossings.

Although the last-mentioned alternative is not favoured, it, and the others,

will remain possibilities until more relevant material from the regions in question

is collected and studied.

SUMMARY

The status of recorded late Tertiary seals of the South Atlantic Ocean is

revised. Prionodelphis rovereti from the late Miocene/early Pliocene of Argentina

was identified by Frenguelli (1922) on the basis of five teeth belonging to a

cetacean and one, or possibly two, monachine seals (Phocidae, Monachinae).

P. rovereti is regarded as a cetacean. Prionodelphis capensis Hendey & Repenning,

1972, from South Africa is assigned to a new genus, Homiphoca.

A nearly complete skull and mandible, designated as a paratype of the

species H. capensis, and most of the more significant postcranial bones, are

described. These indicate that the genus is morphologically intermediate

between monk seals, Monachus (Monachinae, Monachini), and Antarctic seals

(Monachinae, Lobodontini) excluding Mirounga. The structure of the auditory

region suggests a closer relationship with the Lobodontini, which are here

informally divided into two groups, namely, Leptonychotes/Ommatophoca and

Loboden/Hydrurga. There is evidence to suggest that the affinities of Homiphoca

lie with the latter group, and that it is likely to be more closely related to

Lobodon than any other living seal. Homiphoca may have been derived from the

North Atlantic Monotherium (Monachinae, Lobodontini), but the poor fossil

record of primitive monachines makes this uncertain.

The possible migration routes followed by early monachines are examined,

and it is suggested that the South Atlantic monachines probably followed the

route suggested earlier by Hendey (1972).

RESUME

La position des Phoques du Tertiaire supérieur de l’Atlantique Sud est ici

révisée. Prionodelphis rovereti du Miocéne supérieur/Pliocéne inférieur

d’Argentine fut décrit par Frenguelli (1922) sur la base de cing dents appartenant

a un Cétacé et un ou deux Monachinés (Phocidae, Monachinae). P. rovereti est

126 ANNALS OF THE SOUTH AFRICAN MUSEUM

considéré comme un Cétacé. Prionodelphis capensis, Hendey et Repenning 1972,

d’Afrique du Sud est assigné 4 un genre propre Homiphoca gen. nov.

Un crane et une mandibule presque complets, désignés comme paratype de

Vespéce H. capensis, et la plupart des éléments postcraniens les plus significatifs

sont aussi décrits. Ce matériel montre que H. capensis est morphologiquement

intermédiaire entre les Phoques moines Monachus (Monachinae, Monachini) et

les Phoques antarctiques (Monachinae, Lobodontini) excluant Mirounga.

La structure de la région auditive suggére un lien étroit avec les Lobodontini qui

sont ici divisés en deux groupes, Leptonychotes/Ommatophoca dune part et

Lobodon/Hydrurga d’autre part. Plusieurs arguments suggérent un rapproche-

ment d’Homiphoca avec le second groupe et plus précisément avec le genre

Lobodon. Homiphoca pourrait avoir son origine dans le genre Monotherium

(Monachinae, Lobodontini) de |’Atlantique Nord mais cette hypothése reste

incertaine compte tenu de la pauvreté du matériel des Monachinae fossiles.

Les routes de migration possibles, suivies par les premiers Monachinés, sont

envisagées et il est suggéré que les Monachinae de |’Atlantique Sud ont

probablement utilisé la route proposée par Hendey (1972).

ACKNOWLEDGEMENTS

We are indebted to Mr C. A. Repenning, U.S. Geological Survey, Menlo

Park, California, and Dr C. E. Ray, National Museum of Natural History,

Washington D.C., for casts and other comparative material used in the course

of the study. We are also indebted to Drs L. Ginsburg and R. Hoffstetter of the

Institut de Paléontologie, Paris, for helpful comments on the manuscript.

We thank Miss J. Nolte, South African Museum, for the photographs.

The Langebaanweg Research Project is supported by Chemfos Ltd, the

South African Council for Scientific and Industrial Research, the South African

Museum, and the Wenner-Gren Foundation for Anthropological Research,

New York (grant no. 2752-1834), and the assistance of these organizations is

gratefully acknowledged.

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