rol fi canis wt cubed dons ere saree ree: Lele Spaeyine 14 treba laf Tphb Os Hie 4a 7 coeds D-o Neate cd Lice piel wal ae tet lt Melee oe rs — ee
y yeoteds r ee s - eu 3 ride f eR os wi 4 rear? seh i ge EF htm LE Ap TO LE OL ON A A ll BE A he
ts hodint nT Is Pm ae gee ges $ aed mee ey ; trie Sea get pee Fae see eae Crs) Shaken ta Peer RPA p NM sgian Tete ab Dele tees ee oo yaar dwn

j re 7 RF 8 CEG IN TLS Hite) PA Ee Felt LEE BG PE my 2) Pry oly Pa ath DLs 230-448 4
Spiess 2
o set Shri BEERS GMP,
Tah ii hire Toe

A oF Be pag kde nal AS OER EN OA. Fe I tlhe a a
= PPE EAE Dal 2M Le ENO Ty cla es oh BE
So a PES PNAS DR Ae H. Bp Ber eA

nee MS eee Sef
fa find Baa fre
ger pidaoce :
4 : i : rat oe Soren r
ot ey Se Ppa e te : sf was -. Peeterd . . eet : ae Dae eee

a pr sen gad = r a 7 alan an : -' ry yy 425
4 2 Ah F - ed Pe Pd ° = Aakeahe te P ha > he a

A A ne nlite ~ %
Fee te i Pt lt te tte Mit Ne Ei yy Pe
a Pn OAL PLD AP Ena ogy tek ie ee
Cet Lait a etl Rat een MS ve? et ln eB?
‘ © <i” nM CR tee PL Di MR te OM ap S
ee eae a ee ee ee ea ek

PRR Pt ge Lhe ee ee eee ee

—

ee Pot

~_ - a - a owe ee ea en ee et ed
a a ~ “i ~ ater — : ~ tS t Boh - Pi Me eR wh ey amen tn ee NE ing Mele ae aos hy glen > lene tl
ne or-o —- S page « 2 —* o - Ra ee ie. Tomcat las oie MOP tin nella alt teen oT eMac a lp Na aa Mn "ay enh. ‘tn walitn- Ona

— anes ew — o POC Ee ae ~ ee * Ae tle = im ti) aelllay Sct itcn RAIN Menage mee atten he Ter Seeley Mn, at tae el ei te

obs Rial he Basin phe

eat te ae ae Pinan, Gaia 4 i 4 “ athe ° - poe a oe aad tals deseo 4 RE en IE UR ae tre ee EE
oF RBA mA REG Mi a oem nae rum taon #% sere 2 Bt An ret nn Oma PAM L tyne oe # . —— ~ atti ite tt ot. i ll het dein, ap ln a er et Te PTT Aas al, Peterlee ala i alli
wt era os seu+ 4 . — ~—_ e 7 ~~ a ry ems te a line a Scag tet NN Sty tots Bip i Spine esl eatin as
> - ~ . ~ ~ . - an ae tng Sc nL EAT i te Mlle Rea eT Ie
RAE) ferent ger writ ihn” AF Bato e ot. °. io tow 2 . a a ver —— CF ie te etl eae anh al ann Mage telat Attach a 4 ee *. ie S
LP ete Poentten nn Gite SOA a Re Mais ba Soe TE Re hed> -: tint “ red - ne > ~etn _ . et. -
6 aLelesP Dy A. eet 9% nal > “- vm ee
« ¥ - at en . - ade ed bed ee en
o — - > - betes aa bind itn
= oon - , . hat a eR me
7 . _— sa ne ne = a eee
my oa > 4 ~ Pinal aga e ae
- ~ * wpenrer eines evendinibettee tiee ee
‘ om = “ » er ees
a aed < -" Pn see? cpt «
- * - d vas . ~ a
¢ “ a . 4 ~ Aut ~ ae
“ - «
° - - ~ : .
. ai ae ne -
~ ~~
- / La —,
- - > » ~~ ae
= - >> att ~~ - ¥
- _ a 7 - al “ - a
. ss . Are ct Sie Sotetons ti
7 . « ~ ~*~
. = a Pe < Rh
‘ vd . Se Pp ig =
— anatiy
_ < - a. : Pe
—
tee
«
.
- es ~
iH =
‘
i
.
: P . o> a ee
ve i ail - eS ae —— ee ———— ee —— - - ” ee oe eee eee eee ee eS” ae tin ee ee ee Se ee oy OS eee

IWS S3IuYvuYdIT LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVYINOSHLINS S31%

z cf = au = me z
° = = .w = wo S
he 2 29 a a >) oe
> > = & > "at > 4 ~

oo: 2 Es : : % R

po : g mi & 3 i Xe
mm NOILALILSNI NVINOSHLINS S3ZIYVYaIT- LIBRARIES SMITHSONIAN INSTIT
Fis Ww Fat 2) 8 aris ~” i j z =e a
<x = —< = Fe = <x ;
— ee 4 -* — _—

S a S a a = 5

ie 3 + rae? r O- r

4 = = a = =

= se : = > | = = =

w” a. Pi Ww = ap) ; = ‘i
IWS Saliyvugi7 LIBRARI ES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLIWNS S31 4%
_ _— . te ; ‘ Ww ;

tu rr te £ re = = ul

nee + ae a Be by : | rae’ ~| ee,

: se. a: — oO pay

ee 5 <vf ‘Up S z= <

ane 5 i) > = nd
a S 1 “GY o se 5 via

x aug Ps ay a oul XN ay

VIAN INSTITUTION NOILONLILSNI NVINOSHLINS S3I1YVY¥EIT LIBRARIES SMITHSONIAN INSTI
aad Pu iz % z= c iN ra

— 2 : 2 =e SC ft «

Z - — E
Pe) A wD a

Yn = > = > “s >

a ee a E a 7 4 2B

” m WS n* m
A = wn ae - a = . a 4

LIWS SAIYWYRAIT LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3IUV
Ww : za ‘= Y” zz + w ae t w t
Eo ae, = = z = ae od
4 za =| —_ tJ =
: s Ne 3 : : 8%? 1 GY
7 ne) \_ WS ” ” w LO nD Hn GG
fe) fe WANQKY OC na O VW tl, 0 ¢
= SENN SEE Ee 2 2"
= ee ae er : a.

MIAN INSTITUTION NOILNIILSNI_NYINOSHLINS Sa 'YVYE!IT LIBRARIES SMITHSONIAN INSTIT
= > = w : = ‘

Ge : g 5 g - a ee

ha a < a ~ = = eer
= ss a : = m By
fe) a 2) pie: _ fe) ws ie 08
Zz ay = — 4 a i :

IWS S3IYVYEIT LIBRARIES SMITHSONIAN INSTITUTION NOILALILSNI NVINOSHLINS S314 %
= “3 = 2 5 D WA =

\ ra > oa > ~ $ N R =
be =: ' — > VK =
=. a aie A = s) . Es

te : D - D = -

NIAN INSTITUTION NOILALILSNI NVINOSHLINS S31YVYEIT LIBRARIES SMITHSONIAN INSTIT
z * w” z. wn = aa 21 = a
< = ae = es = es re
z = Zz = De aaa: Zz ar =
2 = S 7 LD ‘Mp os WN \ = 5
= O = Oo % he I \ O 4
ie 2 2 z iy E » 2 E
Y ~. z Ww) za Ww) *. we w” ea

LINS S3IYVYSII INSTITUTION NOILNLILSNI NVINOSHLINS S314
9) = w = Oo = Lie ie Hs

’ a. f Pyf Ay, 4 poe Y . = r

A ag = < Pup = a = ae

om = mo “i = om 5 ree

Seats) z a, © Zz a z= eat

NIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3!IYVYUEIT LIBRARIES SMITHSONIAN INSTIT
: : a ee : ; —

Bi = oo = oo — w

Vi ne a = 0 i >)

Jee > ms > = > = >

IBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S31uvudi7 LIBRARII

= ot = Le & Zz oc
Sa) = ow = wo Ns Oo w
=> Us a =) A — & Ls Pe)
ere = = Pe) = De)
22) m 7) an w” =
cS wo = wo = wn
NI NVINOSHLINS S3I1YVYEIT LIBRARIES
Zz. wn” = oe: ” z= 22)
< = <= = < =
Z a "ise = = =
ah 8) ae, O . — oO aS
: x 2 g 2 a
= = = zZ - S
= > = > = >
” z 2) gs mee Ww =
SMITHSONIAN INSTITUTION NOILNLILSNI LIBRARI
7) > 7) == wn =
Nw. te oes ox = ow =
7 AY a =
¥2) & 3 ¢ ce c e =
FS Y = m aa. a =
1S <a “le oO _ oO ais O
| -_ sl = ad an ar =
HINLILSNI NVINOSHLINS S3IYVYUSIT LIBRARIES SMITHSONIAN INSTITUTION NOILNLILS
ee z Soe :
ry = - 2 2 5
a i = Ss Ee
pas FE = - = -
ee wo ”
o = o z wo Zz
3RARIES INSTITUTION NVINOSHLINS S3SIYVYUEIT LIBRARI
_ o = <x = < = P Yy =
. =a a hbivi,fs
= 2 Ne 3 2 2 Gy, ?
\ 2 2 wR 2 a ay ge
> Re a S = “oN >’ = > =
| 7) Zz 7) ra a

DILALILSNI_ NVINOSHLIWS S3INVYGIT LIBRARIES SMITHSONIAN INSTITUTION NOILMLILS

Zz ” = wn = ”
Py a oe = cc .= a
9-) c < = <t i a
sip a o sr oh ar oc
Novant” Oo = cn.” 5 .
he RE - Zz me Za
IBRARIES SMITHSONIAN INSTITUTION NOILALILSNI_NVINOSHLINS Saiuvugiy tl Pham
_- z iit z
ot SON ee @ | Biz — aia)
) > = be “ > Ws 2° ;
2 : : 3 : :
s} FE: >
y= 2 2 2 = :
Neos” 2 | m o z a m
5 - ce _ 7) ee ”
OLLNLILSNI_ NVINOSHLIWS S3I1YVUGIT_LIBRARIES SMITHSONIAN INSTITUTION, NOILNLIL
< = a ee =e ang
S = ‘SE WL 7 ZN
a: AS a oO a. YY
> oY DER” 2 2 A O
& 2 = Ls 2 i Be | see
.. s : >
a S a = = =

SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLIWS Saluvugi7_-LIBRARI

Te) = 79) =
aie: 2 uw : us 5
x. 7 fp = o = 0: =
LZ Lp ef < = < c
« Qe = a = a =
a 1% 5 = ‘i Z 5
4 ie ; 2 Re 2 eee =
NOILALILSNI NVINOSHLINS S31YVYAIT LIBRARIES SMITHSONIAN INSTITUTION _
; + "i - Came,
; ° a S ty, a -
0) a a a We 5
a = D “=a = r

OF THE SOUTH AFRICAN
MUSEUM

CAPE ‘TOWN

ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

QY

|

Volume 63 _ Band

January 1974 Januarie

THE LATE CENOZOICG CARNIVORA
OF THE
SOUTH-WESTERN CAPE PROVINCE

By
Q. B. HENDEY

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
becomes available

Obtainable from the South African Museum ,P.O. Box 61, Cape Town

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na beskikbaarheid
van. stof

Verkrygbaar van die Suid-Afrikaanse Museum Posbus 61, Kaapstad

OUT OF PRINT/UIT DRUK
I, 3, 5, 73) B=, 5> t.—p.1.), 51S 2s 5> 9)
(ij EIc)5 TB B GL=2)y 102)

6(1,
WG 25 5, 7> = Dalla)|s 24(2), 27; 31(1-3), 33

T, 2(
I

Price of this part/Prys van hierdie deel
R20,00

Trustees of the South African Museum © _ Trustees van die Suid-Afrikaanse Museum
1974

ISBN 0 949940 34 8

Printed in South Africa by In Suid-Afrika gedruk deur
The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk.,
Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

THE LATE CENOZOIC CARNIVORA OF THE
SOUTH-WESTERN CAPE PROVINCE

By
Q.B. HENDEY
South African Museum, Cape Town

(With 78 figures and g1 tables)
(MS accepted 15 March 1973)

CONTENTS

General introduction
The south-western Cape Province . ;
Modern mammals of the south-western Cape Browne
Zoogeography of the south-western Cape Province .
Fossil mammal occurrences in the south-western Cape Province

Quaternary occurrences

Late Tertiary occurrences .
Fossil mammals of the south-western Gane Province
Non-mammalian fossils of the south-western Cape Province
Late Cenozoic mammal ages in southern Africa
Carnivora of the Pliocene

Introduction

Family Phocidae .

Family Canidae

Family Ursidae

Family Mustelidae

Family Viverridae

Family Hyaenidae

Family Felidae

Incertae sedis

Unclassified specimens .
Carnivora of the Pleistocene and Fidéeete

Introduction

Family Phocidae .

Family Otariidae .

Family Canidae

Family Mustelidae

Family Viverridae

Family Hyaenidae

Family Felidae

Unclassified specimens .
General discussion on the Carnivora :
The role of Carnivora in the origin of bone ect auteniens
Conclusion.
Summary .
Pete od ledsements
References .

Ann. S. Afr. Mus. 63, 1974: 1-369, 78 figs, 91 tables.

PAGE

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

GENERAL INTRODUCTION

The systematic investigation of the fossil Mammalia of South Africa was
begun towards the end of the 19th century with the description of a new species
of an extinct long-horned buffalo (Seeley 1891). This specimen, which had
been found in 1839 by Andrew Geddes Bain and which is now housed in the
South African Museum, came from the banks of the Modder River in the
Orange Free State. The interior plateau of South Africa has remained the
principal focus of local palaeomammalogical studies and numerous publications
have appeared which deal with material recovered from gravels along the
Vaal River, cave breccias in limestones and dolomites of the northern Cape
Province and Transvaal, and from sites such as Cornelia and Florisbad in the
Orange Free State. The Pleistocene mammal faunas of the interior of South
Africa are now moderately well known (see Cooke 1963).

Until about the middle of the present century little was known of the
fossil mammals of the coastal regions of the southern African subcontinent.
Scott (1907) described a small assemblage of Pleistocene mammals from the
Zululand coast, while series of fossils of late Tertiary age from the coasts of
Namaqualand and South West Africa were described by Stromer (1926,
1931a, etc.) and Hopwood (1929). Broom (1909), in one of his earlier palae-
ontological contributions, described a new equine species from a specimen
washed up on a beach near Cape Town, and this was the first description of a
fossil mammal from the south-western Cape Province, the area under considera-
tion in the present investigation. Following on this discovery was a period of
about 40 years during which very little collecting of fossils was undertaken
in this region.

In 1951 the prolific Pleistocene fossil occurrence on the farm Elands-
fontein near Hopefield, which is about 90 km north-north-west of Cape Town,
was visited by a scientist for the first time and its importance recognized (see
Singer 1957). The discovery of the remains of a fossil hominid (the ‘Saldanha
skull’) at Elandsfontein in 1952 led to considerable local and international
interest in this site and thereafter a number of publications appeared which
dealt with geological, palaeontological and archaeological aspects of the
occurrence. These investigations were climaxed by a series of systematic
excavations undertaken during 1965/6 (Singer & Wymer 1968).

Another major event in the south-western Cape was the discovery of
vertebrate fossils in the phosphatic deposits at Langebaanweg, 105 km north-
north-west of Cape Town (Singer & Hooijer 1958). The deposits at this locality
have yielded the largest and most important assemblage of Pliocene vertebrates
known from southern Africa.

Many other Quaternary vertebrate fossil occurrences are now recorded
in this region (Hendey 1969), and the south-western Cape now features more
prominently than before in the internationally-based investigations on the
fossil Mammalia of Africa. Particular interest centres on Langebaanweg,
which is the most prolific of the local sites and the best source of information

LATE CENOZOICG CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 3

on the nature of the Pliocene mammalian fauna of southern Africa.

The present widespread interest in the late Cenozoic of Africa is due largely
to the fact that it was on this continent and during this period in time that
important developments in the evolution of the Hominidae took place.
Although the skull and mandible fragments from Elandsfontein are still the
only important fossil hominid remains known from the south-western Cape,
the study of other fossil mammals from local Pliocene, Pleistocene and Holocene
occurrences are a significant part of the investigations into the life and events
of the late Cenozoic of Africa as a whole.

The present report summarizes the existing knowledge of the fossil
mammals of a geographical region which is limited in extent, and it deals in
detail with the known remains of just one mammalian order, namely, the
Carnivora. In general, carnivores tend to be less well represented in the fossil
record than other orders and this does apply in the case of those from the
south-western Cape. Nevertheless, the material available represents a significant
assemblage of largely unstudied specimens made up of a wide variety of species.

Investigations on African fossil mammals are still essentially a matter of
determining their systematics and it is the study of the relationships of the
local Carnivora which make up the greater part of the present report. An
appreciation of the taxonomy of the species represented does, however, allow
for interpretive work of various kinds.

One of the most critical problems evident in southern African Cenozoic
studies is the uncertainty which surrounds the relative and absolute age of
many of the local occurrences which are relevant to these studies. At present
only those which fall within the time limits of the radiocarbon dating technique
are potentially datable in absolute terms. This means that most of the impor-
tant fossil occurrences can only be dated in a relative sense and, although
relative dating may be undertaken in a number of ways (see Oakley 1964),
in dealing with fossil faunas it is usually most convenient to use the fauna
itself as the basis for dating. Kurtén (1957a) has demonstrated the usefulness
of the larger Carnivora in making temporal correlations between faunas in
widely separated areas. Consequently, one of the aims of the present study
was to determine the relative ages of the local assemblages on the basis of
their carnivore faunas and other evidence and to make a temporal categoriza-
tion of southern African fossil mammal faunas on a formal basis.

In addition to the determination of the age and phyletic relationships of
the species described, account was taken of their past and present distributions.
Viewed in relation to other mammals, some aspects of the evolution and dis-
persal of African Carnivora have become evident. Although the interpretation
of the available factual evidence is necessarily subjective, there is the basis of
an understanding of the evolutionary history of the carnivorous element in the
African mammalian fauna.

Since predation and scavenging by carnivores may lead to bones being
accumulated in certain areas and in certain ways, the possible relationships

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

between carnivore activity and local fossil occurrences was also investigated.

The African Cenozoic fossil record is poor and, because of the nature of
this record, account is taken only of the late Tertiary and Quaternary of the
south-western Cape. It was during this period in time that the character and
composition of the now declining modern fauna was developed and it is thus
only the climax of the ‘Age of Mammals’ which can be investigated.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 5

THE SOUTH-WESTERN CAPE PROVINCE

The definition of geographical regions is frequently difficult and the final
boundaries selected are often arbitrary. This is true of the south-western Cape
Province and, although it is one of the most easily defined regions in southern
Africa, its boundaries can. be varied according to the nature of the study being
undertaken.

In the present instance this region is taken to include that area between
the coast from St. Helena Bay to Cape Hangklip and the Cape Folded
Mountains, with the lower course of the Great Berg River forming its northern
boundary. It may be subdivided as follows:

(1) Cape Peninsula—'That area between Table Bay in the north and Cape Point in the south,
and including Table Mountain and the mountains extending south from it.

(2) Coastal plain—The remainder of the region and including the areas known as the Cape
Flats, the Sandveld and the Swartland (see Talbot 1947).

The maximum north-south extent of the region is about 190 km and the maxi-

mum east-west extent about 90 km (Fig. 1).

The regions adjacent to the south-western Cape (Fig. 2) and which will

_ be referred to from time to time are as follows:

(1) West coast—The coastal plain north of the Great Berg River.

(2) Southern Cape—The southern coastal plain of the Cape Province stretching from Cape
Hangklip in the west and Cape St. Francis in the east.

(3) Cape Folded Mountains—The mountains which separate the west coast, south-western
Cape and southern Cape from the inland plateau (Karroo).

King (1951) has discussed the geomorphology of the south-western Cape
and adjacent areas under the headings ‘Cape Folded Belt’ and ‘Southern
Coast’. The former is a complex system of mountain ranges which may be
conveniently divided into two groups, namely, a group which trends north—
south roughly paralleling the Atlantic coastline and those which trend east-
west approximately parallel to the Indian Ocean coastline. The former group
intersects the coastline in the south at Cape Hangklip, effectively separating
the south-western Cape from the southern Cape. Since this group is parallel
to the Atlantic Ocean, the west coast region is in the nature of an open corridor
stretching northwards from the south-western Cape. The east-west group
approaches the southern Cape coast at intervals so that the coastal plain is
significantly narrowed for a distance of about 300 km west from Cape St.
Francis.

In brief, the most southerly part of the African continent is made up of a
complex system of mountain ranges and valleys fringed by a coastal plain,
which in the south-western Cape has an average width of about 65 km. The
physiography of the southern continental extremity is of considerable zoo-
geographic significance and will later be discussed again in this connection.

The geology of the south-western Cape is relatively simple. There are
exposures of Archaeozoic and Proterozoic igneous and metamorphic rocks,
mainly in the coastal plain, while the Cape Folded Mountains and Cape

ANNALS OF THE SOUTH AFRICAN MUSEUM

St. Helena
Bay

€

>

SALDANHA x WN
LANGEBAANWEG Great, Be Ker

e
RODIN) Q
go op ~
hy
=
X

ELANDSFONTEIN

S
=
x
7)

MELKBOS

Atlantic
Ocean

Table
Bay

Cape
Town

CAPE FLATS

SWARTKLIP

False Bay

Cape
Hangklip

Fig. 1. The south-western Cape Province.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 7

Ale
31
EASTERN
CAPE
Sourn- N\N , @6.°~=="">2=>>-=:-
WESTERN, | ‘SED Mibrs wee
35 | |
17 21 25 29
0 500
km

Fig. 2. The Cape Province, showing the regions referred to in the text.

Peninsula are made up principally of sedimentary and metamorphic rocks of
the Cape System (Archaeozoic).’The Cape Peninsula is connected to the
mainland by an isthmus of largely unconsolidated late Cenozoic sediments,
which extend along the coast to the northern limits of the region and beyond.
Most of the Tertiary element of these deposits was apparently reworked during
the Quaternary. Overlying some of the deposits of the coastal plain are
unconsolidated aeolian sands which are probably largely Holocene in age.

The late Cenozoic deposits are significant because they contain the fossil
occurrences dealt with in the present report. All the major, and most of the
minor vertebrate fossil occurrences in the south-western Cape are located in
either the Cape Flats or the Sandveld. In most instances the exposure of the
fossiliferous deposits has resulted from the stripping of cover sands by erosion
or human agency.

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

MopERN MAMMALS OF THE SOUTH-WESTERN CAPE PROVINCE

‘In this Land of prester Fohn ther was seene by our Men Lyons and monkeyes,
Babownns a multid, with divers other Strange beastes as Antilops and many
other deformed creattures verie strange to be Sene.’
Standish-Croft Journal, Cape of Good Hope, 23 June 1612.
(Raven-Hart 1967: 59).

Like most of Africa, the south-western Cape Province is part of the
Ethiopian faunal region and its mammalian fauna is unmistakably ‘African’
in character. It is an important part of this region since it was in this area that
Europeans first encountered many of the mammals which characterize the
Ethiopian fauna (Table 1).

By the time that the first permanent European settlement was established
on the shores of ‘Table Bay in 1652, ships of the great trading nations of Europe
had already been calling at the Cape for more than 150 years and there are
many log and journal references to the rich fauna of the region (see Raven-
Hart 1967). Records such as that quoted above are, for the most part, brief
and imprecise, although they indicate very clearly that the Cape had as rich
a mammalian fauna as any other part of sub-Saharan Africa explored sub-
sequently.

The indigenous inhabitants of the region were the beach-combing and
pastoral Hottentots, whose presence apparently had little or no effect on the
numbers and variety of mammals occupying the territory. The arrival of the
European settlers, however, initiated the decline of both the indigenous people
and animals. Hunting, wanton slaughter and destruction of natural habitats
steadily escalated as the settlements grew and spread further afield and by the
time that the first modern scientific records of fauna were being made in the
18th century, the fauna of the south-western Cape was already much depleted.
Consequently, it is now impossible to make an accurate reconstruction of the
nature and composition of the local mammalian fauna as it was early in the
historic period.

The lists of local historic period mammals (Table 6, 89) were compiled
on the basis of historical records, place names and such scientific accounts as
are available and, although they are probably fairly comprehensive, they are
of necessity provisional and subject to revision. The historical records are often
vague, confusing and even fanciful, while the early scientific accounts also
have their shortcomings. For example, the holotypes of some of the species
listed in Table 1 were probably not from the south-western Cape region as it
is here defined since locality records are vague. Most are simply recorded as
being from the ‘Cape of Good Hope’ (see Ellerman e¢ al. 1953), a term which
is not necessarily synonymous with the south-western Cape. It is, however,
probable that the holotypes of these species were representatives of populations

which did, or still do, include the south-western Cape in their area of distri-
bution.

LATE CENOZOIC CGARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 9

TABLE I

Some Ethiopian land mammals, the

holotypes of which are from the south-

western Cape Province or immediatelv
adjacent regions.

*Chrysochloris asiatica Linnaeus, 1758

*Papio ursinus Kerr, 1792

* Otocyon megalotis Desmarest, 1822
Canis mesomelas Schreber, 1775

*Ictonyx striatus Perry, 1810

* Mellivora capensis Schreber, 1776

* Aonyx capensis Schinz, 1821

*Genetta tigrina Schreber, 1776

* Herpestes pulverulentus Wagner, 1839
Atilax paludinosus G. Cuvier, 1829
Hyaena brunnea Thunberg, 1820
Felis serval Schreber, 1776

*Felis caracal Schreber, 1776
Orycteropus afer Pallas, 1766

*Procavia capensis Pallas, 1766
Diceros bicornis Linnaeus, 1758
Equus zebra Linnaeus, 1758
Phacochoerus aethiopicus Pallas, 1766

*Sylvicapra grimmia Linnaeus, 1758

* Raphicerus campestris Thunberg, 1811

* Raphicerus melanotis Thunberg, 1811
Oreotragus oreotragus Zimmermann, 1783

*Pelea capreolus Forster, 1780
Taurotragus oryx Pallas, 1766

*Lepus capensis Linnaeus, 1758

* Bathyergus suillus Schreber, 1782

*Georychus capensis Pallas, 1779

*Cryptomys hottentotus Lesson, 1826
Pedetes capensis Forster, 1778

*Praomys verreauxi A. Smith, 1834

* Mus minutoides A. Smith, 1834

* Acomys subspinosus Waterhouse, 1838

* Otomys irroratus Brants, 1827

* Tatera afra Gray, 1830

* Species of which indigenous populations still occur in the region.

During the 18th and 1gth centuries, the European settlements spread
beyond the confines of the south-western Cape and the destruction of flora and
fauna became more widespread. By this time, however, scientific records were
being made more frequently and specimens were being preserved in scientific
institutions in Europe. Consequently, while the extinction of the blaauwbok
(Hippotragus leucophaeus) in about 1790 passed unnoticed in South Africa,
descriptions of this animal were already in existence and specimens were
preserved in European museums. In addition, the 19th century saw the advent
of the practice of wildlife conservation so that even in settled areas the preser-
vation of species was assured. Unfortunately these developments came too
late for the south-western Cape and by the end of the 1gth century most of the
larger mammals of this region were already extinct.

Io ANNALS OF THE SOUTH AFRICAN MUSEUM

Such conservation as was undertaken in the Cape Province during the
19th century, and indeed for much of the present century as well, largely
excluded the south-western Cape, at least in as far as the mammals were
concerned. One hundred years ago there remained only the vestiges of the
mammalian fauna which had flourished a scant 200 years earlier, and even
this has continued to be depleted and disturbed.

Although the most obvious effect of human activities has been the
reduction in the size of populations of locally occurring mammals, this is not
a general rule. For example, in recent years there have been periodic plagues
of gerbils (Tatera afra) in the grain-growing areas of the region, and it is the
reduction or elimination of the natural enemies of this species which has
largely contributed to this phenomenon.

Several of the mammals which had become extinct in the region have now
been re-introduced, mainly into nature reserves, while other species not known
to have occurred locally in historic times have also been introduced. The
former category includes the eland (Taurotragus oryx), and the latter includes
Burchell’s zebra (Equus burchelli), the bontebok (Damaliscus dorcas dorcas) and
springbok (Antidorcas marsupialis). Even exotic species have now become
established locally. Apart from those species such as the brown rat (Rattus
norvegicus) which have an almost world-wide distribution, there are the grey
squirrel (Sc7urus carolinensis), the fallow deer (Dama dama), the sambar deer
(Cervus unicolor) and the Himalayan thar (Hemitragus jemlahicus).

In few other regions in sub-Saharan Africa has the natural pattern of the
mammalian fauna been disturbed to the extent which is evident in the south-
western Cape.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE II
ZOOGEOGRAPHY OF THE SOUTH-WESTERN CAPE PROVINCE

INTRODUCTION

In the course of investigations into the fossil mammals of the south-western
Cape it became evident that, in order to place these studies in perspective,
account had to be taken of past and present patterns of distribution of African
mammals. Pinniped distributions as they relate to the seal from Langebaanweg
have already been discussed elsewhere (Hendey 19720), and so the observations
which follow concern only terrestrial species.

A study of late Cenozoic palaeozoogeography is necessarily based on
an understanding of the modern biology and geography of the region in
question, although even then a clear and unequivocal interpretation of past
events is not always possible. This is at least partly due to the fact that there
were marked changes in climate during this period in time and the nature and
effects of such changes may be difficult to determine. The late Cenozoic
climatic changes of southern Africa are not well documented or clearly under-
stood and consequently many of the statements which follow are tentative
and based on inadequate factual information. They could, however, provide
a basis for future studies. As more becomes known of the fossil mammals of
Africa and the nature of past climatic changes become better understood, the
present conclusions may be tested and, when necessary, modified.

Darlington (1957: 420) summed up the basis of the present zoogeographic
study by stating that it ‘is only in the present that we can see exactly how
animals really are distributed and how their distributions are related to space,
climate, barriers, other organic factors, plant cover and each other’. The
human activities and influences referred to earlier have, however, so altered
the character of the modern flora and fauna of the south-western Cape that
much of the information fundamental to the present investigation can now
be obtained only by indirect methods. This is, of course, by no means a unique
situation and many parts of the world are in a comparable or even worse state
in this respect. Nevertheless, since sub-Saharan Africa includes large tracts of
unspoilt countryside, it is as well to emphasize that this does not apply in the
case of the south-western Cape.

This region differs from the rest of sub-Saharan Africa in other respects
as well. It is the only part of the sub-continent to lie within a warm-temperate
climatic zone and have a Mediterranean type of climate (Cs of Koeppen).
Its climate is of considerable biological significance.

The rainfall, which is mostly cyclonic, falls mainly in the winter and the
summers are long and dry. The precipitation varies considerably from place
to place, the Cape Flats receiving only 400-500 mm per annum, while the
annual total in the mountains only a few kilometres away may exceed 3 000
mm (Schulze 1965). The depressions which cause most of the precipitation
pass from west to east in southern mid-latitudes and it is usually only in winter
that they reach far enough north to affect the south-western Cape. There is a

12 ANNALS OF THE SOUTH AFRICAN MUSEUM

marked decrease in the rainfall northwards along the Cape west coast and north
of the Orange River true desertic conditions prevail (the Namib Desert).
The inland plateau is a summer-rainfall area and that part closest to the south-
western Cape (the Karroo) is semi-arid. The southern Cape receives rain at all
seasons, although in the western parts the totals are not high. Some of the
higher mountains in these areas are snow-covered for short periods during
winter.

The cold Benguela Current along the Cape west coast has a moderating
effect on temperatures in the area. Cape Town has a mean annual temperature
of 17°C and a relatively small mean annual range of 12,8° to 21,7°C. Frost is
rare in the lower-lying areas and on the mountain slopes below 600 m.

The climate has been an important factor in the development of a unique
vegetation in the region. As in other winter rainfall areas, evergreen bushes
and shrubs with leathery or waxy leaves are predominant. The flora is very
diverse and endemic species are abundant. This vegetation, which is usually
referred to as ‘Cape Macchia’, may be subdivided into a number of groups,
the distribution of which is largely dependent on the amount of rainfall.
Forests are confined to well-watered and sheltered parts of mountain slopes,
although most indigenous montaine forests have now been destroyed as a
result of human activities. Grasses constitute a relatively insignificant element
in the vegetation in all areas, but this applies particularly in the Sandveld and
Cape Flats. The unique character of the local natural vegetation has attracted
the interest of botanists for generations and there is an extensive literature on
the subject. This vegetation is believed to have originated within the south-
western Cape winter rainfall area (Levyns 1962).

The definition of the south-western Cape as a distinct geographical region
is based principally on its climate and vegetation, and these in turn are deter-
mined by the latitude and physiography of the area.

PALAEOGEOGRAPHY

Brain & Meester (1964: 332) have pointed out that a ‘fuller understanding
of past climates will elucidate many of the problems concerning the distribution
and systematic position of South African endemic (mammalian) species’.
Unfortunately little generally acceptable work has been done on the late
Cenozoic climatic changes of this country, although such changes undoubtedly
occurred and, just as certainly, did influence the evolution and dispersal of local
mammals.

There is evidence which shows that these climatic changes had a marked
affect on the vegetation of the south-western Cape and adjacent areas. For
example, the trunks of large trees have been found in excavations on the Cape
Flats, an area which prior to the introduction of various exotic trees was
covered by Cape Macchia and unvegetated sand dunes.

Much has been written about the Pleistocene climatic changes in South
Africa and the principle of ‘glacio/pluvial’ correlations was once widely

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 13

accepted. While it is now certain that such correlations are not generally
applicable, it does seem likely that they have a certain validity in the case of
the south-western Cape. At those times during the Pleistocene when there was
a general lowering of world temperatures, there was apparently a shift towards
the Equator of the world’s climatic zones. In the south-western Cape the most
obvious effect of such a shift would have been to bring the rain-bearing
depressions into the area throughout the year, rather than just in winter as is
presently the case. Van Zinderen Bakker (1967) has already suggested as
much in a discussion on South African Pleistocene climates. If this was indeed
the case, then the south-western Cape would have experienced a cool-temperate
rather than warm-temperate climate and received rainfall throughout the
year.

_ Butzer (1961) has stated as a general principle that the areas bordering
the Mediterranean Sea experienced pluvial conditions synchronously with the
higher latitude Pleistocene glaciations. He referred to these as ‘Mediterranean
pluvials’ and concluded that they were ‘Early Glacial’ (Butzer 1961: 455).
It is assumed that this principle applies for Mediterranean regions in the
Southern Hemisphere as well.

The degree to which local climate changed during the colder phases of
the Pleistocene has yet to be determined. Linton (1969) has recorded what he
believed to be evidence of ‘cryonival phenomena’ in the south-western Cape.
If his interpretation of the geological record is correct, it means that the
lowering of temperatures in this region at certain times during the Pleistocene
was sufficient to produce perennial snow cover on the mountains. Even today
the higher peaks of the Cape Folded Mountains are snow-covered for short
periods during winter and under colder conditions a snow cover for the duration
of winter at least, is not difficult to visualize. Even without the perennial snows
suggested by Linton, the presence of snow gathered for prolonged periods during
winter would have had a marked effect on the local environment. For example,
it would have served to keep local rivers and streams at higher levels during
summer, in contrast to the position today when many local water-courses have
a poor flow, or are completely dry during the largely rainless summer.

Thus, in theory at least, during those periods in the Pleistocene when
the higher latitudes were glaciated, the south-western Cape is likely to have
been far more verdant than at present. Conversely, the existing relatively dry
phase would have been repeated during the Pleistocene ‘interglacial’ periods.

During the colder phases there was probably a northward expansion of
the Cape flora and extensions to the areas covered by forest. Relict patches of
such vegetation which are today found beyond the limits of their expected
distribution (see Acocks 1953: 12) may thus be accounted for, although it is
now difficult to distinguish between the effects of natural change and that
induced by human influences.

It would be unrealistic to suppose that uniform changes in rainfall were
experienced over the whole of the south-western Cape at any given time. Many

[4 ANNALS OF THE SOUTH AFRICAN MUSEUM

local geographical factors would have to be taken into consideration and it is
almost certainly impossible to reconstruct a detailed picture of = and
vegetation at various times during the late Cenozoic.

This region was probably never much more arid during the late Cenozoic
than it is today, since many indigenous plant species would probably have
become extinct if this was the case. Under more arid conditions moisture-
demanding species may still have survived in isolated places where conditions
remained favourable but, whereas major northward shifts of the flora were
possible during wetter periods, the converse is not possible because of the
situation of the south-western Cape at the southern continental extremity.
This restricting influence, coupled with repeated climatic changes may have
contributed to the extreme diversification of the local flora in which ‘the
majority of the species will vary from mountain to mountain’ (Acocks 1953:
153).

The late Cenozoic climatic changes in the south-western Cape must have
had a profound effect on the fauna of the region, perhaps most significantly
in the limitation or otherwise of contacts between locally occurring populations
and those further north (vide infra).

FAUNAL DISTRIBUTION IN THE SOUTH-WESTERN CAPE

In a discussion on the past and present dispersal of animals, Darlington
(1957: 572) concluded that the ‘South African fauna as a whole has evidently
formed by southward movement (extension) of parts of the tropical African
fauna. Replacements must have occurred both in the tropics and southward,
but the absence of outright barriers has prevented the persistence of many
relicts in South Africa. .. .’

Later he went on to qualify these remarks and in referring to the most
southerly parts of the country he stated that ‘climate has importani effects on
distribution of plants and animals in South Africa. Although the southern
tip of Africa is not cold, it is climatically differentiated and also is isolated from
the main part of the continent by a barrier of aridity that eventually retards
southward dispersal of water-demanding organisms’ (Darlington 1965: 110).

These quotations are relevant to many of the observations which follow
and conveniently set the stage for a consideration of the south-western Cape
fauna.

In an account of the biotic regions of southern Africa as indicated by the
Amphibia, Poynton (1964: 206) commences his discussion on the pattern of
amphibian distribution as follows:

‘A cardinal feature of amphibian distribution in southern Africa is the
north-east to south-west polarization of the fauna, there being one focal
point in the Mozambique plain, and another in the south-western Cape.’ The last
part of his statement is emphasized here for obvious reasons.

In an earlier paper (Poynton 1960) he concluded that the south-western
Cape amphibians are geographical equivalents of a north-temperate fauna and

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE T5

he regarded them as true south-temperate forms. In both papers he stresses the
unusual character of the amphibian fauna of this region, which has ‘a taxonomic
bias quite different from that of the tropical fauna’ (Poynton, 1964: 208).

While zoogeographic patterning is so marked in the Amphibia, Poynton
(1964) found that this was little influenced by ecological factors and that
similar habitats in the south-western Cape and south-east tropical Africa are
occupied by completely different types of Amphibia. All these observations
led to the conclusion that it is the aridity of the southern part of the central
plateau which was the principal limiting factor in local amphibian distributions.

Although the south-western Cape is a significant and, in some ways,
unique area in as far as the distribution and representation of one group of
lower vertebrates is concerned, it does not necessarily follow that the mammals .
would exhibit the same degree of regionality of character. Nevertheless, the
mammalian fauna does differ in certain respects from those of regions further
north.

For example, a comparison between the modern mammals of the south-
western Cape and those of south-eastern Kenya shows a considerable variation
in the commonality of species in different orders (Table 2). Regional differences
are most evident amongst small mammals (e.g. insectivores) and those which
are highly selective as to habitat (e.g. primates). They are least evident in that
group which is least dependent on environmental factors (i.e. the carnivores).
Such differences as do exist amongst the carnivores are largely confined to the
smaller species (viverrids), which might therefore be grouped with other small
mammals for the purposes of accounting for observed regional differences.
The larger herbivorous species (e.g. bovids) occupy a more or less intermediate
position in respect of the species commonality of the two regions.

TABLE 2

Mammalian species occurring in south-eastern Kenya! and the south-western
Cape Province.

Insectivores Equids
and and Carnivores
primates bovids
South-western Cape species . 10 21
South-eastern Kenyan species 25 27
Species common to both areas . 6 17
Index of resemblance (%)? . 60 81

1 Based on records for the Tsavo National Park (Williams 1967) and supplemented from
records given by Dorst & Dandelot (1970).
2 Simpson 1967.

In dealing with the distribution of present-day species it is clear that the
narrow definition of the south-western Cape given earlier is not the most
convenient, but that in this instance the Cape Folded Mountains and southern

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

Cape regions should also be included. Meester’s (1965) ‘South-West Cape
Biotic Zone’ covers all these areas and it is here termed simply the ‘Cape
Biotic Zone’, of which the south-western Cape is a subdivision.

The following small mammals are listed by Meester as being endemic to
the Cape Biotic Zone:

Bathyergus suillus, Praomys verreauxi, Acomys subspinosus and Tatera afra.

Two endemic forms are included amongst the larger mammals, namely, the
recently extinct blaauwbok (Aippotragus leucophaeus) and the bontebok (Damalis-
cus dorcas dorcas).

The blaauwbok was known in historic times only from the southern Cape,
but it formerly occurred in the south-western Cape as well. Its status as a
species distinct from the roan and sable antelopes (H. equinus and H. niger)
has never been seriously questioned and it was either an autochthonous species,
or, less probably in the light of available evidence, a more widespread species
which survived later in the Cape Biotic Zone than elsewhere.

The bontebok was formerly given full species status (D. pygargus), but is
now regarded only as a subspecies of D. dorcas. It is recorded in a natural state
only in the western parts of the southern Cape, several hundred kilometres
from the nearest record of its close relative, the blesbok (D. d. phillipsi) in the
eastern Cape. This species presumably had a continuous distribution in the
fairly recent past (? late Pleistocene) and the split between the two populations
was probably the result of environmental changes which will be discussed later.
D. dorcas is endemic to South Africa.

Another bovid which is largely confined to the Cape Biotic Zone is the
grysbok (Raphicerus melanotis), although its range also extends into the eastern
Cape where it is less common. By contrast, the mountain zebra (Equus zebra)
occurs in the Cape Folded Mountains and also in parts of South West Africa.
These two species are mentioned here as examples of mammals which, on the
one hand, range from the Cape Biotic Zone into the comparatively well-
watered south-east of the subcontinent and, on the other, range from the Cape
Biotic Zone into the arid south-west.

Another significant feature of the distribution patterns of larger southern
African mammals is that there are many essentially tropical species whose
ranges extend into South Africa but stop short of the Cape Biotic Zone. These
include the giraffe (Giraffa camelopardalis), the impala (Aepyceros melampus) and
various species of waterbuck (Kobus spp.).

It is thus evident that the mammalian fauna of the most southerly parts of
Africa is distinguishable from that of the subtropical and tropical parts of the
continent, and that there is a distinct and significant patterning in the dis-
tribution of species. Some forms occur in the Cape Biotic Zone and elsewhere
in the Ethiopian Region, some are endemic to this zone, some range from this
zone into the arid south-west of the continent, others range into the south-east,
and finally there is that group whose ranges do not extend into the Cape Biotic
Zone. This zone might, therefore, be regarded as a focus and area of overlap

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE EJ

of two distinct elements of the Ethiopian mammalian fauna, namely, that of
the arid south-west and that of the well-watered south-east. These groups are
superimposed on a third element, the endemic species of the region. On purely
theoretical grounds it is possible that in the past the relative importance of each
of these three elements may have changed in accordance with prevailing climatic
and other factors.

Patterns of animal distribution are determined by geographical factors
and are largely dependent upon the ease with which migrations can take place.
In this connection Simpson (1967) has defined three categories under the
heading of ‘paths of faunal interchange’, and although his work was based on
inter-continental migrations of animals, the definitions can be applied equally
well in a more restricted sense. They are as follows:

(1) Corridor route—‘along which spread of many or most of the animals of one region to
another is probable’ (p. 87).

(2) Filter route—‘across which spread of some animals is fairly probable but spread of others
definitely improbable’ (p. 87).

(3) Sweepstakes route—‘across which spread is highly improbable for most or all animals but
does occur for some’ (p. 88).

Migrating mammals can approach the south-western Cape by any of

three routes:

(1) From the north along the western coastal plain.

(2) From the interior plateau across the Cape Folded Mountains.

(3) From the east across the most south-westerly parts of the Cape Folded Mountains (the
Hottentots Holland Mountains).

Each of these routes has its limitations and each could conceivably
have had its character altered by past climatic and physiographic changes.

The northern route is the only one at present which might be regarded
as a true ‘corridor route’ since there are no barriers of any significance along
the Cape west coast and only the most southerly of the rivers traversing this
route are perennial. Beyond the northerly limit of the Cape Folded Mountains
the escarpment into the interior is low and discontinuous, so this route also
offers easy access to the interior plateau. However, both forks of this route lead
into arid (the Namib Desert) and semi-arid (the Karroo) regions and under
existing climatic conditions it is likely to be used only by those animals which
are adapted to dry conditions.

On the previously stated assumption that the most southerly parts of
South Africa were never much more arid at any time during the late Cenozoic,
it is concluded that the arid south-west element in the fauna of the extreme
south was probably never much more prominent than it is today.

During those periods when the south-western Cape and immediately
adjacent areas were wetter than at present, the northern route may have been
used by animals originating in the interior plateau which had previously not
been able to penetrate the semi-arid Karroo.

The alternative for such animals would have been to use the second of the
access routes, namely, that across the Cape Folded Mountains, although even

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

under the most favourable environmental conditions this was probably never
more than a ‘filter route’ because of the obstacles presented by the mountains.
There is a fairly easy route through the mountains from the Karroo into the
southern Cape by way of the Hex and Breede River valleys, but passage into
the south-western Cape would always necessitate the crossing of mountains.

The last of the access routes, that from the southern Cape, has a number of
disadvantages and it is the one which would have had its character most altered
by past climatic and physiographic changes. At present the Hottentots Holland
Mountains form a barrier between the southern and south-western Cape which
is probably sufficient to inhibit the free movement of some mammals. Conse-
quently, it might now be regarded as a ‘filter route’. The absence from the
modern fauna of the south-western Cape of species such as the bontebok,
blaauwbok, reedbuck (Redunca arundinum) and bushbuck (Tragelaphus scriptus)
may in part be due to the presence of this mountain barrier.

At those times during the Pleistocene when the sea level was lower than at
present, access to the south-western Cape around the southern tip of what is
today Cape Hangklip must have been much easier. Such conditions probably
did prevail during the late Pleistocene (the last glacial period) and both the
blaauwbok and reedbuck were present in the south-western Cape at this time.
Their extinction locally during the Holocene may have been due to a variety of
factors, including the possibility that they failed to adapt to the changing
environment. In this connection it is worth recording that bontebok introduced
into the Cape Peninsula from the southern Cape have not adapted well to their
new surroundings and remain in poor physical condition if left to the natural
resources of the area.

The southern route also has disadvantages at its eastern end. Much of the
southern Cape between Mossel Bay and Humansdorp, a distance of about
300 km, is an area of natural forest under existing climatic conditions, although
little forest remains as a result of recent human activities in the area. This
forested area, coupled with the relatively narrow coastal plain, must serve
as a barrier to the free movement of many animals along this route.

This part of the southern route would, however, have changed character
when sea level was lower than at present. The proof that this did indeed happen
during the late Pleistocene and the effect it had on the local fauna was recently
recorded by Klein (1972). He has suggested that the exposed continental shelf
in the Plettenberg Bay area may have been a grassy plain about 75 km wide
during the last glaciation, and that it was populated by an appropriate plains
fauna drawn from the interior plateau. It may have been at this time that the
distribution of Damaliscus dorcas was continuous between the areas occupied
by the surviving populations of the bontebok and blesbok. With the rise in
sea level again during the Holocene part of the late Pleistocene population of
this species became cut off in the western part of the southern Cape. Here it
survives as a relict population subspecifically distinct (D. d. dorcas) from the
main body of the species (D. d. phillipsi) in the eastern Cape and interior.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 19

If the wide, grassy coastal plain envisaged by Klein extended as far as the
south-western Cape, and there is no reason to suppose that it did not, then it
would have provided an ideal ‘corridor route’ between this region and the
eastern Cape. Consequently, not only southern Cape mammals would have
gained access to the south-western Cape as suggested earlier, but also species
from still further afield.

While the situation in the late Pleistocene can be gauged with some con-
fidence, it becomes progressively more difficult further back in time to determine
the nature of both climatic and physiographic changes. Nevertheless, it is
fairly certain that similar changes did take place earlier and that the character
of fossil faunas was dependent upon the accessibility of the south-western Cape
to the rest of the subcontinent.

One final point concerning the southern access route relates to the nature
of the easterly outlet to the southern Cape. At present the semi-arid Karroo
stretches almost as far as the east coast in the vicinity of Port Elizabeth, near to
the easterly terminus of the Cape Folded Mountains. At any time when the
rainfall in that area was lower than at present, the semi-arid barrier may have
extended through to the Indian Ocean (see Cooke 1964: fig. 10). The Cape
Biotic Zone would then have been completely sealed off from the north by
desert and semi-desert stretching from the Atlantic to the Indian Oceans.

It may be concluded that while the south-western Cape was never com-
pletely isolated from the rest of the subcontinent in a zoogeographic sense, it is
so situated that under certain conditions faunal interchange between it and
adjacent regions may have been very limited. The south-western Cape has
probably been occupied by at least some species endemic to the most southerly
parts of Africa throughout that period covered by the fossil mammal record
(late Pliocene to Holocene), while the arid south-west element of the Ethiopian
fauna was never much more prominent than it is today, although the south-
east African element probably has been more prominent on occasions.

FAUNAL DISTRIBUTION IN SOUTH AFRICA

Although a consideration of the zoogeography of South Africa as a whole
is beyond the scope of the present report, some comments on this subject are
appropriate.

During the Cenozoic the fauna of southern Africa can have changed its
character in only two ways, namely, by evolution in situ and by the immigration
of animals from the north. There is no evidence to suggest that this area has
been a major centre of mammalian evolution in the late Cenozoic and not even
the endemic mammalian genera are necessarily autochthonous. Almost all
the recorded genera of the local late Cenozoic are also known from elsewhere
in Africa, or even further afield and there are probably very few which had
their origins in southern Africa. There are, however, possible exceptions. For
example, the curious antelope, Pelea, is today confined to southern Africa
and is also unknown as a fossil elsewhere. There are probably a number of

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

exceptions amongst the smaller mammals as well (e.g. Chrysochloris, Bathyergus).

The contribution that the subcontinent has made to the Ethiopian mam-
malian fauna is most likely to have originated largely in the arid south-west,
an area which includes the Kalahari and Namib Deserts. This is Africa’s most
extensive arid region south of the Sahara. The previous existence of an arid
corridor between south-west and north-east Africa has been the subject of a
number of investigations (see Van Zinderen Bakker 1969: 139), and the desertic
faunas of these areas do have some mammalian species in common (e.g. Oryx
gazella, Madoqua kirki). Such species could perhaps be southern African in
origin.

It is likely that, for the most part, the mammals of the subcontinent had
their origins elsewhere and that these were added to the local fauna by immi-
gration from the north and that local differentiation of these forms was confined
to lower taxonomic categories. Apart from the endemic bovid genera, Pelea
and Antidorcas, larger mammals which are endemic to the sub-continent include
Hyaena brunnea, Connochaetes gnou and Damaliscus dorcas, while recently extinct
forms include the quagga (Equus quagga) and blaauwbok. |

The last two are part of a category of extinct ‘Cape’ mammals, so named
because they were largely confined to what is today the Cape Province.
Certainly none is recorded from as far north as the Tropic of Capricorn. The
Cape lion (Panthera leo melanochaita), the Cape warthog (Phacochoerus aethiopicus)
(see Ewer 1957a) and the Cape hartebeest (Alcelaphus buselaphus caama) are
other members of this category.

The suggestion has been made that the southern African fauna includes,
or has included late survivors of species which are, or were already extinct
elsewhere (e.g. Hendey 1969). There can be little doubt that this supposition
has some validity, but locally endemic species are not necessarily primitive
forms. For example, Hyaena brunnea is more specialized in certain respects than
its East African counterpart, Hyaena hyaena.

On the other hand, the occurrence of a boselaphine and agriotheriine
in the late Pliocene fauna from Langebaanweg when these two groups were
apparently unrepresented elsewhere in sub-Saharan Africa, suggests that there
was a tendency for certain species to persist for longer in the extreme south.
In East Africa during the late Pliocene and early Pleistocene, the Bovidae
were already represented exclusively by the genera which characterize the
modern bovid fauna of the Ethiopian region, but in South Africa both bosela-
phines and ovibovines (Makapania) were still present. Southern Africa during
this period in time may thus be regarded as a zoogeographic counterpart of the
Indian subcontinent, with tropical Africa as a centrally situated region in
which much of the development of the Ethiopian mammalian fauna took place.
East Africa had an African bovid fauna at this time, but boselaphines and ovi-
bovines survived in southern Africa and hippotragines, reduncines and alcela-
phines survived in India.

In addition and based on purely negative evidence, the late Pliocene fauna

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 21

of at least the extreme south of southern Africa had yet to be supplemented by
mammals such as Hippopotamus, although they were already widespread in
East Africa at the same time.

It is certainly not surprising that the fauna of an area situated at a conti-
nental extremity, with no possible access to other land masses, should develop
certain unique characteristics. Similarly it is to be expected that the most
southerly part of South Africa would itself have a fauna which differs in certain
respects from that of the rest of the subcontinent, especially in view of the
factors discussed earlier. Interpretation of the southern African fossil record,
and that of the Cape Biotic Zone in particular, must, therefore, take zoogeo-
graphical factors into account.

22 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fosst. MAMMAL OCCURRENCES IN THE SOUTH-WESTERN CAPE PROVINCE

General observations on the more important fossil occurrences in the south-
western Cape have already been published (Hendey 1969), while a few more
detailed accounts of some are also available (e.g. Singer 1957; Hendey 1968).
Some earlier statements require revision in the light of more recent investi-
gations and some occurrences not previously recorded will be mentioned in the
course of the present report. An account is therefore given of those which are
relevant.

A. QUATERNARY OCCURRENCES

1. Coastal middens

A large number of coastal hominid occupation sites dating from the Late
Stone Age are recorded in the south-western Cape. All are Holocene in age
and all those specimens from coastal middens which will be referred to are
likely to date from the latter part of this epoch. The earliest date recorded for
an excavated midden in the south-western Cape is 3220 +55 B.Pp. (Van Noten
1967) and it is unlikely that any of the relevant specimens are older than this.
They may in fact be much younger, since the local middens were still being
added to early in the historic period (fost 1500 A.D.).

Although the middens are a potentially useful source of dateable faunal
material, such specimens as were available were acquired in a haphazard
manner and are thus a poor reflection of the numbers and variety of mam-
malian remains represented in the middens. Only three carnivore species,
namely, Arctocephalus pusillus, Mirounga leonina and Canis mesomelas, were repre-
sented in the South African Museum’s collections from local middens and only
the first-mentioned is common.

2. Fish Hoek (34° 7S, 18° 25’E)

There are a number of sites of archaeological interest near Fish Hoek
in the Cape Peninsula, notably Peer’s or Skildegat Cave. Areas of wind erosion
on the northern and southern slopes of the ridge which includes Peer’s Cave
have yielded archaeological and palaeontological material from time to time.
Such material from the northern slope includes a leopard mandible which is
described in this report. Although this specimen is regarded as Holocene in
age, others from the same vicinity apparently date back to the Pleistocene.
The latter include the fragmented skull of a Megalotragus, the giant alcelaphine
which is not known to have survived into the Holocene. Although this specimen
and the leopard mandible appear identical in preservation, they are not
necessarily contemporaneous.

3. Tygerfontein (Approximately 33° 23'S, 18° 12’E)

This is one of the many localities in the region from which only a single
specimen is recorded, in this instance an almost complete skull of Mellivora
capensis, which was presented to the South African Museum in 1968. The

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 23

farm Tygerfontein is situated on the coast approximately 65 km north of
Cape Town, but nothing is known of the mode of occurrence of the skull and it
was apparently without associations. It is regarded as being Holocene in age.

4. Cae ieee Saldanha (33°S, 17° 57’'E)

This previously unrecorded locality is situated on the north-western shore
of Saldanha Bay and has yielded a fairly large assemblage of fossils. They occur
in small shelters in a limestone cliff immediately west of the Sea Harvest
Corporation factory. The occurrences are strikingly similar to those at Swartklip
(Hendey & Hendey 1968), except that the deposits incorporating the fossils are
largely unconsolidated. The few that are in consolidated deposit are
undoubtedly older than the remainder and they could date back to the Pleisto-
cene, but they constitute an insignificant proportion of the available assemblage
and none is included in the systematic descriptions of this report.

The nature of the occurrences suggests that, as at Swartklip, the fossils
were accumulated in animal lairs. A feral cat has in fact been observed occupy-
ing one of the shelters and this animal may still be adding the remains of its
prey to the bone accumulations. There are no indications that any of the fossils
accumulated as a result of hominid activity and the shelters are small enough
to preclude the possibility of hominid occupation. They may, however, have
been larger before the cliff in which they are situated was eroded to the extent
it is today. This appears to have been the case with one shelter near the north
end of the cliff and which is now filled with partly consolidated deposit incor-
porating marine shells and stone artefacts. This shelter clearly was a hominid
occupation site, but unlike the others it has produced no fossils other than the
marine shells.

Both Jarge and small mammals are included in the Sea Harvest assemblage,
while non-mammalian remains are comparatively rare. Very large mammals,
such as elephant and rhinoceros, are represented by skull parts of very young
individuals, although medium- and small-sized mammals are represented by
both adults and juveniles. In general the preservation of specimens is good,
although cranial remains tend to be fragmentary. Postcranial bones are often
complete in all respects.

Almost all the species recorded are known to have occurred in the south-
western Cape in historic times. Exceptions are Redunca, Connochaetes and Anti-
dorcas. Most species are apparently indistinguishable from their extant
counterparts, although the Carnivora include varieties which are larger than
the modern forms and the Perissodactyla include the extinct species, Equus
capensis.

As at Swartklip, a wide variety of carnivores is represented and some, if
not all of these species are likely to have been occupants of the shelters. Although
much of the assemblage probably accumulated as a result of carnivore activity,
other factors are undoubtedly involved. For example, a few of the fossils have
been gnawed by porcupines and Hystrix is included in the assemblage. This

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

species is also regarded as a likely one-time occupant, fossils having been added
to the assemblage as a result of its bone collecting activities. In addition, some
specimens almost certainly represent the remains of animals which died in the
shelters.

Although the Sea Harvest and Swartklip assemblages are essentially
similar in the variety of species represented, there are at least two striking
differences in the nature of the assemblages.

Firstly, the Sea Harvest assemblage is characterized by an appreciably
higher proportion of small mammals. These include large numbers of rodents
(e.g. Bathyergus, Georychus, Otomys), a hare (Lepus) and dassie (Procavia). The
latter is particularly common, but is not represented at Swartklip at all. The
cranial remains of this species are very fragmented, which suggests that the
animals were the prey of some carnivore and do not merely represent the result
of natural fatalities in a dassie lair. Although the Sea Harvest and Swartklip
sites are in essentially similar situations adjacent to the present coastline, the
differences in the numbers and varieties of small mammals represented may be
taken to indicate one or both of the following factors:

(1) ‘The environment of the two localities differed at the time that their respective assemblages
were accumulated and they cannot, therefore, be contemporaneous.
(2) ‘The species responsible for adding to the two assemblages were not the same. ;

Confirmation of the first factor is afforded by the other major difference
between the two assemblages. Both penguins (Spheniscus) and seals (Arcto-
cephalus, Lobodon) are recorded from Sea Harvest, but no marine animals are
known from Swartklip, indicating that the sea was close to the former site at the
time when the fossils in the unconsolidated deposit were accumulated and that
it was some distance away at the time that the Swartklip fossils were accumu-
lated. There is a parallel to this situation in the faunal record of a single site
in the southern Cape (Nelson Bay Cave), which was recently described by
Klein (1972). There the advent of marine faunal remains in the succession was
correlated with a rise in sea level from a late Pleistocene minimum to a level
similar to that of the present early in the Holocene. On this basis the Sea
Harvest fossils were tentatively regarded as Holocene in age, while those from
Swartklip were regarded as late Pleistocene. This conclusion is supported by
other faunal evidence, including the nature of some carnivore species repre-
sented (vide infra).

5. Swartklip (34° 5’'S, 18° 41’E)

The occurrences at Swartklip on the False Bay coast have already been
dealt with in some detail (Hendey & Hendey 1968), although a considerable
number of additional specimens have since been recovered and some of the
earlier statements about the fauna and its associations require revision. The
Swartklip faunal list has, however, not been significantly altered. Previously
unrecorded species include two carnivores (Vulpes chama, Felis libyca), which
are described elsewhere in this report.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 25

In the 1968 report a distinction was made between the material from
Swartklip described by Singer & Fuller (1962) and Ewer (1962) and that from
Sites 1 and 2. It was recently established that the ‘Singer/Fuller Occurence’
was actually a block of fossiliferous deposit which had fallen on to the present
beach from Site 1 (i.e. that exposure from which the greater part of the Swart-
klip assemblage is derived). Almost all the specimens from this locality which
are described in this report are from Site 1, the only exceptions being some
belonging to Canis mesomelas and Hyaena brunnea which are from Site 2. The
source of these specimens is indicated in the relevant specimen lists.

An additional point regarding the origin of at least a part of the Site 1
assemblage is that the porcupine must also have been involved in the bone-
collecting at the site. A few of the Site 1 fossils have now been found to show
porcupine gnaw-marks, so it is likely that at least a part of the assemblage
resulted from the bone-collecting activities of this animal. The greater part of
the assemblage is, however, still regarded as representing the results of carnivore
activity and the remains of animals which died in the shelters.

The Swartklip fauna was reported in 1968 to be late Pleistocene or Holocene
in age, but the latter alternative is now dismissed in the light of evidence
discussed in connection with the Sea Harvest occurrence and the fact that
ostrich eggshell from Site 1 has given a C14 date of more than 40 000 years
(I-6840). In addition the geological context of the fossiliferous deposits suggests
that they date from the earlier part of the last glacial (K. W. Butzer, pers.
comm.). The statement that ‘the lairs in which the fossils accumulated were
(possibly) still being occupied after the arrival of the first settlers from Europe
in 1652’ (Hendey & Hendey 1968: 71) can no longer be accepted, although it is
fairly certain that there was at least one hyaena lair in the area at some time
during the historic period.

Of all the more important assemblages from the south-western Cape, that
from Swartklip is the least problematical. The fauna has an essentially ‘modern’
character but the species represented are in many cases sufficiently different
from their extant counterparts to indicate that the fauna must be pre-Holocene
in age. This applies in the case of the carnivores which are to be described later.
Some species are sufficiently well-represented to allow good definition of their
dental and osteological characters and the Swartklip fauna provides a great
deal of information on the nature and character of the local late Pleistocene
mammalian fauna, exclusive of the very small species.

6. Lime Quarry, Saldanha (Approximately 33°S, 17° 57’E)

A number of specimens acquired by the South African Museum in 1918
and 1919 are recorded as being from a lime quarry, south of Hoedjies Bay,
which is the old name for the town of Saldanha. Cooke (1955: 166) has already
made reference to this assemblage, part of which is now lost.

This occurrence is clearly not the same as the Sea Harvest one, although
the two must be in the same general area. The preservation of the Lime Quarry

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

fossils suggests that they are older than those from Sea Harvest. The Lime
Quarry assemblage also includes the seal, Arctocephalus, which indicates that it
too must date from a period when relative sea level was similar or higher than
at present. The Lime Quarry Arctocephalus is, however, a variety which is
distinct from the extant A. pusillus and this suggests that it might be pre-Holocene
in age. Unfortunately the characters which distinguish the Lime Quarry
Arctocephalus from A. pusillus cannot be observed in the Sea Harvest material,
so that the relative ages of the two faunas cannot be determined on these
grounds. It is, however, tentatively concluded that the Lime Quarry fauna is
earlier and that it probably dates from the latter part of the Pleistocene.

7. Melkbos (33° 40'S, 18° 26’E)

This occurrence has already been dealt with in detail (Hendey 1968)
and little of significance has been added to the assemblage. One previously
unrecorded species which was recently recognized is the -giant alcelaphine,
Megalotragus, and this lends support to the inferred Pleistocene age of the
fauna from the site. The view that it probably predates the Swartklip assem-
blage is maintained.

The possibility that the fauna has Middle Stone Age associations requires
further qualification since Early Stone Age artefacts have recently been. found
in surface association with the fossils. The cultural associations of the fossils can
now be established with certainty only by controlled excavations at the site.

8. Bloembos (Approximately 33° 17'S, 18° 11’E)

Reference to this site has already been made by Cooke (1947, 1955) and
nothing further can be added to his comments. No additional specimens have
been acquired from this locality since 1906.

g. Elandsfontein (33° 7'S, 18° 14’E)

The fossil occurrences on the farm Elandsfontein near Hopefield are
probably the best known in the region. Many of the species, and their geological
and archaeological associations have already been described (see Singer &
Wymer, 1968).

The age of the Elandsfontein fauna is more problematical than would
appear from the various publications relating to the site. It is usually regarded
as a single unit dating from the Vaal-Cornelia Faunal Span of the South African
Quaternary (e.g. Cooke 1967), but it is evident that the specimens are not all
contemporaneous, a fact which is clearly demonstrated by the carnivores
which are described in this report. Artefacts from the Early, Middle and Late
Stone Ages, as well as some dating from the historic period, have been recovered
in surface association with fossils and the suggestion has been made that some
elements of the fauna may be associated with each of these periods of hominid
occupation (Hendey 1969). Most of the fossils are apparently contemporaneous
or broadly contemporaneous with the Early Stone Age (‘Final Acheulian’)

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE =

occupation and the only definitely recorded cultural/faunal association relates
to this period in time (Singer & Wymer 1968).

In contrast to the other faunas already mentioned, that from Elands-
fontein includes a number of extinct genera (e.g. Szmopithecus, Megantereon,
Libytherium), as well as a larger proportion of extinct species. Almost all the
identified species have already been recorded elsewhere, but the Bovidae
(A. W. Gentry & L. H. Wells, in preparation) and the Carnivora (vide infra)
do include some apparently unique species.

The Elandsfontein carnivores have already been the subject of one detailed
study (Ewer & Singer 1956), but many additional specimens are now available
and a number of species not previously recorded at the site are known. Although
most of the recorded carnivore specimens from Elandsfontein are described in
this report, some specimens were not available as they are no longer in the
South African Museum’s collection.

8. Elandsfontein Wes (33° 7'S, 18° 13’E)

A few kilometres west of the Elandsfontein site is a smaller wind-eroded
area from which a few fossils have been recovered. Such occurrences are not
uncommon in the Sandveld and that at Elandsfontein Wes is included because
Hyaena brunnea is represented in the assemblage. On the basis of its characteristics
this material is concluded to be late Pleistocene in age.

B. LATE TERTIARY OCCURRENCES

There is only one late Tertiary occurrence relevant to the present report.

Langebaanweg (32° 58'S, 18° 9’E)

Since the first reported discovery of fossils in the phosphate quarries at
Langebaanweg (Singer & Hooijer 1958), these quarries have become one of the
most prolific sources of fossil vertebrates in South Africa. The remains of a
large number of marine, fresh-water, terrestrial and flying vertebrates have
been recovered, the fossils coming principally from the only quarry which is
still being mined, namely, ‘E’ Quarry (Hendey 1970a). Some invertebrates,
coprolites and fossil roots are also known. The fauna is significant not only
because of its abundance and diversity, but also because it is the only one of
Pliocene age being actively investigated in southern Africa (Hendey 1970), 1972c¢,
1973b). It predates the more widely known South African australopithecine
sites (Brain 1970), and is comparable in age to East African faunas such as
Koobi Fora I (Maglio 1971) and Kanapoi (Patterson 1966).

Several phosphate occurrences are recorded in the south-western Cape
(Du Toit 1917; Haughton 1932a), but those at Langebaanweg are the only
ones being commercially exploited at present. Unpublished studies on the
geology of the Langebaanweg area have been undertaken in addition to those
referred to above (African Metals Corporation and Chemfos Limited records),
and a further detailed study is in progress (A. J. Tankard, in preparation).

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

On the basis of the geological and palaeontological information presently
available, the deposits at Langebaanweg are categorized as follows:

Varswater Formation—An occurrence of largely unconsolidated clastic sediments located on the
farms Varswater and Langberg Suid and which are rich in phosphate. The deposits are
made up mainly of marine, estuarine and terrestrial sands in which fossils of late Pliocene
age are incorporated. The largest exposures of these deposits are at the scene of current
mining operations (‘E’ Quarry), while those in an earlier open-cast mine (‘C’ Quarry)
are now obscured by water and sand infill.

Unnamed deposits— Mined-out phosphate occurrences located on the farm Langberg, approxi-
mately 2 km east of the Varswater deposits which yielded fossils of late Pliocene age as well
as some of apparently (? early) Pleistocene age. This mine (Baard’s Quarry) has now
been back-filled and the relationships of these deposits to the Varswater Formation are
uncertain, although records suggest that those which contained the Pliocene fossils were
fluviatile in origin. (Table 3)

TABLE 3

The stratigraphy of the Langebaanweg area.

Varswater area Langberg area
Age (‘E’ Quarry) (Baard’s Quarry) —
Pleistocene/Holocene . . . Surface bed Surface bed
Pleistocene!) =" =a. ec — Unnamed deposits
Bed 3b
PHOGERE: os. Sy oe 8, =e: ee ee = Unnamed depoat

Bed 1

The most significant palaeontological discoveries have been made in ‘E’
Quarry (Fig. 3). Although a detailed account of the geology of this site will
be published elsewhere, observations made in the course of the fossil recovery
program are summarized here.

DEPOSITS OF “E’ QUARRY (Table 4)

The exposed deposits consist largely of unconsolidated or partly consoli-
dated medium-grade sands in which the commercially exploited granular
phosphate occurs. These deposits are fossiliferous only in the lower levels.
They are underlain by a clay of undetermined thickness and overlain by more
recent aeolian sands which vary in depth from about 2 to over 40 metres.
There are considerable differences in the appearance of the ‘E’ Quarry sediments
both vertically and horizontally, and there are some important differences
in the succession in the eastern and western parts of the quarry. The two areas
are separated by a drainage channel (‘Main Stream’) which runs from near
the north wall to the south-west, following the general dip of the deposits.
In most parts of the quarry mining has extended below the level of the water-
table, which has fallen as a result of dewatering of the mine. Main Stream has

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 29

tae
4

WEST WALL

SCALE: 1:5000

Fig. 3. Sketch plan of SE? Quarry, Langebaanweg (July, 1972), showing collecting areas.
The deposits exposed are as follows:

Bed 3a— West Wall
Bed 2 —All areas
Bed 1 —1st Sump, 2nd Sump, South Wall, Far East and July 1971 areas.

become progressively less active and is now ephemeral. It is, nevertheless, a
significant feature, since apart from being the boundary between the east and
west successions, the deposits of its bed and banks differ somewhat from those
elsewhere in the quarry in that they are coarser-grained and completely
unconsolidated. Main Stream may follow a drainage line of considerable
antiquity.

Three stratigraphic units are recognized in the ‘E’ Quarry exposures of
the Varswater Formation (Fig. 4).

30 ANNALS OF THE SOUTH AFRICAN MUSEUM
TABLE 4

The stratigraphy of ‘E’ Quarry, Langebaanweg.

Stratigraphic Depositional Faunal
Age Unit Lithology Environments Unit

Pleistocene/Holocene . Surface bed Sands, calcrete, Terrestrial —-
etc.

Bed 3b | Medium-grade —

ZG — —-—-—] sands Estuarine |—-—---—-----
e Bed ga | (Phosphatic) Estuarine
EH 7,5 m Faunal Unit 2
< )
5 Medium-grade Estuarine Estuarine

Pliocene © Bed 2 sands and Faunal Unit 1
~ 2m Terrestrial
ea
FH Sand with
S boulders,
N Bed 1 cobbles and Marine Marine
E pebbles of Littoral Faunal Units
> phosphate rock 1 and 2

Im
5 — Clay ? —
Bed 1

The lowest horizon is characterized by the presence of phosphate rock.
It was originally believed to underlie those deposits in which the bulk of the
vertebrate fossils occur (Hendey 1970a), but subsequently it was thought to
postdate and laterally truncate these deposits (Hendey 1970b). The original
conclusion has now been shown to be correct, although it is still uncertain
whether or not it underlies the whole of the next horizon in the succession.

In the absence of detailed petrographic studies it is difficult to interpret
the origins and development of Bed 1, since it evidently had a complex history.
However, at least two generations of phosphate rock are represented. ‘The first
is an extremely well-indurated rock, brown in colour, which occurs as large
irregularly-shaped boulders with abraded surfaces, around which are scattered
well-rounded cobbles and pebbles of the same material. Rare instances are
known of embedded and unidentifiable bone fragments. The age and mode of
origin of the rock is still conjectural.

Interspersed with this indurated phosphate rock is a partly cemented
brown phosphate rock, which locally forms a matrix embedding elements of the
older and more indurated rock. A suite of well-preserved marine invertebrates
has been recovered from the second phosphate rock (Marine Faunal Unit 1)
(Kensley 1972). The invertebrates included in this fauna represent both rocky
and sandy shore species. Their presence indicates that a marine shoreline

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

WEST | EAST
SECTION | SECTION
Elevation

50m

gin biel ay

Elevation the
40m eran nue
FORMATION
sos 3b
Phosphatic
sand
BED 3a "
2.5m
Re ahatic BED 2
sand
Phosphate

Phosphatic
sand

Silcrete

Non-
phosphatic

sand

Phosphate

rock in
see :
ps" |

ai Not to scale

Fig. 4. The stratigraphy of ‘E’ Quarry, Langebaanweg.

32 ANNALS OF THE SOUTH AFRICAN MUSEUM

existed in the area and that abrasion of the indurated phosphate rock was by
wave action. The partly cemented phosphate rock has also been subjected to
wave action, as shown by the abraded surfaces of infills of this material in
Bed 1, and rounded fragments lie mixed with the cobbles and pebbles of the
indurated rock. The abrasion of the second phosphate rock may be associated
with a second period of marine erosion or with the later part of a single marine
incursion. A wide variety of marine fossils (Marine Faunal Unit 2) is preserved
in an unconsolidated, light-coloured and sandy matrix incorporating the two
generations of abraded phosphate rock. Occasional remains of terrestrial
vertebrates also occur and these are usually heavily rolled.

An exposure of Bed 1 east of Main Stream (BDT 1/1971) revealed that the
rocky element is considerably less prominent than it is in more southerly
exposures of this horizon. ‘The BDT 1 exposure has cobbles and pebbles of
both primary and secondary phosphate rock, the latter being more common,
which is again in contrast to the more southerly exposures. In addition, rolled
fragments of less consolidated and non-phosphatic sandstone were present.

Bed 2

The light-coloured, medium-grade sands of Bed 2, which are exposed
over most of the floor of the quarry, are often not readily distinguishable from
the unconsolidated matrix of the underlying bed. The sediments and fossils
suggest that deposition was primarily in a relatively calm and shallow estuarine
environment, but at least some of Bed 2 may have accumulated subaerially.
In the vicinity of Main Stream, and sometimes also near the base of the horizon,
coarse-grade sands suggest deposition by higher-velocity waters. ‘Towards the
eastern limit of the quarry, Bed 2 becomes very thin, no more than a capping
on Bed 1, and perceptible mainly because of its fossil content.

Bed 2 is largely non-phosphatic and includes large numbers of non-marine
vertebrate fossils, while marine vertebrates are rare (Estuarine Faunal Unit 1).

This horizon apparently began accumulating when a river, which had
previously discharged into the sea elsewhere, altered its course and met the
sea somewhere in the immediate vicinity of the present ‘E’ Quarry. This
probably occurred during a period of marine transgression and Bed 2 is
regarded simply as a facies change in the cycle of deposition which resulted in
the marine erosion and deposition of Bed 1.

In the eastern part of ‘E’ Quarry the upper limit of Bed 2 is marked by a
discontinuously-developed capping of phosphate nodules and phosphate rock.
This rocky horizon was previously confused with Bed 1 and, although termed
the ‘basal marker’ (Hendey 19700), this name is appropriate only in that this
horizon marks the lower limit of mining in this part of the quarry. Many of the
nodules have built up around fossils and crushed and broken fossils at this level
are often cemented together by a phosphatic matrix.

About 150 m east of Main Stream is a smaller and less well developed
drainage course (‘East Stream’). Between these two features and immediately

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 33

below the phosphatic nodules is an extensive and well-developed zone of siliceous
induration of the light-coloured sands of Bed 2. The origins of the nodules and
‘silcrete’ are not known, although the former may be connected with the depo-
sition of the phosphatic sands which overlie Bed 2.

Beds 3a and 36

The phosphatic nodules of Bed 2 are absent west of Main Stream and this
bed is overlain by the brown, phosphatic and fossiliferous sands of Bed ga.

Some of Bed 3a may have been laid down subaerially, but most of it was
apparently deposited by faster-flowing water than was the case with the
subaqueous element of Bed 2. Remains of both terrestrial and aquatic verte-
brates tend to be concentrated towards the base of the bed and become
progressively less common upwards. There are, however, several levels above
the base of the bed where fossils occur in greater numbers and these levels, and
the base of the bed, follow the general south-westerly dip of the deposits. They
may represent old land surfaces over which successive periods of subaqueous
deposition took place, perhaps at times when the river feeding the area was in
_ flood. Even if such land surfaces existed, they were clearly not accessible to
very large land mammals, which is a contrast to the situation prevailing at the
time of the deposition of Bed 2, since such species are commonly represented
in this horizon. The surfaces on which the Bed ga fossils were apparently
accumulated may actually have been subaqueous all the time and the land
mammals represented in this bed may have been carried to their site of
deposition by water.

Unweathered pebble-sized feldspar crystals, rolled feldspar and quartz
pebbles and fragmentary fossils apparently derived from Bed 2 are included
in the sedimentary suite of Bed 3a. Their presence indicates that at least some
of Bed 3a was deposited by strongly flowing water. Bed 3a may have been laid
down along the path of the periodically flooding ‘Langebaanweg River’ rather
than in the backwaters of its estuary as is inferred for Bed 2. Bed ga is far less
extensive in area than Bed 2 and, although its distribution has yet to be finally
determined, there is a suggestion that it has a linear north-east to south-west
spread. If this can ultimately be demonstrated, it will be a further indication of
deposition along or ahead of a river channel.

Bed ga is presently regarded as a different facies of the estuarine sedi-
mentation of the Varswater Formation and, although the zm sztw fossils cannot
always be distinguished from those derived from Bed 2, its fauna is termed the
‘Estuarine Faunal Unit 2’.

Overlying Bed ga, or Bed 2 where ga is absent, are the phosphatic sands of
Bed 3b. This is by far the most extensive unit in the Varswater Formation and
it is largely unfossiliferous. The only fossils ever recovered from it have been
a few isolated giraffid limb bones. Along the west wall Bed 3b can be visually
distinguished from Bed ga only by the fact that it is apparently unfossiliferous
and the subdivision of Bed 3 is made solely on these grounds. Evidently though,

34. ANNALS OF THE SOUTH AFRICAN MUSEUM

the environment of deposition of Bed 3b did differ from that of Bed ga since
animal remains were so rarely added to the accumulating sediments. Differences
between the two units may be detected by detailed examinations of the
sediments, although their lithology is superficially similar.

On the assumption that deposition of the Varswater Formation took place
during a marine transgression, it is evident that a shift of the shoreline to the
north-east with the rise in sea level would mean that the ‘E’ Quarry area was
progressively further from the shoreline and no longer in an area of concen-
trated vertebrate activity. Consequently, although the Langebaanweg River
was still discharging sediment into the area, very few vertebrate fossils were
incorporated into the accumulating deposits.

Bed 3b is easily distinguished from the underlying deposits in the eastern
part of the quarry (Bed 2), because it is non-fossiliferous and phosphatic,
whereas Bed 2 is fossiliferous and non-phosphatic.

The upper limit of Bed 3 is marked by a variably developed zone of
induration and it is always readily distinguishable from the overlying and
non-phosphatic aeolian sands of what is informally termed the ‘surface bed’
(Hendey 19702).

FAUNA OF ‘E’ QUARRY
Marine Faunal Unit 1

The fossil fauna of the partly consolidated phosphate rock of Bed 1 is
known only from a single and very limited occurrence. The invertebrate remains
are, however, very well preserved and have been described in detail elsewhere
(Kensley 1972). A single shark’s tooth is the only recorded vertebrate fossil in
this faunal unit.

Marine Faunal Unit 2

The fossils from the unconsolidated matrix of Bed 1 represent a wide
variety of invertebrate and vertebrate species.

The invertebrate fauna of this unit is represented only by internal casts
and although they have been found at several places, most notably BDT 2/1972.
several metres south of BDT 1, they are less well known than the invertebrates
of Marine Faunal Unit 1.

By contrast, marine vertebrate remains are abundant. Shark’s teeth
are particularly common and an array of species similar to that recorded from
‘C? Quarry is represented (see Hendey 1970a: 96). Sting-ray spines and
denticles, eagle-ray tooth plates, skate denticles, vertebrae and teeth of bony
fish, a variety of whale bones and a single seal femur have also been recovered
from the unconsolidated matrix of Bed 1.

Some fragmentary and usually heavily rolled remains of terrestrial
vertebrates occur in association with the marine fossils. Most frequently
represented is a land tortoise and since this is also the most common fossil in
Bed 2, it suggests that Beds 1 and 2 are broadly contemporaneous. A recently

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 35

discovered Hipparion tooth from Bed 1 is a further indication that this faunal
unit is indeed Pliocene in age.

Estuarine Faunal Unit 1

This faunal unit is characterized by the often very well-preserved remains
of large terrestrial mammals. Smaller vertebrates are actually more commonly
but less obviously represented.

The most common species is a land tortoise (Chersina) and even where the
deposit is poorly fossiliferous, remains of this species are likely to occur Other
non-mammalian vertebrates include sharks, bony fish, snakes, lizards, frogs
and birds. Sharks and bony fish are rare and there was clearly not a concentra-
tion of the remains of these creatures by wave action as was the case in Bed 1.
Birds are represented by a wide variety of species including ostrich, francolin,
plover, penguin and cormorant, all of which are represented by modern
counterparts in the immediate area or on the adjacent coast.

Small mammals are common and include insectivores, rodents, a lago-
morph and small viverrids. Medium-sized herbivores are Hipparion, Nyanza-
choerus and a variety of bovids. Large herbivores are Mammuthus subplanifrons
(Maglio & Hendey 1970), a gomphothere, Ceratotherium praecox (Hooijer 1972),
a sivathere and Giraffa. The larger carnivores include a machairodont, a viverrid
and hyaenids, which are described elsewhere in this report. Also represented
is a seal (Hendey & Repenning 1972).

The larger mammals of this unit are those on which the relative dating
of the Varswater Formation is largely based. Comparisons with radiometrically
dated East African faunas indicate a late Pliocene date for Bed 2 and an age
of about 4 million B.P. is inferred (Hendey 1970), 1972C¢).

A study of the fossil occurrences of Bed 2 (vide infra) has provided some
information on the manner in which the fossils came to be accumulated and
the nature of the environment at the time. Some fossils were clearly laid down
on a land surface, with little disturbance subsequently by geomorphological
agents. Damage to bone has resulted from both carnivore action and fires.
Indications are that those fossils which were accumulated subaerially were
subsequently inundated by relatively calm and shallow water and that sub-
aqueous deposition of deposits and other fossils also took place.

Nothing certain is known of the vegetation of the area at the time that the
fossils were being accumulated, but the abundance and variety of large
herbivores suggests that it was more luxuriant than the present flora. The rain-
fall today averages about 250 mm per annum and falls mainly in winter. It is
sufficient to support only a Mediterranean macchia vegetation without
indigenous trees and little indigenous grass. The presence of Giraffa in the Bed 2
deposits indicates that trees must have been present during the late Pliocene,
while the high-crowned teeth of the Ceratotherium and Hipparion suggest the
presence of grassland as well.

The burnt bone in the deposits probably resulted from bush-fires and,

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

since burnt bone is not uncommon and occurs at most levels, the fires must
have been quite frequent, probably towards the end of a season which was long,
hot and dry. On the other hand the inferred presence of trees and other more
luxuriant vegetation points to a period of good rains as well. Together these
factors indicate strong seasonality in precipitation and it follows that the
Langebaanweg River is likely to have varied markedly in the volume of its
flow according to the season.

The environment visualized on the basis of these facts and inferences is
that of a coastal savanna crossed by a river flowing south-westwards and having
its estuary surrounded by an area of low-lying sandy flats which were flooded
during the rainy season. The marine environment is likely to have been fringed
to the west by islands formed by granite outcrops in what is today the Saldanha/
St. Helena Bay area (see Hendey 1970): fig. 4). Alternatively it may have been a
north-east extension of the present Saldanha Bay, and in either case, a relatively
sheltered area with low-energy waves.

A greater number of land mammals may have concentrated in the area
during the dry season owing to the availability of fresh water in the river. The
hypothetical floodplain may thus have been an area of intense terrestrial biotic
activity during the dry season. Amongst other things predation and scavenging
of terrestrial animals would have occurred, with the remains being buried
beneath subaqueously deposited sediment during inundation of the floodplain
in the succeeding rainy season. Judging from the dispersal of some of the fossils,
the flooding must have been subdued rather than torrential in most of the area
of Bed 2 which is now exposed in ‘E’ Quarry. This area may have had one or
more perennial water courses and the present Main Stream may follow such a
feature.

Estuarine Faunal Unit 2

The fauna of Bed ga is not as well known as that from Bed 2 and it is
problematical since it is not always possible to distinguish those fossils which are
in situ from those that are derived from Bed 2. There are, however, some obvious
differences between the two assemblages.

The large mammals which are so characteristic of Bed 2 are either not
recorded from Bed ga or are known only from very fragmentary remains, most
or all of which may be derived from Bed 2. For example, Ceratotherium praecox
is represented by occasional tooth fragments in Bed ga, but in Bed 2 it is known
from hundreds of complete teeth, partial and complete dentitions, parts of
skulls and elements of the postcranial skeleton (see Hooier 1972). The large
pig, Nyanzachoerus, is not known from Bed ga, although a miniature one, which
is not known from Bed 2, is present. The most common of the Bed 2 bovids,
a boselaphine, is not recorded from Bed 3a, where the two most common
bovids are aicelaphines. They in turn are not definitely recorded from Bed 2.
Each of the two estuarine faunal units has at least one hyaenid which is not
recorded in the other. The seal, which is rare in Bed 2, is common in Bed ga.

LATE CENOZOICGC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 37

There are also differences in the nature of the fossil occurrences. Even
amongst the zn situ Bed ga fossils, there is a greater proportion of fragmented
bone and there have been fewer indications of close associations of different
parts of single skeletons, although the latter feature may result from the smaller
sample size.

There are probably a number of factors which could have caused the
differences between the two estuarine faunal units. For example, it has already
been suggested that the environment of deposition of Bed ga differed from that
of Bed 2 and this may have contributed to some of the observed differences.
In addition, Bed ga clearly postdates the underlying Bed 2 and, although the
time difference may not have been very great, it may have been sufficient for
changes to have taken place in the composition of the local fauna.

Unless otherwise stated all references in this report to the Langebaanweg
vertebrate fauna relate to the ‘E’ Quarry occurrences.

DEPOSITS AND FAUNA OF ‘C’ QUARRY

The observations which have already been made on the ‘C’ Quarry
occurrences (Hendey 1970a) have not been supplemented. The ‘C’ Quarry
marine fossils apparently occur in deposits which are a westerly extension of the
‘E’ Quarry Bed 1, while the overlying deposits apparently include extensions
of Beds 2 and/or 3. One puzzling feature of the ‘C’ Quarry succession is the
occurrence of terrestrial vertebrate remains below deposits containing marine
fossils (Hendey 1970a: fig. 4), a situation which has not been encountered in
‘E? Quarry.

The actual relationships between the ‘C’ and ‘E’ Quarry deposits will
only be determined as the area between them is mined away and exposures of
the linking deposits become visible.

The few fragmentary carnivore remains from ‘C’ Quarry have already
been referred to (Hendy 1970a: 97), and this material is not described in the
present report since none of the pieces could be positively identified.

DEPOSITS AND FAUNA OF BAARD’S QUARRY

As with ‘C’ Quarry, no further progress has been made with investigations
into the geology of the Baard’s Quarry area. It is, however, clear that fossils of
Pleistocene age are included in the Baard’s Quarry assemblage (Hendey 1972¢),
although most may be contemporaneous with those from the Varswater Forma-
tion, and the earlier element in the Baard’s Quarry succession is tentatively
regarded as a fluviatile facies of the Varswater Formation.

The few specimens from Baard’s Quarry which are included in the syste-
matic section of this report are regarded as belonging with the Pleistocene
element of the fauna from this site.

38 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fosst. MAMMALS OF THE SOUTH-WESTERN CAPE PROVINCE

The late Cenozoic fossil mammal record of the south-western Cape is
potentially good, but at present only the faunas of the latter part of the
Quaternary are comparatively well known. The Pleistocene element of the
Baard’s Quarry assemblage apparently dates from the earlier part of this
epoch, but there are so many uncertainties relating to this occurrence and only
a limited number of poorly represented species are involved, which means that
they constitute a largely insignificant addition to the record. The Tertiary is
known only from the late Pliocene Langebaanweg occurrences. The record is,
therefore, far from complete and an additional disadvantage is that many of
the available fossils have yet to be studied in detail.

In the account which follows, the record as it is presently known is reviewed.
The Carnivora are excluded as they will be dealt with in detail later and of the
remaining orders some are better known than others (Tables 5, 6). For example,
the Chiroptera are completely unrepresented as fossils, whereas the Artio-
dactyla are ubiquitous and often abundantly represented.

The mammalian microfaunal remains are still largely unstudied, but a
cursory examination of the available material has revealed nothing which is
obviously out of character with the modern fauna.

The Chrysochloridae, which are endemic to Africa and which still occur
commonly in the south-western Cape, are represented locally as fossils only
at Langebaanweg. Apparently only a single species occurs and its remains are
common in both Bed 2 and Bed ga. The fossorial habits of this animal probably
contributed to its preservation in these deposits. The fact that it is not recorded
at other local sites is almost certainly due to the nature of the collecting which
has been undertaken, since Langebaanweg is the only local occurrence where a
determined effort has been made to recover microfaunal remains.

The Macroscelididae are another group which are endemic to Africa
and they are also quite.common at Langebaanweg but unrepresented in the
Pleistocene. Once again there is apparently only one species represented at
Langebaanweg and it occurs in both Bed 2 and Bed ga.

The relative abundance of chrysochlorids and macroscelidids in the south-
western Cape Pliocene is of special interest in view of the exclusively African
distribution of these groups and the fact that neither has a good fossil record.
Miocene, Pliocene and Pleistocene macroscelidids are known (Patterson 1965),
while chrysochlorids are recorded from the Miocene of East Africa (Butler &
Hopwood 1957; Butler, 1969) and the Pleistocene of South Africa (see De
Graaf 1960). The Langebaanweg material includes both skull and postcranial
remains and the large number of specimens available should allow for confident
definition of the species represented. Both groups are, however, taxonomically
complicated and the Langebaanweg species are unlikely to add much to the
understanding of their phylogenies.

Soricidae are poorly represented as fossils in the south-western Cape and
once again Langebaanweg is the only site from which they are recorded.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 39

TABLE 5

The late Pliocene mammalian fauna from ‘E’ Quarry, Langebaanweg (May, 1972).*

Marine | Estuarine | Estuarine
Faunal Faunal Faunal Faunal
Unit 2 Unit 1 Unit 2 Unit
(Bed 1) (Bed 2) (Bed 3a) | unknown
ORDERS MENOTYPHLA AND LYPOTYPHLA (IN-
SECLIVORA)
Elephantulus sp.
Soricidae (2 species) .
Chrysochloris sp.
ORDER PRIMATES
cf. Cercopithecidae
ORDER PHOLIDOTA
cf. Manis sp.
ORDER TUBULIDENTATA
Orycteropus sp.
ORDER PROBOSCIDEA
Gomphotheriidae.
Mammuthus subplanifrons
ORDER HYRACOIDEA
Cogcaviaantiqua . . « « 5» « « *
ORDER PERISSODACTYLA
Ceratotherium praecox
Hipparion albertense baardi
ORDER ARTIODACTYLA
Nyanzachoerus sp. ;
Suidae Species B (aff. Diamantohyus)
Libytherium olduvaiense
Giraffa cf. gracilis .
Tragelaphus sp.
Bovini (aff. Pelorovis)
Boselaphini (aff. Taaachortaa)
Reduncini_. hon, |, Aor ee x
Alcelaphini Species A ake ae tae at
PicelapmimkepeciesB . 2°. kl x
Neotragini (aff. Raphicerus) Toh ae
(ecee au. danhoepent =. ww lk le x
Incertae sedis
ORDER LAGOMORPHA
Incertae sedis
ORDER RODENTIA
Bathyergidae (2 species) eer we
Muridae and perhaps others (several
species) .
ORDER CETACEA
DPRCMPNESERC ER yo ek BM x

x XK X
2 OK OS

x x x

x xX

x X
Kw

x

~wy X

Xx XK XK XK

x xX x x
x x x

x
x

* Excluding Carnivora.

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 6

The Quaternary mammalian fauna of the south-western Cape Province (May, 1972).!

Baard’s

Quarry, | Elands- Swart- Sea Historic

Lange- | fontein |Melkbos| klip |Harvest,| Period
baanweg” Saldanha

ORDER PRIMATES
Simopithecus oswaldi hopefield-
ensis : es om ee
OPO Ursinus 3 isn ee Xx
Homo sapiens rhodesiensis . . x
Homosapicnsn a - s x
ORDER PHOLIDOTA
Miantsisn as i) se x
ORDER TUBULIDENTATA
Onjeteropus afer <= 5 ws =. x |
ORDER PROBOSCIDEA
SStCZOQONE SPs. 6) 3) ee x
Loxodonta atlantica . . . . x :
Koxodoniaajmcanad 3) x x x
ORDER HYRACOIDEA
J EOGLIE GL TG OSISE Sh Be x x
ORDER PERISSODACTYLA
IDET OS WEDS Se te oe x
Ceratotherium simum :
Equusictaicapensis, “5 3: x
Equus cf. plicatus tot
JIGS FAA, BB ee ef. x
ORDER ARTIODACTYLA
Potamochoerus porcus. . . .
Mesochoerus paiceae \ Probably
Mesochoerus lategam ; conspecific
Phacochoerus aethiopicus. . . ?
Tapinochoerus meadows .
Hippopotamus amphibius
Libytherium olduvaiense . . . ?
Tragelaphus cf. strepsiceros
T aurotragus oryx
Pelorovis sp. Seer
SYNGCTUS SPaet fa, na ake x
Syncerusicayien- . . .
Redunca arundinum subspp.
Aippotragus gigas
Hippotragus leucophaeus .
? Beatragus sp. .
Damaliscus niro . :
‘Rabaticeras’ arambourgi
Megalotragus sp.
Connochaetes sp(p). . . . .
Alcelaphus buselaphus caama ch x
Syluicapra grimmia . . . . x
Raplicerus spp) 45 - x x x ~<
Raphicerus campestris
Raphicerus melanotis .
Oreotragus oreotragus . . . x

x OS XK
x
x

ba)

*K 1% KK KX x
ae)

e)
oo

KS OK OK OK OK OK

a

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 4I

TABLE 6 (continued)

Baard’s

Quarry, | Elands- Swart- Sea_ | Historic

Lange- | fontein |Melkbos} klip |Harvest,| Period
baanweg? Saldanha

ORDER ARTIODACTYLA (continued )
Gazella sp. .
Antidorcas recki .
Antidorcas australis .
Pelea capreolus dene Ears
Bovidae incertae sedis (2 species)
ORDER LAGOMORPHA
Lepus sp.
Lepus capensis
Lepus saxatilis
ORDER RODENTIA
Bathyergus suillus
Georychus capensis
Hystrix africae-australis .
Others .

MS PSPS PS OS
x
x
v

x
x
x X

x &X x
x

x & x

w

WS PS Pe PS

OS

ADDITIONAL SPECIES:
Elephas transvaalensis(=1iolensis) from near Melkbos
Giraffa camelopardalis from Bloembos

1 Excluding Insectivora, Chiroptera, Carnivora and Cetacea.
2 Pleistocene species only.
3 Indirect evidence only.

Compared to the chrysochlorids and macroscelidids the amount of material
which is available is limited, but in this instance at least two and possibly more
species are represented. Soricids are not uncommon as fossils elsewhere and a
detailed study of the Langebaanweg specimens should complement the studies
on those from the early Pleistocene of the Transvaal (see Meester 1955).

Rodents are more commonly represented as fossils than the insectivores
and, although especially abundant at Langebaanweg, they are also known
from local Quaternary occurrences.

About a dozen species are known from Langebaanweg and the most
commonly occurring forms are Bathyergidae, a family with an essentially
African distribution. As with the chrysochlorids, it is probably their fossorial
habits which has led to their being more commonly preserved as fossils than
other contemporary small mammals. Bathyergids are also known from all the
more important Quaternary occurrences of the south-western Cape, their
relatively large size and abundance probably having contributed to their
discovery.

Their fossorial habits raises the possibility that they may sometimes become
accidentally associated with fossils already buried in the ground, a situation
which may easily arise because most of the local occurrences are in unconsoli-
dated deposits in areas where bathyergids still occur. This possibility has

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

already been mentioned in the case of the Melkbos bathyergid (Hendey 1968:
112), and also applies particularly in the case of the specimens from Elands-
fontein. Owing to the cliff-side situation of the Swartklip and Sea Harvest
sites, and the consolidated deposits of the former, there is less likelihood of
non-contemporary bathyergids being associated with the rest of the fauna.
It is also unlikely that the Langebaanweg bathyergids are later intrusive
fossils since the levels at which they occur have been deeply buried ever since
deposition of the Varswater Formation ceased. In addition, the species repre-
sented evidently differ from those of the Quaternary and there is no reason
to believe that they are inconsistent with the Pliocene age of the deposits.

One of the Langebaanweg species is apparently a small form of Bathyergus,
while the other, which is much less common, is tentatively referred to Georychus.
Both these genera are still commoniy represented in the south-western Cape
today, and both are recorded from the local Quaternary occurrences.

The other small rodents from sites in the south-western Cape are, for the
most part, unclassified. Otomys is recorded from some of the Quaternary occur-
rences and still occurs in the region today. It is, however, not known from
Langebaanweg, which is surprising in view of the nature of the environment
at the time of the deposition of the Varswater Formation.

Another notable absentee from the Langebaanweg fauna is the porcupine,
Hystrix. This is a rodent of particular significance in local palaeontological
investigations, since its bone collecting habit has resulted in it being at least
partly responsible for some of the important fossil accumulations (e.g. Elands-
fontein, Swartklip, Sea Harvest). Even when skeletal remains of this animal
are not recorded in an assemblage, its presence may be deduced from the very
characteristic gnaw-marks it leaves on the bones which it has collected. The
fact that not a single one of the many thousands of bones recovered at Lange-
baanweg shows any signs of porcupine gnaw-marks suggests that this animal
did not occur in the immediate vicinity in late Pliocene times. By contrast,
porcupine gnaw-marks are a not uncommon feature of the fossils from Elands-
fontein (Singer 1956), one of the few local sites from which porcupine skeletal
remains have also been recovered.

Several recent porcupine lairs have provided an invaluable record of the
historic period fauna of the Cape Folded Mountains and southern Cape regions,
but no significant assemblages of such material from the south-western Cape
was available in the course of the present study.

The Lagomorpha are another group of small mammals which are, as yet,
unstudied. They are not particularly well represented locally and even in
recent times they appear to have been a relatively insignificant element in the
small mammal fauna.

One of the more remarkable features of the fossil and modern faunas of the
south-western Cape is the small number of primate species and specimens
which are represented. The only primates, other than Homo, which are definitely
recorded locally are baboons.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 43

Papio ursinus still occurs in the region today, but no Pafio has been recorded
locally as a fossil and the only fossil baboon which is known is the Elandsfontein
Stmopithecus oswald: (Singer 1962). This giant cercopithecoid was almost certainly
not arboreal and consequently its association with the remains of other terrestrial
mammals is not surprising. Remains of this species are comparatively rare,
although specimens in addition to those already described have been found.
Cranial remains of nine and possibly ten individuals are now known (Q. B.
Hendey, unpublished manuscript), but the additional specimens are no longer
in the South African Museum’s collections. The Elandsfontein Simopithecus
was described as a distinct subspecies and it is apparently the most recently
occurring representative of the genus in South Africa. A few isolated teeth from
Langebaanweg are tentatively identified as belonging to a small cercopithecid.

If arboreal primates were ever present in the south-western Cape, it is
highly unlikely that their remains would be found at the more important fossil
sites. Such species are in any case rare as fossils and in the south-western Cape
it is probable that wooded areas would always have been confined to more
mountainous parts, away from the areas in which the principal fossil occurrences
are located. Palynological evidence from Elandsfontein suggests that trees
might have grown there in the past (Singer & Wymer 1968), while the presence
of Giraffa at Bloembos and Langebaanweg suggests the presence of trees in
these areas as well. However, this does not necessarily indicate suitable habitats
for arboreal primates.

Both orders of Old World ‘edentates’, the Pholidota and Tubulidentata,
are represented in the south-western Cape fossil record. The former are known
from a braincase from Elandsfontein and some postcranial bones from Lange-
baanweg, while the latter are represented by a single cheektooth from Lange-
baanweg. Neither of these groups has a good fossil record and the local specimens
are of little significance, except in as far as past distributions are concerned.
Pangolins are not recorded in the historic period fauna of the south-western
Cape, but aardvarks still occurred locally until fairly recently.

Hyracoidea are known from several local fossil sites. Procavia capensis is well
represented in the Sea Harvest assemblage and is also known from the Saldanha
Lime Quarry site, while a few isolated teeth, tentatively referred to Procavia
antiqua, are recorded from Langebaanweg. It is unlikely that the Langebaanweg
species can be positively identified on the basis of the material presently avail-
able, but it is in no way inconsistent with P. antiqua as defined by Churcher
(1965). This would be the earliest record of an otherwise Pleistocene species,
but its presence is not unexpected if Churcher’s phylogeny for the group is
taken into consideration.

Fossil Proboscidea, which have proved so useful in many parts of the world
for relative dating purposes, are not particularly well represented in the south-
western Cape. During the Pliocene and Pleistocene the Elephantidae arose and
diversified in Africa, giving rise ultimately to the African elephant (Loxodonta
africana), the Asiatic elephant (Elephas maximus) and the woolly mammoth

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

(Mammuthus primigenius) (Maglio 1970a). All three lineages are represented in
the south-western Cape and, in addition, gomphotheres are recorded from
ILangebaanweg and from Milnerton, near Cape Town.

The Langebaanweg gomphothere has been referred to the genus Anancus
(Hendey 1970a), but its relationships are actually uncertain, as are those of the
Milnerton species. The latter is known only from a single tooth fragment, while
the Langebaanweg species is represented by a number of complete teeth and
several tooth and tusk fragments.

The Langebaanweg elephant is a primitive form of Mammuthus subplanifrons
(Maglio & Hendey 1970), and is one of the earliest recorded members of the
Mammuthus lineage. Its identification contributed towards the recognition of the
Pliocene age of the Langebaanweg fauna. The Langebaanweg species was
identified largely on the basis of the remains of a single individual, of which the
mandible, an isolated upper molar and a number of elements of the postcranial
skeleton are now known. Although this material is superior in some respects to
previously recorded M. subplanifrons specimens, the skull is unknown and
‘the reference of the Langebaanweg elephant to Mammuthus subplanifrons is
more a matter of necessity than one of direct evidence’ (Maglio & Hendey
1970: 87). ,

This additional record of M. subplanifrons lends support to the observation
that it is an essentially southern African species (Maglio 1970a), while Loxodonta
adaurora was the ‘dominant elephant during the late Pliocene and early Pleisto-
cene of east Africa’ (Maglio 1970b: 19, 20). The latter was subsequently
replaced in East Africa by Elephas recki, while the apparent derivative of M.
subplanifrons was the North African M. africanavus.

By the Pleistocene Elephas had also appeared in South Africa and there is a
single record of this genus in the south-western Cape, namely, E. transvaalensis
(=z2olensis; see Cooke & Maglio 1972) from near Melkbos (Hendey 1967).
The actual age and associations of the Melkbos specimen are unknown, although
it is unquestionably a Pleistocene species.

The elephant remains from Elandsfontein have been the subject of a
number of unpublished studies, but the status of this material is still uncertain.
The accompanying faunal list (Table 6) reflects the view that only one species,
namely, Loxodonta atlantica, is represented (Cooke & Maglio 1972).

With a single exception, all other elephant fossils from the south-western
Cape have been referred to Loxodonta africana, a species also recorded in the
area in historic times. The exception is a tooth from Baard’s Quarry which
has been identified as belonging to Stegodon (see Hendey 1970a: 94).

As with most other important fossil sites in Africa, those from the south-
western Cape have their assemblages dominated by the remains of ungulates.
Both perissodactyls and artiodactyls are represented and, as is also usual in
Africa, the latter are predominant.

The black and white rhinoceroses (Diceros bicornis and Ceratotherium simum)
are both recorded as fossils in this region, although only the former survived

Senet ce

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 45

into the historic period. This conservative group of mammals has been repre-
sented by the extant species for most, if not all, of the Pleistocene and the only
extinct rhinoceros species presently recorded in southern Africa is that from
Langebaanweg. This species, which is close to the dichotomy of the Diceros
and Ceratotherium lineages, is actually an early form of the latter and is referred
to the species C’. praecox (Hooijer 1972).

The remains of Equidae occur quite commonly in the south-western Cape
and most of the specimens belong to a large species of Equus. The numerous
isolated teeth, partial dentitions and elements of the postcranial skeleton of this
form appear to represent the same species as the mandible on which Broom
(1909) based his description of FE. capensis. Wells (1959) rejected this name, but
subsequently Churcher (1970: 149) resurrected it and took ‘E. capensis to
include E. helmei, E. cawoodi, E. kuhni, E. zietsmani and some of the teeth referred
to E. harrisi and E. plicatus’. This array of names provides some indication of
the confusion which has prevailed in the nomenclature of South African fossil
Equidae and, while the final word in the controversy over the status of E£.
capensis may still be to come, Broom’s name is retained in the present report since
it seems likely that the E. capensis holotype is conspecific with other later
Quaternary Equus specimens from the same region.

Clearly there was at least one species of now extinct equine of large size
which was present in South Africa through most of the Pleistocene. E. capensis
is recorded from the early Pleistocene Makapansgat assemblage (Churcher
1970), through a series of younger Pleistocene contexts in various parts of the
country and it apparently survived into the Holocene of the south-western Cape.

The Baard’s Quarry Equus was referred to the species helmet by Boné &
Singer (1965) and in view of Churcher’s synonomy it is here listed as E.
capensis (Table 6). The single Eguus tooth previously included with the ‘E’
Quarry assemblage (Hendey 1970a, 1970) is now believed to have come from
Baard’s Quarry (Hendey 1972c¢).

At Elandsfontein the Equidae are represented by a large number of
specimens, including one nearly complete skull. Singer & Inskeep (1961: 23)
state that ‘the majority of specimens . . . may be referred to Equus plicatus’
and that some ‘may belong to E. helmei’ (i.e. E. capensis). In the absence of a
detailed study these conclusions are regarded as tentative.

Apparently only two species are represented in the late Pleistocene and
Holocene assemblages from the south-western Cape. The large EF. capensis
is most common and its most recent record is from the ? early Holocene Sea
Harvest occurrence. The second species is a smaller one and is identified as
E. zebra. No remains of E. quagga or E. burchelli have as yet been positively
identified locally, although these species are commonly recorded as fossils
elsewhere in South Africa. While it is fairly certain that FE. zebra occurred in
the south-western Cape in historic times, it is by no means certain that E.
quagga was present, although it was very common in the adjacent Karroo
and probably also in the west coast region.

46 ANNALS OF THE SOUTH AFRICAN MUSEUM

Since E. capensis apparently did survive into the Holocene of the south-
western Cape, it is possible that this species was the local equivalent of the
plains-dwelling E. quagga and E. burchelli, while E. zebra occupied the inter-
vening mountainous areas and also the mountainous parts of the south-
western Cape itself. HE. zebra may thus have formed a species barrier between
E. quagga in the Karroo and E. capensis in the south-western Cape. A general
mutually exclusive relationship between E. capensis and the plains zebras
cannot, however, be concluded. Elsewhere in South Africa E. capensis has been
recorded in association with both EF. quagga and E. burchelli (see Churcher 1970:
Table 8). A possible explanation of the available evidence is that E. capensis
managed to survive fairly late in the south-western Cape because E. quagga
did not extend its range into this region, but that elsewhere it had earlier failed
in competition with both EF. quagga and E. burchellz.

Yet another curious feature of the record of the South African Pleistocene
zebras is the reported presence of both E. burchelli and E. quagga in the early
Pleistocene of the Transvaal (Churcher 1970), even though the latter is often
regarded as being no more than the most southerly variety of the plains zebra
group. Even if it is accepted that E. quagga and E. burchelli are specifically
distinct, it is difficult to accept that they were recognizably different and could
have co-existed as far back as the early Pleistocene. Although this problem has
no direct bearing on the present discussion, it is worth noting that if FE. quagga
did once extend its range as far north as the Transvaal, its absence from the
fossil record of the south-western Cape is more unexpected than ever.

In spite of the fact that much has been written on the Pleistocene equines
of South Africa and that their systematics have been reviewed several times
(Haughton 1932b; Cooke 1950; Wells 1959), it is evident that there is still
further scope for a further reappraisal of the available material. Until this has
been done, interpretation of past patterns of distribution and determination of
phyletic relationships of the recorded species will be difficult.

Apart from Equus, the only other equid known from the south-western
Cape is Hipparion albertense baardi from Langebaanweg (Boné & Singer 1965).
The described material was from ‘C’ and Baard’s Quarries, but the species is
best represented by the material from ‘E’ Quarry.

The relationships, if any, between the Langebaanweg species and Noto-
hipparion namaquense (Haughton 19326) and the South African Pleistocene
Hipparion steytleri (Churcher 1970) have yet to be clearly demonstrated. Boné &
Singer (1965: 389) included both NV. namaquense and H. steytlert in the synonymy
of H. libycum, although they stated that VV. namaquense ‘may prove to be a tran-
sitional form between the typical Hipparion and the more progressive Hipparion
(Stylohipparion)’. The opinion held by the present author is that WV. namaquense
is earlier and perhaps ancestral to the Langebaanweg H. albertense, which is
earlier and perhaps ancestral to H. steytleri. There is evidently scope for a further
re-evaluation of African Hipparion as well as Equus.

The perissodactyls have not proved particularly useful for relative dating

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 47

purposes in South Africa, although both the Ceratotherium praecox and Hipparion
albertense from Langebaanweg have served to substantiate the inferred Pliocene
age of the Varswater Formation, while the Baard’s Quarry Diceros bicornis
is one of the species which served to confirm that a Pleistocene element is
included in the fauna, a conclusion originally based on the Equus capensis
from this site. None of the perissodactyls can yet be used to indicate precise
relative ages of Pleistocene faunas. The black and white rhinoceroses, E. capensis
and the plains zebras apparently occurred through most of the South African
Quaternary, while Hzpparion steytleri persisted for most of the first half, or more,
of this period. The perissodactyls are, however, potentially useful in palaeo-
ecological studies.

The Artiodactyla are completely dominated in most of the fossil faunas of
the south-western Cape by the Bovidae, although Suidae, Hippopotamidae and
Giraffidae are also recorded locally.

The African late Cenozoic fossil record is remarkable for the variety of
Suidae which are represented, but there are surprisingly few species of this
family recorded in the south-western Cape. It is not even certain that the
bushpig (Potamochoerus porcus) and the southern warthog (Phacochoerus aethiopicus)
were present in the region during the historic period, although they were both
widespread elsewhere in sub-Saharan Africa. The latter species is known from
Sea Harvest by a single tusk fragment, which to date is the only suid fossil
known from a local late Quaternary context.

At Elandsfontein three extinct species have been recorded, namely,
Mesochoerus paiceae, M. lategani and Tapinochoerus meadowsi (Singer & Keen
1955; Keen & Singer 1956). It seems unlikely that there are indeed two species
of Mesochoerus represented and the material thus identified may, at most,
represent no more than two varieties of a single species. A reappraisal of the
available material is required. The Tapinochoerus is very poorly represented.

The only other fossil suids recorded locally are from Langebaanweg. The
most commonly represented is a species of yanzachoerus which was appreciably
larger than the extant bushpig. The other is an extremely small and as yet
unidentified species, comparable in size to the pygmy hog of India, Sus salvanius.
It may have affinities with the small Diamantohyus africanus from the Miocene
of South West Africa (Stromer 1926), a genus which is now also known from
the Miocene of East Africa (Walker 1969), but has yet to be recorded from the
Pliocene. The small species is poorly represented, but the Nyanzachoerus is
comparatively well known. It differs in certain respects from the recorded
East and North African species of Nyanzachoerus (H. B. S. Cooke, pers. comm.),
but has yet to be studied in detail.

The Hippopotamidae are represented in the south-western Cape only by
the extant species, Hippopotamus amphibius. The largest number of specimens are
from Elandsfontein (Hooijer & Singer 1961), while other Quaternary occur-
rences have yielded only limited numbers of fragmentary specimens. This species
is frequently mentioned in historical records and only became extinct in the

48 ANNALS OF THE SOUTH AFRICAN MUSEUM

region during the 19th century.

The absence of Hippopotamus at Langebaanweg has already been remarked
upon elsewhere (Hendey 1970a: 98), and this remains one of the most remark-
able absentees from the fauna of this site. Occurrences of comparable age in East
Africa have yielded large numbers of hippopotamus specimens and the deposi-
tional environment at Langebaanweg in the late Pliocene was such that remains
of this animal would have been preserved if it had been present. The negative
evidence suggests that the Hippopotamidae had not penetrated to the southern
continental extremity by the late Pliocene.

The extant giraffe, Giraffa camelopardalis, which was only recorded as far
south as the Orange River area in historic times, is known locally from a single
tooth from Bloembos (Cooke 1955). Giraffa is otherwise recorded only from
Langebaanweg, where it is represented by a comparatively small species
tentatively identified as G. gracilis.

The short-necked and large-horned Sivatheriinae are represented locally
by what is apparently a single species, Libythertum olduvaiense, at both Lange-
baanweg and Elandsfontein (Singer & Boné 1960). Much more material has
been recovered at Langebaanweg since 1960 and elements of both skull and
postcranial skeleton are represented. With the exception of Ceratotherium
praecox, Libytherium is the best represented large mammal from Bed 2 of the
Varswater Formation.

As is, or was the case elsewhere in Africa, the Bovidae were once numerous
in the south-western Cape and four antelope (Raphicerus campestris, R. melanotis,
Syluicapra grimmia, Pelea capreolus) are amongst the largest of the surviving
indigenous mammals of the region. Bovidae are known from all the more
important local fossil occurrences and, with the exception of the Cephalophini,
all extant African bovid tribes, as well as representatives of nearly all the extant
genera, are represented as fossils. In addition, the only South African repre-
sentative of the Boselaphini, a tribe now restricted to southern Asia, is from the
south-western Cape.

The Tragelaphini are represented by at least three species, one of which
(Taurotragus oryx) still occurred in the region in historic times. The eland is,
in fact, one of the more commonly represented fossil bovids and although all
the known specimens are referred to T. oryx, the Elandsfontein assemblage
apparently includes a more primitive variety of this species.

The kudu, Tragelaphus strepsiceros, is not known to have occurred in the
region in historic times, but an extinct form, which is at least subspecifically
distinct, is recorded from both Elandsfontein and Melkbos (Hendey 1968).
This form has not been recorded elsewhere and it may warrant recognition as a
distinct species, just as Hippotragus leucophaeus is recognized as a species distinct
from Hippotragus equinus.

The small tragelaphine from Langebaanweg may be ancestral to the
modern nyala (Tragelaphus angasi) (Gentry, in Hendey 19702).

Bovini are well represented only in certain assemblages. Although abundant

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 49

in the southern Cape during the historic period and often featuring in historical
accounts of this region, the African buffalo, Syncerus caffer, was apparently rare
in the south-western Cape. S. caffer has been certainly identified only by a single
- horn core from Ysterfontein, south of Saldanha Bay. This specimen does not
appear to be very old. Some postcranial remains from Elandsfontein are
tentatively referred to S. caffer. A large form of Syncerus, which might be
specifically distinct from S. caffer, has been recorded from Méelkbos
(Hendey 1968).

Still larger are the long-horned buffaloes previously referred to Homoio-
ceras, but which are now being included in Pelorovis (A. W. Gentry, pers.
comm.). These belong on a lineage distinct from that of Syncerus and they are
recorded locally from Elandsfontein and Bloembos. The Elandsfontein Pelorovis
assemblage is probably the best of its kind in southern Africa.

The only other bovine recorded in the region is an as yet unnamed and
poorly represented species from Langebaanweg. The only horn core fragments
known suggest that it might be an early member of the Pelorovis lineage. A
bovine of comparable age from East Africa is thought to be an ancestor of
Syncerus (A. W. Gentry, pers. comm.), so the indications are that the two
African bovine lineages were differentiated during the Pliocene. The latest
recorded occurrence of Pelorovis in southern Africa is that from Nelson’s Bay
Cave in the southern Cape, which is dated at about 12 000 B.P. (Klein 1972).
There is no comparable very late Pleistocene record from the south-western
Cape.

The only South African record of a boselaphine is the Pliocene species
recorded from Langebaanweg (Gentry, in Hendey 1970a). This species could
perhaps derive from Protragocerus labidotus from the late Miocene from Fort
Ternan in Kenya (Gentry 1970). The Langebaanweg species is considerably
larger than P. labidotus, it differs from the Fort Ternan species in some morpho-
logical details as well and apparently has affinities with Tragoportax of the
Dhok Pathan stage of the Siwaliks of India. The Boselaphini are an essentially
Eurasiatic group and the available record suggests that only a single lineage,
which is not known to have survived the Tertiary, was present in Africa.

The Reduncini are represented in the south-western Cape by at least three
species. Curiously, this tribe was apparently not represented in the region in
historic times, although Redunca fulvorofula may have occurred in the adjacent
Cape Folded Mountains. The nearest recent record of this species, and of
R. arundinum, is in the southern Cape. R. arundinum, or closely related forms, is
one of the most commonly occurring bovids in the Pleistocene of the south-
western Cape. It is well represented in the Elandsfontein, Melkbos and Swart-
klip assemblages and is also recorded from Sea Harvest. The Elandsfontein
and Swartklip forms differ from another and both differ from modern R.
arundinum. These three probably represent varieties of a single lineage, of
which R. dart: from the early Pleistocene of the Transvaal (Wells & Cooke
1956) may bea more primitive member.

50 ANNALS OF THE SOUTH AFRICAN MUSEUM

Kobus is not recorded in the Quaternary of the south-western Cape,
although it is not uncommon elsewhere in sub-Saharan Africa.

The other two reduncines recorded locally are from Langebaanweg, one
from ‘E’? Quarry and the other from Baard’s Quarry. The former is tentatively
referred to Kobus and while that from Baard’s Quarry was originally identified
as Redunca ancystrocera (Gentry, in Hendey 19702), its affinities are now un-
certain.

No hippotragines were recorded in the south-western Cape in historic
times, although the southern Cape was the last refuge of the recently extinct
HMippotragus leucophaeus. This species is, however, recorded as a fossil in the south-
western Cape. It was initially described as H. problematicus by Cooke (1947)
on the basis of a specimen from Bloembos and has since been recorded from
Swartklip (Hendey & Hendey 1968), Melkbos and Elandsfontein. Confirmation
of the identity of the south-western Cape fossils has come from an increasing
number of undoubted H. leucophaeus specimens from porcupine lairs and hominid
occupation sites in the southern Cape. This species is of special interest since it
was endemic to the most southerly parts of Africa and the available fossil
material is currently being studied in detail (R. G. Klein, in preparation).

The only other locally occurring hippotragine is H. gigas, which is recorded
from Elandsfontein where it is more commonly represented than H. leucophaeus.
It apparently belongs in the earlier element of the Elandsfontein fauna, while
H. leucophaeus is probably more recent.

The Alcelaphini are among the most diverse and abundant of the African
Bovidae and are represented at all the more important south-western Cape
fossil sites. Surprisingly, the wildebeest (Connochaetes) was not recorded in the
region in historic times, although C. gnou was very common on the interior
plateau and C. taurinus is still one of the most common bovids of the savannas
further north. Damaliscus was also not recorded in the region in recent times,
although D. dorcas still occurs in the southern Cape. The only alcelaphine which
was definitely recorded locally was the southern hartebeest, Alcelaphus buselaphus
caama. This species is also recorded from a number of Holocene occurrences
such as Late Stone Age coastal middens, but is not known from a Pleistocene
context. By contrast, both Connochaetes and Damaliscus are represented at Pleisto-
cene occurrences.

The identification of alcelaphine species on the basis of relative tooth size
is not always reliable. In most instances the local fossils were identified on the
basis of horn cores and other skull characters, but the Swartklip Connochaetes
and Sea Harvest Alcelaphus are represented only by teeth and their relationships
are, therefore, not certain. Nevertheless, the present indications are that
Alcelaphus replaced Connochaetes in the’ south-western Cape during the Holocene.
Such replacements and the apparent local extinction of species such as
Ceratotherium simum may well be related to environmental factors. It has already
been suggested elsewhere that climatic conditions were more favourable in the
south-western Cape during the late Pleistocene.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 51

A number of now extinct alcelaphines are recorded locally. The spectacu-
larly large Megalotragus is represented in both the Elandsfontein and Melkbos
assemblages. Like Pelorovis, this giant bovid apparently became extinct in
South Africa at the end of the Pleistocene (Klein 1972).

At least three other alcelaphines are known from Elandsfontein. One is
tentatively referred to Beatragus, a genus now represented only in East Africa
(B. hunteri), although its affinities may lie with the tsessebe, Damaliscus lunatus.
The apparently recently extinct D. niro is represented by only a few specimens,
which may belong in the later element of the Elandsfontein fauna. One of the
most commonly represented bovids at the site is a form of ‘Rabaticeras’, a genus
which is otherwise known from Swartkrans (Vrba 1971), Olduvai Gorge
(A. W. Gentry, pers. comm.) and Morocco (Ennouchi 1953).

Two medium-sized and as yet unnamed alcelaphines are represented at
Langebaanweg. Both are primitive forms and at a stage of development not
inconsistent with the inferred Pliocene age of the Langebaanweg deposits
(A. W. Gentry, pers. comm.). One may be ancestral to ‘Rabaticeras’, which in
turn may be ancestral to Alcelaphus.

The Cephalophini, a group which is widespread in Africa, are not known
as fossils in the south-western Cape, although the grey duiker, Syluicapra
grimmia, occurs in small numbers in the region today. It may be a relatively
recent immigrant to the area.

By contrast, neotragines are common as fossils and are the most commonly
occurring indigenous antelopes in the region today. Two of the three extant
species of Raphicerus are represented locally, namely, the steenbok (R. campestris)
and the grysbok (R. melanotis). Fossil specimens of these two species are usually
difficult to distinguish from one another and no attempt has been made to
identify the local fossil Raphicerus at the species level. There is clearly more than
one variety represented in the Quaternary assemblages.

Two neotragines are represented at Langebaanweg, one from ‘E’ Quarry
and the other from Baard’s Quarry. Both are now tentatively referred to
Raphicerus. The Baard’s Quarry form is poorly known, but it apparently differs
from the ‘E’ Quarry species, which in turn is clearly not conspecific with any
extant species.

Another neotragine recorded locally is the klipspringer, Oreotragus oreo-
tragus. It was formerly common on mountains in and adjacent to the south-
western Cape and is known as a fossil only from Sea Harvest, where it is repre-
sented by a single metacarpal.

The only antilopine recorded from South Africa in recent times is the
springbok, Antidorcas marsupialis. During the Pliocene and Pleistocene, however,
several species of Gazella, a genus which still survives elsewhere in Africa and
in Asia, were represented in the country. There are two extinct species of
Gazella recorded from the south-western Cape, one from Langebaanweg and
the other from Elandsfontein.

Although the modern springbok was not recorded in the south-western

52 ANNALS OF THE SOUTH AFRICAN MUSEUM

Cape in historic times, Antidorcas is not uncommon as a fossil and at least two
species are represented. The subspecies A. marsupialis australis (Hendey &
Hendey 1968) is now regarded as a species distinct from A. marsupialis and it
is also clearly distinct from the extinct species A. reckt, which is locally recorded
from Elandsfontein. The southern springbok is best known from Swartklip,
but is also represented in the Elandsfontein and Melkbos assemblages and may
also be present at Sea Harvest.

The horn cores of A. recki are more like those of A. marsupialis than are
those of A. australis, which suggests that A. marsupialis could be a descendant of
A. recki and that A. australis belongs on a distinct lineage. The latter species is
so far known only from the south-western Cape, although fossil specimens
from the southern Cape may also belong to this species. Like Aippotragus
leucophaeus, it is regarded as a species which was endemic to the southern parts
of the African continent.

A. recki is less well represented at Elandsfontein than A. australis and, if the
former is indeed ancestral to A. marsupialis, its normal habitat may have been
the inland plateau and its presence at Elandsfontein the result of penetration
into this area of occasional groups of this plains species. Movements of the
modern springbok into the Cape Folded Mountains and west coast region were
recorded in historic times and it is not impossible that they also moved into the
northern parts of the south-western Cape, that is, the area in which Elands-
fontein is situated. A. australis is thus regarded as the locally enedmic springbok
which occasionally may have met its plains counterpart near the northerly
limits of its range.

The possibility that the Elandsfontein A. reckt belongs to the early element
of the fauna and was ancestral to A. australis, which would therefore belong in
the later fauna, was considered but rejected. In all those instances where it is
certain that early and late forms exist as, for example, amongst the Carnivora
(vide infra), the former are more abundantly represented. Since A. australis
is more common than A. recki, this would mean a reversal of the usual pattern
in the case of the springboks. Furthermore, the possibility that certain specimens
were not contemporaneous with others of the same or related species, was often
suspected because of the nature of their preservation. The Elandsfontein
A. australis assemblage includes at least one specimen which differs in preser-
vation from the others, which suggests the possibility that early and late forms
of the species are represented at the site.

The somewhat aberrant antelope, Pelea capreolus, which is endemic to
southern Africa, still occurs in the south-western Cape today. Since it is a
species which is largely confined to more mountainous areas, it would not be
expected to occur in any of the more important local fossil occurrences. It is in
fact extremely rare and is known only from a few fragmentary specimens from
Elandsfontein and Sea Harvest.

In addition to those species already mentioned, there are two others from
Elandsfontein and one from Langebaanweg which are unclassified. All three

LATE CENOZOICG CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE ey

are moderately large forms and all are clearly not conspecific with any extant
species.

Apart from the Carnivora, the only other mammals which are recorded
locally as fossils are Cetacea. Holocene cetacean remains are not uncommon
in hominid occupation sites and other deposits adjacent to the present coast
and have also been recovered during building operations on Cape Town’s
reclaimed foreshore area. Heavily mineralized cetacean remains are frequently
washed ashore on the beach at Milnerton near Cape Town, in association with
other marine fossils and the remains of terrestrial mammals. The latter have
included the gomphothere tooth fragment referred to earlier. Most of this
material is in private collections and is unstudied. The cetacean remains from
Langebaanweg (Hendey 1970a: 103), all of which are from Bed 1 of the
Varswater Formation, are also unstudied.

The only extant orders of African mammals which are not represented in
the fossil record of the south-western Cape are the Chiroptera and Sirenia.
The former are well represented in the modern fauna of the region and although
they may occur as fossils in caves, no such occurrence was investigated in the
course of the present study. No sirenians have ever been recorded near the
southerly parts of Africa in recent times and their absence from the local fossil
record is, therefore, not surprising.

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

NON-MAMMALIAN FOSSILS FROM THE SOUTH-WESTERN CAPE PROVINCE

The invertebrate and non-mammalian vertebrate fossils from the south-
western Cape form a significant part of the local fossil record. Although such
fossils are recovered and studied only incidentally to current investigations on
local fossil mammals, they have sometimes proved useful in interpreting the
origins of the assemblages in which they occur, as well as giving indications of
the nature of past environments. For example, the occurrence of marine
invertebrates and vertebrates in Bed 1 of the Varswater Formation provided
evidence that the Bed 1 deposits were laid down on a marine shoreline.

Most of the non-mammalian vertebrate fossils from this region have yet
to be studied in detail, but some specimens have already been partly or com-
pletely classified.

Chondrichthyes are known locally in association with mammalian remains
only from Langebaanweg and Milnerton, the latter being a littoral occurrence
of little significance. In the Langebaanweg assemblage at least six species of
shark are provisionally identified, as well as a skate, an eagle ray and a sting
ray. The remains are very incomplete and mostly only isolated teeth or tooth-
plates are found.

The only fossil mammal locality from which teleost remains have been
recovered in any numbers is Langebaanweg, although Swartklip and Sea
Harvest have produced a few isolated specimens. At Langebaanweg vertebrae
have been most common, but occasional teeth are recorded from Bed 1 and
catfish fin spines are known from Beds 2 and ga.

Anuran remains are also quite common at Langebaanweg and have
occasionally been found at Quaternary occurrences as well.

The remains of Reptilia are more common and Chelonia are almost
ubiquitous. Skeletal elements of land tortoises occur at all the important south-
western Cape sites and are also known from many of the less significant fossil
occurrences. At Langebaanweg the individuals represented range from very
small to moderately large and vast numbers of specimens are known. No
attempt has ever been made to compute the number of individuals involved,
but the figure must run into thousands. As far as is known, only one or possibly
two species of land tortoises are represented in the Varswater Formation, while
at Baard’s Quarry a land tortoise and a water turtle are recorded.

Reptilia other than Chelonia are far less common, but both lizards and
snakes are known and once again the largest assemblage is from Langebaanweg.
A feature of the south-western Cape fauna, both modern and fossil, is the absence
of crocodilians from the record. Elsewhere in Africa these reptiles are, or were
common and it seems that, as with many mammalian species, they failed to
penetrate to the more southerly parts of the continent.

With the exception of Langebaanweg, birds are not common as fossils in
the south-western Cape, although the ostrich is known from all the more
important occurrences. The Langebaanweg fauna includes a wide variety of
smaller birds, of which only a penguin (Spheniscus predemersus) has been described

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 55

(Simpson 1971). Another penguin, apparently Spheniscus demersus, is known
from Sea Harvest.

Relatively little attention has been paid to invertebrate fossils which occur
in association with vertebrate remains. The shells of land snails are not un-
common at many of the Quaternary occurrences, but more significant are the
records of marine invertebrates in Bed 1 of the Varswater Formation. They
constitute the largest Pliocene invertebrate assemblage yet recorded from
South Africa and have been described in detail elsewhere (Kensley 1972).

56 ANNALS OF THE SOUTH AFRICAN MUSEUM

LATE CENOzoICcC MAMMAL AGES IN SOUTHERN AFRICA

The age of southern African fossil mammal faunas can be indicated in a
number of ways. It is, of course, common to refer them to particular epochs,
which may be subdivided into ‘Lower’, ‘Middle’ and ‘Upper’, or ‘Early’,
‘Middle’ and ‘Late’ stages. The practice thus far employed in the present report
has been to use the latter alternative on an informal basis (e.g. late Pliocene),
since there never has been an acceptable formal definition of subdivisions of the
southern African Cenozoic epochs. References to the ‘late Pliocene’ and others
such as ‘Plio/Pleistocene’ are not entirely satisfactory because the epochs
themselves are poorly defined in southern Africa. European stage names such
as ‘Villafranchian’ have been used as a substitute and with varying degrees of
circumspection by various authors, but since this procedure is so obviously
unsatisfactory, it is usually considered unacceptable.

It has long been the practice in other parts of the world to devise a system
of local names to which are referred faunas dating from a particular period in
time and with a particular character. This system has obvious advantages and
at the Third Pan-African Congress on Prehistory (1955) it was recommended
that the Pleistocene faunas of Africa be divided into four ‘stages’, namely,
Omo-Kanam, Lower Olduvai, Upper Olduvai and Post Olduvai ie
1957: xxx). This nomenclature never came into general use.

Cooke (1967: 179) has discussed in some detail the question of a formal
stratigraphic nomenclature for the late Tertiary and Quaternary of Africa
based on mammalian faunas and he concluded that, ‘it is probably true to say
that our knowledge is not yet adequate for the designation of Land-Mammal
““Ages’’’, His carefully considered justifications for the use of his own provisional
terminology are undoubtedly sound, but his decision to regard the units as
‘faunal spans’ rather than ‘stages’ is questionable. This decision was based on
the definition of a ‘stage’ as set out in the American Code of Stratigrophic
Nomenclature (1961), and since on this basis his units did not rank as ‘stages’
he refrained from using ‘-an’ and ‘-ian’ endings for his ‘stage’ names.

The application of this code to a palaeontological rather than a geological
problem can hardly be justified if it results in a clumsy and possibly confusing
nomenclature. For example, it is now necessary to refer to the fauna from
Makapansgat as a ‘Sterkfontein faunal span fauna’ rather than as a ‘Sterk-
fonteinian fauna’. Any system of nomenclature, no matter how formal, must
function as an aid to communication and no ‘code of nomenclature’ has been,
or should be completely inflexible. While this is not intended as a plea for
scientific anarchy, it is intended to convey the opinion that too rigid a control
in such matters may ultimately lead to the very situation it is designed to avoid.
Terms such as ‘age’ and ‘stage’ do have different connotations as any good
dictionary will show and the former is used here to indicate a period of time
characterized by a particular mammalian fauna, while ‘stage’ is recognized
as a geological time-stratigraphic unit which is not applicable in the present
instance.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 57

The proposed series of age names (Table 7) are a modification of the
current terminology applied to the Quaternary, with an additional two names
to cover the known late Tertiary faunas. The use of these names, or subsequent
modifications, will eliminate the necessity of using others defined and designed
for use elsewhere and in other fields of study. It must be stressed that the
proposed terminology is intended for use in reference to mammalian faunas
only and its adaptation for use in other disciplines will inevitably lead to
confusion.

TABLE 7

Mammal ages of southern Africa

Proposed Provisional terms
names of Cooke (1967) Sites Epochs
RECENT ‘Recent’ Numerous cave and surface sites Holocene
FLORISIAN Florisbad-Vlakkraal | Elandsfontein (in part), Melkbos,
Faunal Span Swartklip, Nelson Bay Cave (in
part), Wonderwerk, Florisbad,
Vlakkraal, Kalkbank and others
io eS ee ees ss
CORNELIAN Vaal-Cornelia Elandsfontein (in part), Vaal River 5
Faunal Span Younger Gravels, Cornelia, Cave of S
Hearths (in part) ‘3
= en ees A
MAKAPANIAN Swartkrans Taung, Makapansgat, Sterkfontein,
Faunal Span Swartkrans, Kromdraai,
?Langebaanweg (Baard’s Quarry)
Sterkfontein
Faunal Span
LANGEBAANIAN Langebaanweg, Kleinzee Pliocene
NAMIBIAN Elisabethfelder, Langentai, Bohrloch | Miocene

A tentative correlation with European and North American ages is
proposed (Table 8) but, to paraphrase Wood et al. (1941), although the supposed
equivalence of the proposed names to the epochs and Northern Hemisphere
ages is indicated, the new terms are emphatically not defined in relation to
them. The precise limits of the ages is intended to be flexible and may be
modified in the light of later discoveries.

The definitions which follow are based on the system employed by Wood
et al. (1941). The faunal lists are by no means exhaustive and final and, bearing
in mind the earlier comment by Cooke on our present inadequate knowledge
of southern African fossil mammals, the definitions form only the foundations to
which more secure and substantial information may be added from time to
time.

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

FLORISIAN

Source of name: Florisbad, Orange Free State.

Includes the faunas from Elandsfontein (in part), Melkbos, Swartklip, Nelson Bay Cave (in

part), Driefontein (Cradock), Wonderwerk, Florisbad, Vlakkraal, Kalkbank, Cave of Hearths

(in part), Chelmer.

Cultural associations: Middle Stone Age and earlier part of Late Stone Age.

First appearance: Many modern species including Homo sapiens sapiens, Papio ursinus, Loxodonta

africana.

Last appearance: Pelorovis, Megalotragus, Damaliscus niro, Gazella.

Comment: The fauna is composed essentially of modern species, many of which are characterized
by individuals of larger size than their modern counterparts.

References: Cooke 1963; Wells 1970; Klein 1972; this report.

CORNELIAN

Source of name: Cornelia, Orange Free State.

Includes the faunas from Elandsfontein (in part), Vaal River Younger Gravels, Cornelia, Cave

of Hearths (in part).

Cultural associations: Advanced Acheulian.

First appearance: Homo sapiens rhodesiensis, Elephas transvaalensis, Loxodonta atlantica, Stylochoerus,

Phacochoerus.

Last appearance: Simopithecus, Megantereon, Hipparion, Mesochoerus, Tapinochoerus, Libytherium.

Comment: The modern element in the fauna is strong, although the forms represented are usually
readily distinguishable from their extant counterparts. Some archaic elements are
still present.

References: Cooke 1963; this report.

MAKAPANIAN

Source of name: Makapansgat (Limeworks), Transvaal.

Includes the faunas from Taung, Makapansgat, Sterkfontein, Swartkrans and Kromdraai.

Cultural associations: Earliest artefacts and Early Acheulian.

First appearance: Australopithecus, Homo, Papio, Panthera, Megantereon, Crocuta, Diceros bicornts,
Ceratotherium simum, Tapinochoerus, Hippopotamus, Connochaetes, Antidorcas.

Last appearance: Australopithecus, Homo erectus, Parapapio, Hyaenictis, Lycyaena, Dinofelis, chalicothere,
Makapania.

Comment: Archaic forms predominate, but modern genera appear in appreciable numbers.

References: Cooke 1963; this report.

LANGEBAANIAN

Source of name: Langebaanweg, Cape Province.

Includes the faunas from Langebaanweg and Kleinzee.

Cultural associations: None.

Characteristic fossils: Prionodelphis, Agriotherium, Percrocuta, Machairodus, Enhydriodon, Mammuthus
subplanifrons, Nyanzachoerus, Ceratotherium praecox.

Comment: No extant species and many extinct genera.

References: Stromer 19314, b; this report.

NAMIBIAN

Source of name: Namib Desert, South West Africa.

Includes the faunas from Elisabethfelder, Langental and Bohrloch.

Characteristic fossils: Protypotheroides, Myohyrax, Pterodon, Diamantohyus, Propalaeoryx, Austrolagomys,
Parapedetes, Bathyergoides, Diamantomys.

Comment: No extant genera.

References: Stromer 1926; Hopwood 1929; Cooke 1968.

LATE CENOZOIG CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

TABLE 8
Tentative correlation of mammal ages of southern Africa, Europe and North
America
SOUTHERN AFRICA EUROPE NORTH AMERICA
ETHIOPIAN PALAEARCTIC NEARCTIC
REGION REGION REGION
aaa oO
RECENT RECENT RECENT
FLORISIAN = RANCHOLABREAN
Z
o
CORNELIAN CROMERIAN 5 IRVINGTONIAN
2
c 2
MAKAPANIAN VILLAFRANCHIAN F BLANCAN
LANGEBAANIAN ASTIAN REXROADIAN
4
PIKERMIAN
z = HEMPHILLIAN
< el
3)
hel ei ee ©
fe) |
= B&B j--------- 8
VALLESIAN
---l------ CLARENDONIAN
-~-------- 12
TORTONIAN
--------- BARSTOVIAN
HELVETIAN
-~-------- 16
Mag, Sate z
NAMIBIAN - HEMINGFORDIAN
e
=
BURDIGALIAN
--------- 20
ARIKAREEAN
ome eee le 2 4
28

Sources: Gabunia & Rubinstein 1968; Kurtén 1971.

Approximate age in millions of years

59

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

The post-Langebaanian ages defined here differ only slightly in arrange-
ment from the most recent of the subdivisions of the South African Pleistocene
(Cooke 1967: Table 1). There is a precedent for this more simply defined three-
fold division since the Makapanian, Cornelian and Florisian correspond to the
‘Ape-Man’, ‘Hand-Axe’ and ‘Middle Stone Age’ faunas of Ewer & Cooke
(1964). The definitions of these ages was based on the simple concept of the
degree of similarity between the fossil faunas and the modern fauna of South
Africa, with the latter being taken to have spanned the whole of the Holocene.

The most recent of the Pleistocene ages, the Florisian, is that in which the
fauna was little different to that of the Holocene. Most of the species represented
are still extant, although the varieties represented often differed in certain
respects (e.g. size) from their extant counterparts. The only extinct genera
recorded (Pelorovis, Megalotragus) are giant forms which are regarded as highly
specialized rather than archaic in character. Their extinction may be linked
with environmental and/or hominid activity factors rather than the rise of
better adapted, ecologically related species.

The next oldest age, the Cornelian, was the one in which the modern
character of the fauna became discernible, but in which the species represented
were usually readily distinguishable from their modern counterparts by more
than just size differences. Furthermore, some truly archaic genera appear for
the last time. These include primates (Szmopithecus), carnivores (Megantereon),
perissodactyls (Hipparion) and artiodactyls (Mesochoerus, Libytherium), which
were superseded by more advanced and better adapted forms. There was also
at least one other important difference between the Cornelian and modern
faunas. During the Cornelian the Suidae were represented by a wider variety
of forms and the present almost total dominance by the Bovidae of the medium-
sized herbivore fauna had yet to be achieved. The Cornelian is characterized
by a fauna which bridges the gap between the essentially modern Florisian
fauna and the first one to have a pronounced archaic character, namely, the
Makapanian.

During the Makapanian there were few extant species represented amongst
the larger mammals. Those that are conspecific with modern forms are either
rather generalized representatives of their kind (e.g. Canis mesomelas, Hyaena
hyaena) or specialized forms adapted to ecological niches in which they had no
active competitors (e.g. Hippopotamus amphibius, Ceratotherium simum). Patterns
in the representation of certain groups also differed from those in the Holocene
fauna. Amongst the larger carnivores the dominance of the sympatric machairo-
donts and hyaenids was evident (see Ewer 1967: 120). The South African record
of Makapanian Suidae is relatively poor, but judging from contemporaneous
East African records this family was at about the peak of its late Cenozoic
radiation in Africa (see Cooke & Maglio 1972: fig. 3). The fauna as a whole
was, however, clearly ‘African’ in character and ‘Eurasian’ elements such as the
ovibovine, Makapania, were exceptional.

The subdivision of the Makapanian into two or more units would mean

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 61

that the whole basis for defining the Pleistocene ages would have to be altered.
It would necessitate more involved and more precisely worded definitions and
would probably serve no useful purpose. It should however be noted that in a
recent review of the Pleistocene ‘faunal spans’ concept, Wells (1969: 94)
suggested that ‘Kromdraai should be dissociated from Swartkrans, and either
included in the Cornelia span or made the type locality of an intermediate
span’. The latter alternative would have resulted in the creation of five rather
than four units. The present arrangement of only three units is an obvious
reversal of at least one trend of thought.

The present definitions of the South African Pleistocene mammal ages
are of some significance to anthropologists, since the local mammals include
both Australopithecus and Homo. The Makapanian is characterized in part by
the presence of both Australopithecus and Homo erectus and is, therefore, the age in
which the genus Homo was differentiated. The only hominid recorded in the
Cornelian is Homo sapiens rhodesiensis, which is included here on the basis of the
Saldanha skull from Elandsfontein. Homo sapiens sapiens is not recorded until
the Florisian. As a result, the ages can be broadly correlated with units of the
local cultural succession.

The Tertiary mammal ages may also be considered in relation to the
modern fauna of southern Africa, but the connections are more remote and
would be largely meaningless for ages earlier than the Langebaanian. The
Namibian and any other Tertiary ages which might be recognized in the future
will have to be defined in relation to one another rather than in relation to the
modern fauna.

The Langebaanian is differentiated from the Makapanian largely by the
fact that no extant species of larger mammals are included. The fauna is
definitely archaic in character and includes many extinct genera. Those genera
which are still extant are conservative forms such as Ceratotherium or highly
specialized and successful forms with long fossil histories such as Felis. In
addition the ‘Eurasian’ element in the fauna is more marked than in the
Makapanian and includes Agriotherium, Percrocuta and a boselaphine. The
machairodont/hyaenid combination is similar to that of the Makapanian,
but the Pantheriinae were evidently not yet present. The Suidae were appar-
ently not yet diversified and the type fauna from Langebaanweg includes only
two species. The Proboscidea recorded are a primitive elephantid and a gom-
phothere.

Although the proposed nomenclature is intended to be used instead of
other systems, it is obviously desirable that the position of the ages relative to
an absolute chronology and to the geological epochs be determined.

In terms of absolute age, the Pleistocene/Holocene boundary is here taken
to be 12 000 B.P. There is evidence which indicates that at about this time
there were significant changes in local environmental conditions which were
associated with changes in the fauna and cultures (see Klein 1972). The South
African Pleistocene/Holocene boundary may thus be defined to be more or

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

less synchronous with that in other parts of the world.

The Plio/Pleistocene boundary is taken to date back 3,5 million years
(see De Heinzelin 1969), although this is an arbitrary figure since there is at
present no satisfactory basis on which this boundary can be defined in South
Africa. A date of 3 million B.P. would be as convenient (see Savage & Curtis
1970), and other alternatives have been suggested so that the situation is
still far from being resolved (see Flint 1971).

Similarly the Mio/Pliocene boundary is arbitrarily dated at 12 million
B.P. In North America and Europe it is variously placed between 10 and 13
million B.P. (see Gabunia & Rubinstein 1968), to as little as 5 million B.P.
(Van Couvering 1972).

Many East African late Cenozoic faunas have been dated by radiometric
determinations on associated igneous rocks and this enables them to be related
to the absolute chronology outlined above. Absolute age determinations in
South Africa have only been possible for the late late Pleistocene and Holocene,
so it is only by fixing the age of the local pre-Holocene faunas relative to those
of East Africa that the mammal ages may be tentatively.related to the absolute
chronology.

The fauna of the Namibian is poorly known and comparisons with faunas.
in East Africa have been inconclusive (Cooke 1968). It is, however, fairly
certain that it does predate the 14 million-year-old Fort Ternan fauna and is
probably broadly contemporaneous with those from sites such as Rusinga,
Napak, Songhor and others. The Namibian is for the present regarded simply
as a Miocene mammal age with undefined limits.

The Langebaanian type fauna has an inferred age of about 4,5 million
years, so that this age is definitely in the Pliocene and clearly dates from the
latter part of the epoch. On this basis it is concluded to be broadly equivalent
to the Astian of Europe, the Dhok Pathan of India and the early Blancan
(Rexroadian) of North America. The upper and lower limits of this age cannot
be defined, but for the sake of convenience the former is taken to coincide with
the Plio/Pleistocene boundary (i.e. 3,5 million B.P.).

The present indications are that the Namibian and Langebaanian may be
separated by a time interval of 10 million years or more. This interval is without
a significant recorded fossil history, although Namaqualand in the north-
western Cape Province is an area with some potential in this respect. A
previously unrecorded fossil mammal locality, Bosluispan near Gamoep, has
yielded a limited and poorly preserved fauna in which the only identified
specimen is a tooth of Protanancus macinnesi Arambourg, 1946. (Information
supplied by the South African Geological Survey.) Protanancus is recorded from
the ‘Upper Miocene’ of East Africa (Leakey 1967), and it thus seems likely
that the Bosluispan fauna is intermediate in age between the Namibian and
Langebaanian. Another fossil from Namaqualand which is apparently Pliocene
in age and which apparently pre-dates the Langebaanian type fauna is the
holotype of Notohipparion namaquense (Haughton 1932)). This specimen is from

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 63

Areb, which is about 80 km northwest of Bosluispan. On the basis of these two
records it seems possible that a post-Namibian/pre-Langebaanian fauna, or
faunas, may one day be recorded from Namaqualand.

The absolute ages of the upper and lower limits of the three Pleistocene
mammal ages are also not certain, although the lower limit of the Makapanian
and upper limit of the Florisian are taken to coincide respectively with the
Plio/Pleistocene and Pleistocene/Holocene boundaries. The date of the
Makapanian/Cornelian boundary is unknown. The faunas from Cornelia
and Elandsfontein have been correlated with that of Olduvai Bed IV (see
Cooke 1963), and recently the base of Bed IV was dated to about 0,7 million
B.P. (Leakey 1971). The Makapanian/Cornelian boundary may therefore be
somewhere between 0,5 and 1 million B.P. The Cornelian/Florisian boundary
is also not fixed, but it may date back to something of the order of 100 000
years.

Although the dating of the mammal ages is so imprecise it is perfectly
evident that those of the Pleistocene differ very considerably in the lengths of
time which they cover. Even in the arrangement of ‘faunal spans’ discussed by
Cooke (1967) the earlier units were clearly far longer than the later ones.
This problem is by no means unique to the southern African systems and it
does not necessarily detract from the value of defined mammal ages.

In a discussion on the geochronology of North America based on fossil
mammals, Tedford (1970) pointed out the value, and shortcomings, of the
system devised by Wood et al. (1941). There can be little doubt that were a
comparable arrangement to come into general use in Africa, communications
would be considerably facilitated. The present attempted emulation is certainly
less satisfactory than the North American model and it would have been
preferable for the nomenclature to have been based on the superior East
African fossil record. Recently Coppens (1972) did indeed propose a subdivision
of the African Pliocene and Pleistocene based on this record, although in this
instance the two epochs were divided into ‘Zones’ (I to VIII in descending
order of age). Coppens’s nomenclatural system and that proposed here both
have shortcomings and it is quite possible that neither will prove generally
acceptable. However, it seems inevitable that agreement on a system for Africa
will eventually be reached and although the American system is itself not
faultless it does perhaps provide the best available model.

64 ANNALS OF THE SOUTH AFRICAN MUSEUM

CARNIVORA OF THE PLIOCENE

The Carnivora from the late Pliocene deposits at Langebaanweg are
treated as a unit separate from the Quaternary species since the two groups have
little in common.

The study of the Langebaanweg material revealed that no extant species
are represented, an entirely predictable conclusion in view of the age of the
fossils. In the systematic accounts which follow, some of the fossils are compared
with corresponding parts of modern species and it was intended that such
comparisons should serve largely to clarify the descriptions of the fossils. The
comparisons were not made in such a way as to prove that the fossils are
taxonomically distinct from the modern species. In other words, no attempt
was made to fully catalogue the differences between any two species dealt with
in this fashion. The intention was rather that characters such as size of fossil
species or of individual skeletal components might be referred to in terms of
material which is more widely available and familiar.

In the case of the Quaternary fossils, references to modern comparative
material had a greater significance, since in these instances the relationships
between the modern and fossil species are far closer. Consequently, in order
that the taxonomic status of the latter might be determined, the similarities
and differences between them and the moderns had to be clearly established.
The fact that inadequate modern osteological series limited the value of studies
on certain of the Quaternary species did not alter the basic difference in
approach to the treatment of the Pliocene and post-Pliocene fossil material.

The lack of direct access to relevant fossil specimens from outside the
south-western Cape has undoubtedly had an adverse affect on the present study
of the Langebaanweg Carnivora. However, it is hoped that by placing on record
the material which is now available, future investigations on contemporaneous
or related fossil species, especially those from East Africa, will be facilitated.

In all the systematic accounts which follow, the tooth cusp terminology
illustrated in Figure 5 has been used. All original measurements under 130 mm
were taken with the same pair of calipers and were corrected to one-tenth of a
millimetre. Measurements over 130 mm were read off ortho-projections onto
a plane surface using a ruler. The latter measurements are obviously less
accurate and were corrected to the nearest millimetre.

All tooth measurements were taken across the longest and broadest points
at, or near to the base of the crown. In the case of the canines, the dimensions
are those taken at the base of the crown in line with the alveolar margin.

Unless otherwise stated, all specimen numbers are those in accession
registers of the South African Museum, Cape Town. Modern specimens are
distinguished from the fossils by the prefix ‘SAM’. Specimens from collections
in other institutions are distinguished as follows:

AMNH — American Museum of Natural History, New York.
BMNH —British Museum (Natural History), London.

GSI — Geological Survey of India.

NHMW — Natural History Museum, Vienna.

65

fx]
zZ
5 piuoBiiy
fe)
my
Ay
fa)
Ay
<
fe)
Z
4
fx)
EX
:
"
Hi
)
o)
i?)
hy
©
: ajAyseieg
o auode1ey
4
<
S
9 3U090}014
O
N 9u0de)a
fe) mn
vA
is ajAyseyaw
fx)
=
<
4
SHYV1OW

"SIIOATUIBS JO YI99} BY} 07 patjdde Asojouruay *S “B14

piluodeled
pluoze,aw
piuoso0jolg

piuojey

yo}ou

ajAise.aw

jeisseuieg

ajAjseiedo}94

ayAyseieg
auoseieg

dsnd AsOssaooe 101J8}UYy

dsn3a jedisutig

dsna Asossa32e
101138}S0q

TWNONIT
H1jdl
u¥iMO1T
1WIING
SYV1IOWIud
dsno Asossaaoe J01/3}UYy
dsno jedisuiig
dsnd jeusajuy
dsnd Ajsossaa9e
alatidk, 1WN9NIT
HiLji1
Ydddn
Ww9IINd

SUVIOWAYd
YOIMIINY

66 ANNALS OF THE SOUTH AFRICAN MUSEUM

During the late Pliocene, Langebaanweg was so situated in relation to
marine, fluviatile and terrestrial environments that Carnivora from all three
habitats were preserved as fossils. In this respect Langebaanweg is unique
amongst the important late Cenozoic fossil sites of sub-Saharan Africa. The
assemblage probably represents a very good cross-section of those carnivore
species which inhabited the most southerly parts of Africa during the late
Pliocene. More carnivore species are represented at Langebaanweg than in
any other single occurrence in the south-western Cape.

Apart from the importance of this material as additional and new records
in the African late Cenozoic fossil record, the carnivores evidently played a role
in the addition of animal remains to the accumulating deposits. They are,
therefore, an order of particular significance in the Langebaanweg assemblage
as a whole.

Suborder PINNIPEDIA
Family Phocidae
Subfamily Monachinae
Prionodelphis capensis Hendey & Repenning, 1972

Discussion

The Langebaanweg pinniped, Prionodelphis capensis, has already been
described in detail elsewhere (Hendey & Repenning 1972), and little additional
material has since been recovered, none of which alters the conclusions already
reached. The holotype and all the referred cranial remains are from Bed ga
and so is most of the postcranial material. Some postcranial bones are also
known from Bed 2, while a single incomplete femur was recently recovered
from the marine deposits of Bed 1.

This species has shed some additional light on the origins of the Antarctic
monachines, a group which includes species that are still occasionally recorded
as vagrants on the south-western Cape coast (vide infra), and it is a significant
addition to the fossil record of the Monachinae in general (Hendey 19722).

Recently P. capensis was tentatively identified at a second locality in
South Africa. A canine (Q 1799), which is virtually indistinguishable from a
Langebaanweg specimen (L 13152—see Hendey & Repenning 1972: pl. oF),
was found at Koingnaas near Hondeklip Bay on the Cape west coast about
250 km north of Langebaanweg. This specimen, together with a Ceratotherrum
praecox tooth found in the same area (Hooijer 1972), and the assemblage from
Kleinzee a little further north (Stromer 19314, 19310), indicates that while the
Langebaanweg occurrences are the most prolific of their kind yet recorded in
the Cape Province, there are, or were other similar occurrences on the Cape
west coast. All these occurrences are at, or near to, river mouths (Langebaanweg

—Great Berg River; Koingnaas—Swartlintjies River; Kleinzee—Buffels
River).

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 67

Suborder FIssIPEDIA
Family Canidae
Incertae sedis

Comment

Reference has already been made to two canid species in the Langebaan-
weg assemblage (Hendey 1970a), but this material has since been referred to
the Viverridae (vide infra). There are, however, several recently discovered
teeth which apparently do belong to a canid.

Material

L 15184B —Left P* and M,, and right M?.
L 15588B/C—Right M1.

155268 ” Left P*.

L 16120B_ ? Right M3.

Locality and horizon

These specimens are from Bed ga, ‘E’ Quarry, Langebaanweg.

Description

With the exception of the ? Ms, all the teeth are fairly worn and they may
belong to a single individual. The ? M, still has an open root and is unworn.
It definitely belongs to a second individual and may even belong to a different
species. It is only provisionally grouped with the other specimens.

The worn teeth belong to a canid which was apparently comparable in
size to modern Vulpes vulpes. The ? P? is a little larger than the P? of the two
available V. vulpes specimens (Table g), but it is otherwise similar. The P*
differs from the comparative specimens in having a more prominent and more
anteriorly situated protocone, while the anterior keel of the paracone is more
lingually directed. ‘The M! and M? of the two species are similar in morphology
but in the fossil the lingual lobes have a greater transverse development and are
relatively narrower antero-posteriorly.

TABLE 9

Dimensions of the canid teeth from Langebaanweg, compared with those of modern Vulpes vulpes.

SE UC | — — | | |§ |S

Fossil

specimens . eids0 7:2 1 9,8 -23;7 4,0

V. vulpes 13.5% G,7°| 10,1 | 12,5 3.4
r3.0. 6,5 | 10,2) —.12;1 3,0 3,0

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

The lower carnassials of the two species are essentially similar to one
another, but in the fossil specimen the trigonid is broader relative to the talonid
and the metaconid is a little larger. |

The problematical ? M, is a small, single-rooted tooth with a crown which
is almost circular in occlusal view, while the occlusal surface is basin-shaped.
It is far larger than the M, of the two V. vulpes specimens.

Discussion

The identification of this material is based principally on the fact that the
M?! and M2? have protocones, which distinguishes them from the M! and/or
M2? of hyaenids and large viverrid which occur in the same deposits.

It is not clear from the material available to which canid genus the material
belongs and comparisons with previously recorded fossil species were incon-
clusive.

Family Ursidae
Subfamily Agriotheriinae
Agriotherium africanum Hendey, 1972

Discussion

The Langebaanweg ursid has already been described and discussed else-
where (Hendey 19720), and the only additional specimen now known is another
incomplete ulna. It is essentially similar to the one previously described.

Family Mustelidae
Subfamily Mellivorinae
Mellivora aff. punjabiensis Lydekker, 1884
(Fig. 6; also Hendey 1970a: pl. 2E)
Material
L 6385—Left mandibular fragment with P, and Mj.

Locality and horizon

This specimen is from ‘E’ Quarry, Langebaanweg.

Description

This specimen has already been mentioned and figured elsewhere (Hendey
1970a), although no conclusion as to its affinities was reached.

Except for some damage to the condyle, the specimen is complete as far
forward as the anterior end of P,. Both P, and M, are intact and this specimen
belonged to a species or individual which lacked M,. In size it compares with
small individuals of the available modern Mellivora capensis assemblage (n = 5),
this species being the largest of the extant African mustelids, excluding the

re eae

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 69

otters. It is to the Mellivorinae that the fossil specimen evidently belongs,
while the absence of M, suggests its affinities to the genus Mellivora itself. It
does, however, differ from the modern species in certain significant details.

Both P, and M, are narrower than the corresponding teeth in the M.
capensis comparative series (Table 51), and the talonid of M, is far smaller.
The basic features of the modern and fossil teeth are, however, similar. The
fossil P, has small anterior and posterior accessory cusps and around the latter
is an expansion of the cingulum which makes the tooth broader posteriorly.
The structure of the M. capensis P, is similar, although in this species the
accessory cusps are more prominent and the development of the cingulum is
more pronounced, especially postero-internally.

Fig. 6. Occlusal views of the Mellivora aff. punjabiensis mandible
(L 6385) from Langebaanweg (left) and a modern Mellivora
capensis specimen.

Although the M, of L 6385 is well worn, its characteristics are still evident.
Except for being far larger, the tooth is not dissimilar to the M, of Poecilogale.
The talonid is small and is narrower than the posterior end of the trigonid.
In this respect it differs considerably from the M, of M. capensis, which has a

70 ANNALS OF THE SOUTH AFRICAN MUSEUM

relatively large and broad talonid. The trigonid of the fossil tooth, which lacks
the metaconid, makes up about 75°% of the total length of the tooth.

The inferior margin of the mandibular corpus of L 6385 is very slightly
convex below the cheekteeth, whereas in the available M. capensis specimens
it is slightly concave. Also in the modern species, the inferior margin is stepped
upwards near the angle and, while there is a suggestion of this characteristic
in the fossil, it is far less prominent. An apparently related feature in M.
capensis is the fact that the dorsal surface of the condyle is above the level of the
cheekteeth. In the fossil the lower pre-angular step goes with a less elevated
condyle, the dorsal surface of which is level with the remaining cheekteeth.

The mental foramen of L 6385 is situated beneath the anterior end of P,
and there are indications that the symphysis extended this far back as well.

Discussion

The differences between the mandibles of the Langebaanweg Mellivora
and modern M. capensis have important functional implications which are
suggested by the nature of the mandibles in other modern mustelids. In those
species in which the carnassial shearing action is important (e.g. Maries spp.,
Gulo luscus), there is a tendency for little upward inflexion of the pre-angular
inferior margin and for the condyle to be low in relation to the cheekteeth.
On the other hand, in those species in which the posterior cheekteeth function
as crushing agents (e.g. Enhydra lutris, Aonyx capensis), there is a sharp upward
inflexion of the inferior margin towards the angle and the condyle is raised
above the level of the cheekteeth.

The Langebaanweg mandible falls into the former category and M.
capensis into the latter, although M. capensis does not show the extreme develop-
ment of these characteristics which are evident in the highly specialized Enhydra
and Aonyx. Similarly, the Langebaanweg Mellivora is not as extreme in the other
direction. It may thus be regarded as showing the beginnings of a trend towards
modification of the masticatory apparatus which culminated in the condition
evident in modern M. capensis.

The similarities between the lower carnassials of L 6385 and Poecilogale
might therefore be coincidental, the latter having retained the more primitive
shearing dentition. Poecilogale is actually more progressive than most other
mustelids in having lost both P, and P,, and it is often without M, as well. It
is possible that the Langebaanweg Mellivora and Poecilogale had a common
ancestor some time fairly late in the Tertiary, although the former can itself
probably be excluded as a possible ancestor of Poecilogale. Even if M, was
sometimes present in the Langebaanweg species, just as it is sometimes present
in Poecilogale, it is perhaps unlikely that a variable feature of this kind would
have persisted for so long a period in time.

It is even less likely that there is any close relationship between the
Langebaanweg Mellivora and the only other non-lutrine mustelid from southern
Africa, Ictonyx striatus, since in this species M, is always present and its M,

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE at

has a metaconid.

Having suggested a relationship between L 6385 and M. capensis, it is
necessary to examine its possible affinities with extinct mellivorines such as
Eomellivora Zdansky 1924, Promellivora Pilgrim 1932 and the early species of
Melliwvora itself.

Eomellivora can be excluded from consideration since one of the character-
istics of this genus is the presence of My. The recorded species are also far larger
than that from Langebaanweg.

Comparisons with Promellivora are not as simple owing to the fragmentary
nature of the material referred to this genus. It is in fact known only from a
single mandibular fragment from the Dhok Pathan stage of the Siwaliks
(Promellivora punjabiensis Lydekker). Pilgrim (1932: 65) stated that this genus
is distinguished from Mellivora “by the presence of P,, the shallower and more
gradual symphysis, the length and compression of M, and the shorter and
simpler premolars especially P; and P,, and the larger canine’.

Lydekker (1884) had previously referred the Dhok Pathan mandible to
Mellivora and Pilgrim’s justification for erecting a new generic name was based
on the principal of horizontal rather than vertical classification, although he
recognized that ‘it is not improbable that (Promellivora punjabiensis, Mellivora
swvalensis and modern Mellivora) are on the same line of descent’ (Pilgrim 1932:
66). Although it is inevitable that there will be differences of opinion in nomen-
clatural problems of this kind and that generic distinctions made within a single
lineage are often arbitrary, it might be preferable, in view of the fragmentary
nature of the P. punjabiensis holotype, to revert to the original generic designa-
tion rather than to accept a new name. The name ‘Promellivora’ is here rejected
and the Dhok Pathan mellivorine is once again identified as Mellivora punjab-
1ensis.

Comparisons between the Langebaanweg Mellivora and both M. punjab-
zensis and the Pinjor M. sivalensis are inconslusive since the bases for comparison
are limited. The three forms are comparable in size and, not surprisingly, each
has certain characteristics which are more primitive than those of the extant
M. capensis.

One of the characteristics of M. punjabiensis which is apparently shared by
the Langebaanweg form is the long symphysis terminating below the anterior
end of P,. If the symphysis of L 6385 did indeed extend this far back, then this
specimen may also have had the compressed anterior premolars characteristic
of M. punjabiensis. Both forms have a slender M, and Pilgrim (1932: 66) stated
that the M, talonid of M. punjabiensis ‘could not have been expanded’. In
addition, both have mental] foramina below P,. Other important characteristics
such as the presence of P, and absence of M, are evident in only one specimen
or the other. Although it is not possible to tell whether or not the two are
conspecific, they do share certain characteristics and there are some grounds
for believing that they are at the same stage in the evolutionary development of
Mellivora.

72 ANNALS OF THE SOUTH AFRICAN MUSEUM

Even less satisfactory are comparisons between the Langebaanweg form
and M. sivalensis, since published descriptions of the latter deal mainly with
skull characters (e.g. Matthew 1929). However, since M. sivalensis is younger
and therefore probably more advanced than M. punjabiensis, the indications
are that the relationship between the latter and L 6385 is closer than that
between MM. sivalensis and the Langebaanweg specimen. This opinion is reflected
in the tentative identification of the Langebaanweg specimen.

As with several other carnivore species from Langebaanweg, the specific
identity of the Mellivora is likely to be resolved only if additional specimens are
recovered. The single available specimen is nevertheless important, since its
observable characteristics are not inconsistent with a Mellivora of late Pliocene
age and it is in character with the fauna as a whole.

Family Mustelidae
Subfamily Lutrinae
Enhydriodon africanus Stromer, 1931

(Fig 7)

Material
L 9138—Right mandibular fragment with part of P,.

Locality and horizon

This specimen is from ‘E’ Quarry, Langebaanweg.

Description

The single positively identified lutrine fossil specimen from Langebaanweg,
although not described in detail, has already been referred to the species
Enhydriodon africanus (Hendey 1970a: 100). The Langebaanweg specimen is
less complete than the E. africanus holotype, which is from Kleinzee in Namaqua-
land (Stromer 19314), but, since both are right mandibular fragments, com-
parisons are facilitated.

The two specimens are similar in size (Table 10), although the mandibular
corpus of that from Langebaanweg is slightly more robust. In this specimen
only the posterior part of P, is preserved, but the alveoli and some of the roots
of the other cheekteeth are present. Both the Langebaanweg and Kleinzee
specimens have a small and single-rooted P,, but in the former the a -eoli
margins are lost so the relative sizes of the two teeth cannot be determined.
Both specimens have a double-rooted P, and these were apparently similar in
SIZe.

The P, of L 9138 is a little larger than that of the holotype, the size dif-
ference being in keeping with that of the mandibular corpora. These differences
are, however, no greater than might be expected in a single species. As far as

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 73

TABLE I0

Dimensions of the teeth and mandible of the Langebaanweg
Enhydriodon, compared with that of the £. africanus holotype.

Enhydriodon

africanus* Lg138

Height of mandible below M, 27 30,1
Breadth of mandible below M, 14,8 17.7
Fo Wie I Sy | €. 55,0
le eee | 28 C. 2555

P, — ] 6.'5;0 —

b Ee a

P, — ] 6 —

b — =
P, — ] 11,9 6. 13,0
b 8,6 10,0
M,— ] 22,0 c. 22,0
b + 12,0 c. 12,5

* Stromer 19314.

can be judged, the P, morphology in the two specimens is similar. Both have a
prominent cingulum and prominent posterior accessory cusp, although in
L 9138 the accessory cusp is more widely separated from the principal cusp.

The lower molars of the two specimens were apparently similar in size.

The tooth-row lengths are also comparable, although the premolars of
the Langebaanweg specimen are more crowded than those of the Kleinzee
mandible.

The Langebaanweg mandible has four mental foramina, the largest of
which is situated high up on the corpus below the posterior root of P;. Only
one mental foramen is visible in the illustration of the Kleinzee specimen
(Stromer 19314: fig. 1) and this is situated in the same position as the principal
foramen of the Langebaanweg mandible.

Both specimens have the most posterior and inferior part of the symphysis
projecting below the adjacent inferior margin of the corpus.

Discussion

Although there are minor differences between the Kleinzee and Lange-
baanweg Enhydriodon mandibles, there can be little doubt that the two specimens
belong to the same species.

74 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 7. Occlusal and buccal views of the Enhydriodon africanus mandible
(L 9138) from Langebaanweg.

E. africanus is still regarded as a valid taxon, even though comparisons
between it and previously described Eurasian species (see Pilgrim 1932: 82)
are limited by the nature of the available material. Undescribed, but apparently
specifically distinct material is now known from Omo in Ethiopia (F. C. Howell,
pers. comm.), while the genus is also known from North America (Repenning
1967). The relationships between these various forms has yet to be determined.

The widely dispersed records indicate that this was a very successful genus in
the late Tertiary.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE ay

Family Viverridae
Subfamily Viverrinae

Viverra leakeyt Petter, 1963
(Figs 8, 9)

Comment

The material referred to this species may include two temporal variants
of a single species and it is listed according to the stratigraphic unit from which
it is derived.

Material from Bed 2, ‘E” Quarry, Langebaanweg

L 12283—Isolated teeth comprising the dentition of a single individual and
including:
Ikight.C, P*, part of P*, and P* to M?; left Gand P? to M?.
Right C and P, to M,; left C, P, and M,.

L 13097—Parts of the skull and skeleton of a single individual and including:
Posterior part of the braincase and right maxillary fragment with M?.
Distal end of left humerus and proximal end of left ulna.

L 20253—Left maxillary fragment with P* and Ml.

The following are the remains of juveniles:

L 14459—Parts of a skull, including:
Left premaxilla and maxillary fragment with dc, dp? and dp*, with
P! just erupting; right maxillary fragment with dp’.
Right mandibular fragment with dc, and dp, to dp,, with P, just
erupting; left mandibular fragment with P, and dpy.

L 14460—Parts of a skull, including:
Right mandibular fragment with dp, and dp,; left mandibular
fragment with dp, to dp.

Material from Bed 3a, ‘E” Quarry, Langebaanweg

The following specimens were recovered from the excavations LBW 1969/1
and 1970/1. The first six teeth listed apparently represent part of the dentition
of one individual, while the last two may represent other individuals.

L 16240A—Left C.

1516204 —Left P*.

L 16240C—Right M1.

L 15174. —Right M?.

L 16051 & L 16055A—Left M, and part of right M,.
L 16197 —Right P?.

L 16055J —? Right P,.

Material from ‘E” Quarry, but of unknown provenance

L 1700—Left mandibular fragment with M, and part of P,.
L 2672—Right mandibular fragment with part of C and P, to P,.

76 ANNALS OF THE SOUTH AFRICAN MUSEUM

Judging from their preservation, it is likely that L 1700 is from Bed 2 and
L 2672 from Bed ga.

Description

These specimens belong to a viverrid of large size (Table 11) and although
it is best represented in Bed 2, the Bed 3a material is described first.

The characteristics of the Gand P* to M? of the Viverra leakeyt holotype,
which is from Laetolil in Tanzania (Petter 1963), are precisely those evident
in the corresponding teeth of the Bed 3a sample. The P* protocone and support-
ing root are lost in the V. leakey: holotype, but they are preserved in the Bed 3a
specimens. Both the protocone and the root are large, the latter being larger
even than the antero-external root. The long axis of the protocone and root
projects antero-internally from the paracone and there is a marked indentation
between the parastyle and protocone. The parastyle itself is very small and is
situated at the base of the prominent paracone keel. It does not project as far
anteriorly as the protocone. The buccal margins of both the Bed 3a specimens
are more or less straight. :

The lower teeth of V. leakeyi are unrecorded and comparisons with those
from Langebaanweg are therefore not possible. The Bed 3a tooth tentatively
identified as a P, is double-rooted, the crown is antero-posteriorly elongated and
consists only of the principal cusp, the apex of which is directed slightly pos-
teriorly. In lateral view the anterior and posterior keels of the principal cusp are
slightly concave. The M, is a large tooth in which the talonid is much reduced
relative to the trigonid. The paraconid is lower than the protoconid and it lies
lingually relative to the protoconid, which results in the buccal surface of this
portion of the tooth having a markedly convex outline in occlusal view. The
metaconid is prominent, but is much smaller than the protoconid.

The mandible L 2672 belonged to a very aged individual, the remaining
teeth are very worn and the specimen is poorly preserved. At least a part of M,
must have been lost during life, since the alveolus of the anterior root is closed.
The premolars appear to be essentially similar to the ? P, (L 16055J) in
morphology and there is a progressive increase in size from P, to P,. Although
the M, is lost, it must have been about the same size as the Bed 3a M,, L 16051.
The Mg, which is also lost, was relatively small and single-rooted. ‘The mandible
is much larger than that of modern V. civetta, but its proportions are essentially
similar.

In most respects the teeth from Bed 2 are similar to those from Bed ga.
The observed differences are not very marked and, since the assemblages
are so small, their significance, if any, cannot be established at this stage.

The upper canine of L 12863 has pronounced grooves on both lingual
and buccal surfaces and a not very prominent anterior keel. The nature of the
anterior keel and the lingual grooves differentiates it from the Bed ga canine,
and apparently also that of the V. leakey: holotype.

The P! is a small tooth with a rather conical crown and slightly bulbous

77

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

‘S61 wast z
‘EQ61 sa9}19g |

O.g1 ‘9 | ag Pol ors OSI L Vo BOT 9 — — olga] ef pog¢
Pe Se ot | OS ee ae i ee SS (ef ee | ee
(19091'T) ({SSog1'J) pareos] v& pog
Pesce eke ea ae ee urea ere |
6° 6°G gil G“Gr G‘G g‘b1 ‘By ake 7

64:9 — ov 9 sO! 3°9 1‘OI EQRQzI'T & pog

eipunanp
iz Eee sal pet mak C61 = sen es a — — — — SuIINUuysr A
ridayba}
08 0G G1 ror -— oh 6G OSI = — — — oo G‘g DLINA
6g L‘G OP aldoy Wee ARNG oy 1A — = aa — — _ o‘L 6g ‘9 4199}
(bi 1S 17) (QobzgrT) (bosgt'T) (yorzgr'T) pare[os] v& pog
— —_ oGr sor oor Sor — — — = — — — — €Szoz'y]
Oro oy Dy i ie Gine 8°60 -— —— — = -- — — — — — Lo6é1'T % pog
aL, EP Vai Lake, nite eran L°G oS bb Por gf g‘P o‘L £6 CQQzI'T
q I q I sl I q I q I q I I ae |
aN IW rd ed od td 0)

“QBULIIDATA IOYIO JO BSOY} YIIM paredusod ‘2ayva) visaaig BOMULLqISUL'] IY} JO YII9} JO sUOISUDWHICT

Il ATaV],

78 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 8. Occlusal views of the left P? to M? (A) and right M, and M, (B) of the
Viverra leakepi from Langebaanweg.

root. Ihe P? and P? are similar in structure to the ? P, from Bedi3a these
is much smaller than the P?.

The P* of L 12863 differs from that of the Bed 3a form in certain respects.
It lacks a parastyle, although there is a slight shelving where the parastyle
would have been if present. The paracone lacks an anterior keel and the tooth is
broader in the region of this cusp than in the Bed ga specimens. The protocone
is similar in size and situation to those of the Bed 3a specimens, but the indenta-
tion between the protocone and buccal cusps is not as sharply notched. The
buccal margin of the L 12863 P* is markedly concave, rather than straight as
in L 16197 and L 16224. The P* of L 20253 is similar to that of L 12863, except
that it does have a small parastyle, in which respect it resembles the Bed ga
specimen, L 16224.

The M? is known from three specimens from Bed 2 and they are essentially
similar to L 16240C from Bed ga. There are, however, quite marked differences
in the M? from the two beds. This tooth is lost in L 13097 and L 20253, but its
alveolus is present in both specimens. These accommodate the M? of L 12863
quite well, so that at least in respect of size the M? of the three Bed 2 individuals
are similar. The only known Bed 3a M?is appreciably larger and the individual
features of the crown are more prominently developed than in the M? of
L 12863. In L 13097 and L 20253 the buccal margins of M1 and M? are at an
angle of about 115° to the long axis of P* and their lingual limits form a line
parallel to this axis.

The lower canine of L 12863 is similar in size to the upper canine, but
lacks grooves on its lingual surface. Of the lower premolars of this individual,
only P, and P, are reasonably intact and they are similar in morphology to P?
and P3. The P, of L 1700 does, however, differ from these teeth in that it has a

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 79

prominent posterior accessory cusp, which is flanked posteriorly by a well-
developed cingulum. This is the only premolar amongst all those which are
referred to this species in which more than just the principal cusp is present.

On the other hand, the M, of L 1700 is virtually ideutical to that of
L 12863. These teeth are in turn very similar to the M, of the Bed ga form,
except that their trigonids are less enlarged relative to the talonids. The over-
all size of the Bed 3a specimens is also greater and they have a more pronounced
cingulum at the anterior end.

The M, of L_ 12863 is a small, single-rooted tooth, which is
slightly elongated antero-posteriorly. There are two small cusps situated at the
anterior and posterior ends of the buccal side of the tooth, while on the lingual
side there are two cusps situated one behind the other at the anterior end. ‘This
tooth is accommodated quite well by the M, alveolus of L 1700. The M, of
the Bed 3a form is not known, but judging from the size of the M, alveolus of
L 2672, it was similar in size to that of L 12863.

The premolars of both Bed 2 and Bed ga forms have markedly divergent
anterior and posterior roots, which indicates that these teeth were well spaced
in both mandible and maxilla. This is evident in both the mandibular fragments
which are known (L 1700 & L 2672), even though the former is much more
slender, comparing in size with that of modern Canis mesomelas.

The fragmentary braincase of L 13097 (Table 12) is a significant specimen,
since, although it is so incomplete, it does exhibit some remarkable charac-
teristics. ‘The posterior constriction of the braincase is far more pronounced
than in any modern viverrid species, while the posterior part of the sagittal
crest is exceptionally high. The transverse compression of the posterior part
of the skull is carried to such an extreme that the width across the occipital
condyles is actually greater than the width of the occiput immediately above

I

I |

Fig. 9. Occipital view of the Viverra leakeyi
braincase (L 13097) from Langebaanweg.

I

£

I

I

§

UII

80 ANNALS OF THE SOUTH AFRICAN MUSEUM

them (Fig. 9). In all extant viverrid species the reverse is true. In L 13097
the ratio of skull height (top of sagittal crest to inferior margin of condyles)
to skull width (transverse diameter of occipital immediately above dorsal
margin of foramen magnum) is 1,98:1, whereas in modern V. civetia (n = 1)
it is 1,11:1 and in Genetta spp. (n = 11) it 1s 1,09:1.

TABLE 12

Dimensions of the skull, humerus and ulna of the Lange-
baanweg Vzverra leakeyz.

Width across condyles .

Skull height from base of songhiles = “ap a
sagittal crest . :

Transverse diameter of ue Sueredicesl

above foramen magnum

58,0

Greatest transverse diameter of distal arti-

ticular end of humerus 26,1
Greatest antero-posterior iiemiwies of sine
shaft 18,6

The skull of this species must, therefore, have been high, narrow and,
judging from tooth size and mandible length, Icng.

The postcranial skeleton of this species is known only from the distal end
of a humerus and the proximal end of an ulna (Table 12), which were found
in association with the incomplete skull, L 13097. The humerus has already
been mentioned elsewhere (Hendey 1973a) as being larger than that of a civet
from the Kromdraai australopithecine site and much larger than that of
modern V. civetta. The ulna is correspondingly large.

In respect of tooth and overall skull size, this species was apparently little
different from modern C. mesomelas, but the humerus and ulna of C. mesomelas
are only about half the size of corresponding parts of the fossil specimens. ‘There
are indications that the size discrepancy is considerably less in respect of the
lengths of the bones, suggesting that the Langebaanweg Viverra was a stoutly-
built and probably non-cursorial animal.

No deciduous teeth of modern or fossil civets were available for comparison
with those from Bed 2. However, the dp, and dp, appear little different in
structure to those of the East African early Pleistocene Pseudocivetta ingens
(Petter 1967: Fig. 3), although the dp, of this species is lower crowned. The
dimensions of the Bed 2 deciduous teeth are recorded here for future reference

(@Qaie 19):

Discussion

The principal differences between the teeth of the Bed 2 and Bed ga forms
of this species are as follows:
(1) The Bed 2 form has a smaller M2.

LATE CENOZOICG CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 81

TABLE 13

Dimensions of the deciduous teeth of the Langebaanweg Viverra leakeyi.

dp? dp? dp, dp; dp,
] b ] b ] b 1 b 1 b
L 14459 | 751 272% 43,0. -,- O36 72 2,31 10,5 3:4) 12,8 4,4
L 14460 = a: ae omy 7,9 255 10,0 335 12,9 533

(2) ‘The Bed ga form has a larger M,, with the talonid similar in absolute size
to that of the Bed 2 form, while the trigonid is larger.

In addition, if the mandibles L 1700 and L 2672 are indeed from Bed 2 and

Bed ga respectively, then it is likely that the Bed 3a form was the larger of the

two.

Even though the temporal difference between Bed 2 and Bed 3a may not
be very great, Bed 3a is undoubtedly later and the differences between the
civets from these two deposits may result from short-term intra-specific evolu-
tion. The Bed 3a form would, therefore, be the more advanced of the two. The
larger M? of the Bed 3a form may represent the beginnings of the trend which
led ultimately to the relatively large posterior crushing teeth of modern J.
cwetta. This speculation can only be tested if larger samples of the two forms
become available.

There can be little doubt that the Langebaanweg Viverra is conspecific
with the Laetolil V. leakeyi (Petter 1963). There are grounds for believing that
an element of the Laetolil fauna is broadly contemporaneous with that from
Langebaanweg and, consequently, the presence of V. leakeyi in the Langebaan-
weg fauna is not surprising (see Hendey 19706: 123, 124).

In her discussion on the affinities of V. leakeyi, Petter considered its possible
relationships to Viverra (Viverra) and Viverra (Civettictis). She found that V.
leakeyi, V. bakerit Bose from the Siwaliks and V. (Civeitictis) could, on morphologi-
cal grounds, represent a phyletic series which derived from a V. (Viverra)-like
ancestor. She concluded, however, that owing to uncertainty about the relative
ages of the fossil species, an actual phyletic relationship of this kind could not
be proven. In fact, since V. bakerii is recorded from the Upper Siwaliks (Pilgrim
1932) and is, therefore, almost certainly early Pleistocene in age, and since
V. leakeyt is a late Pliocene species, such a phyletic series might indeed be
possible. The only objection is that the series is geographically muddled, with
the African V. leakeyi giving rise to the Asiatic V. bakerii, which in turn gave
rise to the African V. civetta. The problem is, however, not serious since there
might well have been a wide dispersal of civets of the V. leakeyi and V. bakeri
types in Africa and southern Asia during the late Pliocene/early Pleistocene,
with the lineage persisting only in Africa.

The Laetolil species was known only from teeth and the very curious

82 ANNALS OF THE SOUTH AFRICAN MUSEUM

braincase from Langebaanweg raises an additional complication to the question
of civet relationships and phylogeny. Pilgrim (1932) defined a new viverrid
genus from the Siwaliks, Vishnuictis, and this must now also be taken
into account. This genus is characterized in part as follows:

‘Viverrinae of medium to large size; . . . braincase exceptionally narrow;
upper molars rather large; P? without internal cusp; premolar series rather
spaced, premolars simple; mandible rather stout but shallow; M, with
relatively long trigonid, relatively short talonid; M, rather large, oblong’
(Pilgrim, 1932: 101).
The Langebaanweg species is accommodated quite well by this definition.
Pilgrim recorded two species of Vishnuictis, namely, V. salmontanus which is
a smaller species from the Dhok Pathan, and V. durandi which is a very large
species from the Upper Siwaliks. In respect of size and other characters, the
Langebaanweg species is apparently most like V. durandi, a species characterized
in part as follows:

‘length of P* greater than united lengths of M! and M?; angle between the
axes of P* and the upper molars about 115°; outer borders of M! and M?
continuous, their inner borders being in the same antero-posterior line’
(Pilgrim 1932: 106).

The Langebaanweg species is, however, apparently not quite as large as
V. durandi and, judging from the illustration of the holotype given by Matthew
(1929), the size of the P* parastyle and the size of M1 and M? relative to P4, are
greater even in the Bed ga form of the Langebaanweg species. In these respects,
V. durandi is the more advanced of the two species, which is in keeping with
their inferred relative ages.

V. leakeyi is apparently not conspecific with either Vishnuictis salmontanus
or V. durandi, but in view of the similarities which do exist between the three
species, they should be regarded as congeneric. A simple and convenient
solution to this taxonomic problem would be to regard Vishnuictis as a subgenus
of Viverra, making it a third category within this genus, the others being Viverra
(Viverra) and V. (Civettictis). Considering all the civets in terms of morphology,
time and geography the relationships reflected in the tentative phylogeny
presented here (Fig. 10) were concluded.

Viverra (Viverra) is regarded as the main lineage from which the others
stemmed. During the Pliocene the first of the off-shoots, Viverra (Vishnuictis)
was evolved and this was represented by an African lineage (V. leakeyr) and an
Asian lineage (V. salmontanus—V. durandi), the latter persisting into the early
Pleistocene. The African lineage is here regarded as the one which gave rise
to the third group of civets, Viverra (Civettictis), which has culminated in the
extant V. civetta of Africa, and which included an Asian off-shoot which is
recorded only in the early Pleistocene (V. bakerit).

V. civetta has a poor Pleistocene record, but the Olduvai Bed 1 ‘Pseudo-
civetta’ ingens (Petter 1967) is here regarded as an early representative of the

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 83
Ww
Zz
O Viverra (Civettictis) Viverra (Viverra)
oO Africa Asia
i
O
zg
Ww Elandsfontein V. (C.) sp.
Q Kromdraai V. (C.) sp.
=
nn
a
a Olduvai V. (C.) ingens
V. (C.) bakerii V. (Vishnuictis)
Asia durandi Asia
V. (Vishnuictis)
leakeyi Africa
Z Fre ry (Vishnuictis)
O salmontanus Asia
Q
=!
a

Viverra (Viverra)

Fig. 10. Tentative phylogeny of the genus Viverra.

Viverra (Cuvettictis) group, while the Kromdraai and Elandsfontein civets
(Hendey, 1973a; vide infra) are regarded as later members of the same group.
The opinions concerning the relationships of the African Pleistocene civets
are largely subjective owing to the nature of the recorded material, but it
seems preferable to visualize a single lineage in which there was apparently a
progressive decrease in body size from the Langebaanian to the Cornelian
rather than to regard the isolated fossil records as representing distinct genera,
as in the case of ‘Pseudocivetta’.

Family Viverridae
Subfamily Viverrinae
Genetta sp.

(Fig. 11)
Material

L 11191—Left mandibular fragment with P,, P; and part of P,.

Locality and horizon
This specimen is from Bed 2, ‘E’ Quarry, Langebaanweg.

84 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

This specimen belongs to a very small viverrid. The mandibular corpus is
relatively long and slender and the inferior margin is arched along its entire
length (symphysis to below the anterior part of the masseteric fossa). The lower
cheektooth row consisted of P, to M, and is approximately 20 mm long. The
preserved premolars are narrow and sectorial. The P, has one posterior accessory
cusp and the P, and P, have two.

Fig. 11. Buccal view of the Genetta sp. mandible
(L 11191) from Langebaanweg.

Discussion

Apart from the fact that this specimen is about one-third smaller than
corresponding parts of the mandible of modern Genetta tigrina, they are otherwise
virtually indistinguishable. There can be little doubt that it belongs to a
species of Genetta.

Stromer (1931a) has described a ? Genetta sp. from Kleinzee, but this is
apparently not the same as that from Langebaanweg, since it is much larger.
No other fossil Genetta from South Africa has yet been described. Petter (in
Leakey 1965) mentions a Genetta of apparently small size from Olduvai Bed 1,
but there is no description of this material.

The Langebaanweg Genetta probably belongs to a new species, but as this
could only be poorly defined on the basis of the available material, it is left
nameless.

Family Viverridae
Subfamily Herpestinae
Herpestes Species A
(Fig. 12)
Comment
As with the Langebaanweg Viverra leakeyi, the material assigned to this
species is separated on the basis of its stratigraphic associations.

| 7 |

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 85

Material from Bed 2, ‘E’ Quarry, Langebaanweg

L 11847—Right mandibular fragment with part of P,, and P,; to M,.

L 13040—Right mandibular fragment with M,.

L 13053—Right maxillary fragment with P? to M®; isolated left M!.

L 13054—Right maxillary fragment with I°*, part of P*, and P3.
The following are the remains of juveniles:

L 14461—Right mandible with dc, P,, dp, and M,, just erupting.

L 14462—Right mandible with de and dp, to dp,.

Material from Bed 3a, ‘E’ Quarry, Langebaanweg

L 15630B/1B—Right maxillary fragment with part of P.
L 15630B/1C—Left maxillary fragment with P®.

L 15630B/1D—Part of left P#.

L16240F —Right C.

L 16055D —Right mandibular fragment.

L 15630B/1A—Left mandibular fragment with M,.

L 161771 — Part of right M,.

L.15797B —Right C.

Description

The Bed 2 material belongs to a species of Herpestes which is a little larger
than the modern H. pulverulentus (Tables 14, 60, 61). Its dental formula is
(ea )29.4.2/(?s).1.4.2.

The I? of L 13054 is appreciably larger than those of available H. pulverulen-
tus specimens (n = 14) and, judging from the size of the alveolus, so was the
canine. The pre-canine diastema is longer than in H. pulverulentus, which would
be in keeping with a larger lower canine as well. The most anterior part of
the snout of the fossil species was thus larger than in H. pulverulentus and the
premaxilla projected further forward.

The cheekteeth are situated close to one another and the relative positions
of these teeth, the infra-orbital foramen and the orbit are as in H. pulverulentus.
The P® has a high principal cusp, a small posterior accessory cusp and a well-
developed internal cusp. The P® of L 13054 lacks an anterior accessory cusp
and in that of L 13053 there is a short horizontal shelf anteriorly from which
projects a minute anterior accessory cusp. The P* is triangular in outline with
the posterior surface of the protocone and the lingual surfaces of the paracone
and metastyle in a straight line. The protocone is more anteriorly situated
than that of H. pulverulentus and the parastyle is smaller relative to the paracone
and metastyle. The M! is relatively broader internally than in H. pulverulentus
and the M? is very small and relatively more reduced than in the modern
species.

In general, the upper teeth of the fossil species are similar in structure to
those of H. pulverulentus.

This applies also to the lower teeth, where the most obvious difference is

ANNALS OF THE SOUTH AFRICAN MUSEUM

86

‘DIEGO IDWIONS x

gL 96 99 C6 C‘Ly = a Be oat —= cos ,IIZUIOP YT VSI OF61
See fore Soe am fol = pas — — — = HL4i91T
ne eee ee ee ee ee ef pog
gil Hee | VO) 21 gigi — = = ~~ == = | Fillets area SS
_—<$— _$. |} |] J J 8
C6 9‘ 9°9 36 °9 — — — = — — — obof1'T =
G6 ae L‘9 ae) PL 9 a ake fest = a = Lbgit 7] 8
: SS SS SS Se ee a i a: |e (eras (ke a opog | 3
— — — — — VG 09 — — — — E9061] >
— — — — — — — OL YD 9G 66 gv VSoE1'T
A MOpEq eulo\seIp (snjooaye)
sndio9 q I I I q I I a ee
JO WYSIOH "WN ‘W-'W Didar rd rd-1d O-el fe)

‘QOZUII[ YJ WOT Saysaquazy ig IYI JO soy} YIM posedusos ‘y soroadsg sasaqduazy Samuveqoasur’y] 24} Jo s[qIpueul puke Y}99} IY} Jo SUOTSUSIICT

VI Adv I,

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 87

3 114 115 116

Fig. 12. Buccal view of the Herpestes sp. A mandible (L 11847) from
Langebaanweg.

that in the fossil the M, talonid is shorter relative to the trigonid. It also differs
from H. pulverulentus in that P, is present. The mandibular corpus is more
robust in the fossil and the masseteric fossa, especially that of L 13040, is much
deeper.

The Bed 3a material is, as far as comparisons are possible, similar to that
from Bed 2. The only observable difference is that the Bed 3a specimens are a
little smaller than those from Bed 2. For example, the mandibular corpus and
M, of L 15630B/1A are comparable in size to those of H. pulverulentus and are
appreciably smaller than specimens from Bed 2.

Discussion

On the basis of the material presently available it seems probable that
the Bed 2 and Bed 3a material is conspecific. There are certainly no major
observable differences and the samples are so small that the significance, if any,
of the size differences cannot be assessed.

Relatively little has been published on the smaller fossil viverrids of
Africa, although this group is a significant element in the modern carnivore
fauna of the continent and presumably the same applied during the late
Tertiary and Pleistocene. None of the small viverrids described from the
Transvaal caves (see Cooke 1963: Table 6) is relevant in the present instance,
but it is likely that the Langebaanweg Species A is conspecific with the ? Her-
pestes sp. from Kleinzee (Stromer 1931a). The mandible L 11847 is virtually
indistinguishable from the Kleinzee specimen which was described and figured
by Stromer.

There is apparently no previously recorded fossil species to which this

88 ANNALS OF THE SOUTH AFRICAN MUSEUM

material can be referred and it is clearly not conspecific with any of the extant
southern African species of Herpestes. As with the Langebaanweg Genetta, it
probably represents a new species. It is, however, not named, since the present
study of the smaller Viverridae was rather superficial and a meaningful diagnosis
of a new species could probably not be given.

Family Viverridae
Subfamily Herpestinae
Herpestes Species B
(Fig. 13)
Comment
As with preceding species, the material from Bed 2 and Bed ga is listed
separately.

Material from Bed 2,‘°E? Quarry, Langebaanweg
L 13055—Left maxillary fragment with P! to P4, part of M!, and M?.

Material from Bed 3a, *E” Quarry, Langebaanweg

L 16240P —Left maxillary fragment with P?.

L 16055Q —Left M!.

L 16177G + —Right CG.

L 16177A, B—Right and left mandibular fragments, the former with C and P,
to M,.

L 15588B/B —Left mandibular fragment with P, and M,.

Description

The Bed 2 maxilla belongs to a very small species of Herpestes (Table 15).
It compares in size to corresponding parts of the skull of a modern H. sanguineus
zombae specimen from Malawi (SAM 15797). Apart from the fact that it is

LL
Ben es

Fig. 13. Buccal view of the Herpestes sp. B mandible
(L 16177A) from Langebaanweg.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 89

much smaller it is essentially similar in detail to the Herpestes Species A from
Langebaanweg.

The Bed ga material is grouped with L 13055 since it also belongs to a very
small Herpestes and there are no grounds for believing that the two samples
are not conspecific. Apart from size, the only observable difference between this
species and the larger Herpestes from ‘E’ Quarry is that the former lacks P,.

TABLE 15

Dimensions of the teeth and mandible of the Langebaanweg Herpestes Species B.

Height of
corpus
below M,

Bed 2 L13055 13,8

L16177A _
Bed 3a — | |
L15588B/B

Discussion

There can be little doubt that this species and the larger Langebaanweg
herpestine are congeneric, but because of the appreciable size difference
between the two, they are clearly not conspecific.

The absence of P, is characteristic of some species of Herpestes (Galerella),
and Species B may have affinities with this subgenus. No other conclusions as
to its relationships were reached.

The specific identity of fossil forms of generalized small viverrids such as
the two Langebaanweg Herpestes is a difficult problem to which there seems
to be no immediate solution. If statistically significant samples of the fossil
species were available, and in the present instance there is a good chance that
this will eventually be the case, and these could be compared with large modern
series, it does not necessarily follow that the fossil forms will be distinguishable
from all modern species, in spite of the great time difference involved. For
example, should it prove that Species B cannot be distinguished from the
Malawian H. sanguineus, it does not logically follow that the two are conspecific.

The best way of making meaningful specific identifications of the fossil
species would be by tracing their lineages to their conclusion. This is clearly
an impossible task at present in view of the poor fossil record of the Herpestinae.
In this respect it is as well to note that the major radiation of the Herpestinae,
at least in Africa, apparently took place during the Pleistocene (vide infra),
and the two Langebaanweg species could each be directly ancestral to more
than one modern species, or genus, as well as other now extinct species. Conse-
quently, even if the fossil record of this group was very much better known,

gO ANNALS OF THE SOUTH AFRICAN MUSEUM

it would probably still be difficult to establish a clear picture of relationships.
The problem is compounded by the fact that during the Pleistocene many, if
not all carnivore species underwent size changes. This may effectively eliminate
an apparently useful method of distinguishing between generalized Herpestinae.

Since no other Pliocene Herpestinae have yet been named in Africa, a
simple solution in the present instance would be to provide new names for the
Langebaanweg species and to list all their observable characters in the diagnoses
in the hope that some might prove genuinely useful in distinguishing them
from other species. On the other hand, since it is almost certain that more
material of both these species will become available, and since relevant material
may yet be described from elsewhere in Africa, it is considered preferable at this
stage to defer the naming of the species. It will almost certainly be more fruitful
to consider this matter again at a later date and this may best be done without
having to consider formal species names already in existence.

Family Viverridae

In addition to the material already listed, viverrid remains have also been
recovered at sites 1/1968 and 12/1968 in ‘E’ Quarry (see Hendey 1970a:
81, 86-88). These occurrences are now regarded as belonging in Bed 2, but it is
still considered preferable to consider these fossils separately for the time being.

The site 1/1968 viverrid specimens include a number of isolated teeth and
two mandible fragments. One of the latter belongs almost certainly to Herpestes
Species A, while the other is referred to the Bed 2 Genetta sp. A similar array of
specimens was recovered from site 12/1968, although in this case a larger
number of isolated teeth were recovered. Once again both Herpestes Species A
and Genetta sp. were represented.

Family Hyaenidae

The Hyaenidae are the most commonly represented Carnivora in the
deposits at Langebaanweg and they are in some respects the most problematical
group in this order. Three species have been identified, while there are two
other tentatively identified and informally named species. The latter may repre-
sent aberrant individuals of one or other of the three named species. Each
of the three identified species, and one which was not formally named, is
represented by cranial and associated postcranial remains of at least one indivi-
dual of known provenance.

It was during the late Pliocene and early Pleistocene that the Hyaenidae
reached the peak of their radiation and a large number of genera and species
dating from this period in time have been recorded, principally from Eurasia.
The importance of the Langebaanweg Hyaenidae lies chiefly in the fact that
they are among the first African Pliocene representatives of this family to be
described. Consequently, they provide information on hyaenids dating from a

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE QI

period in time during which significant evolutionary developments were taking
place and they are from a continent which undoubtedly played an important
role in the evolution of the family.

In the present report no mention is made of the as yet unpublished hyaenid
material from the Pliocene of East Africa, although a few casts of relevant
specimens were available. These specimens were, however, taken into account
in reaching the conclusions on hyaenid phylogeny which are presented here.
Hyaenid classification and phylogeny have been the subject of numerous studies
and some of the conclusions of the present study are a departure from previously
held opinions. They will almost certainly not be generally accepted, but the
records of the Langebaanweg hyaenids themselves should prove useful in other
accounts of the Hyaenidae which are still to come.

Family Hyaenidae
Subfamily Hyaeninae
Percrocuta australis n. sp.
(Figs 14, 15)
Holotype

L 14199—A crushed and incomplete skull, including the following teeth:
Part of right P®; left and right P*; left I, to 1,3, C and P, to M,; right
I, I,, CG and P, to M,.

Referred Material

L 13033—Parts of the skull and skeleton of a single individual and including:
Left and right C; incomplete right mandible with P, and M,.
Parts of at least seven vertebrae; three tarsal bones; parts of five
metapodials; seven phalanges.

Locality and Horizon

These specimens are from Bed 2, ‘E’ Quarry, Langebaanweg.

Referred material which is probably from Bed 2:

L 6381 —Left maxillary fragment with P! and P?.
L 5355A, L g140, L 12102, L 12333—Premolar fragments.

Diagnosis

A Percrocuta similar in size to P. eximia Roth & Wagner 1855, but differs
in lacking P,; P?, and P® without anterior accessory cusps; premolars longer;
M, metaconid very small or absent. Metacarpal I not as reduced as in modern
Hyaeninae.

92 ANNALS OF THE SOUTH AFRICAN MUSEUM

Etymology

From australo meaning ‘southern’, to indicate its geographical position in
relation to other species of Percrocuta.

Description
The skull (Tables 16, 17)

The skull of the holotype is so badly crushed and so incomplete that little
can be said of its characters. ‘The post-glenoid process is not as stoutly developed
as those of available modern Crocuta crocuta specimens (n = 5), a feature which
is probably related to the less enlarged cheekteeth and weaker masticatory
apparatus of the fossil. A very strong posterior buttress to the mandibular
condyle is required by C. crocuta with its greatly enlarged cheekteeth and power-
ful muscles of mastication. In L 14199 the infra-orbital foramen is situated just
above the antero-lateral root of P*, rather than above the anterior root of P?
as in the C.. crocuta comparative series. In this respect it resembles the P. eximia
specimen (BMNH M 4162) from Samos which was illustrated by Pilgrim:
(LOBULE ol, 1).

The maxillary fragment L 6381 has attached that portion of the pre-
maxilla which separates the nasals from the maxilla. It is longer than corre-
sponding parts of the premaxilla of the available C. crocuta specimens, and in
lateral view it is visible along its entire length. In C. crocuta the most distal
portion of the premaxilla curves into a horizontal position and in lateral view
is partially obscured by the maxilla.

The nature of the M! of the fossil is not known.

The P* is comparable in length to that of C. crocuta, while the breadth is
also similar, although the protocone of the fossil is somewhat reduced. The latter
is a characteristic of the genus Percrocuta (Kurtén 19575), although in the present
instance the protocone reduction is not as marked as in species such as P.
tungurensis and P. grandis and nor is it as posteriorly situated. In the P* of L 14199
the parastyle, paracone and metastyle are more or less equal in length, which
is In contrast to the condition in C. crocuta where the parastyle is shorter than
the paracone, which in turn is shorter than the metastyle.

Although only the posterior portion of the holotype P® is preserved, it is
important since it also exhibits Percrocuta rather than Crocuta characteristics.
The posterior accessory cusp is much more prominent than that of Crocuta
and there is a well-developed cingulum which extends from the lingual surface
opposite the principal cusp to behind the posterior accessory cusp. The fossil P?
is also narrower and less high crowned than that of Crocuta.

The P? of L 6381 is much smaller than the P? of the holotype. The posterior
portions of these two teeth are similar in morphology, except that the P? has a
greater development of the posterior-internal region. The P? lacks an anterior
accessory cusp but, like C. crocuta, it has a well-developed anterior keel on the
principal cusp which terminates antero-internally at the cingulum.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 93

in

112

Fig. 14. Occlusal and buccal views of the Percrocuta australis mandible (L 14199) from
Langebaanweg.

The P! of L 6381 is a relatively small tooth, which is similar in size and
morphology to that of C. crocuta.

The upper canines of L 13033 are very worn, but it appears that, compared
to those of C. crocuta, the crowns were small relative to the size of the roots.

Although they belong to different individuals, the left canine of L 13033 fits

O4 ANNALS OF THE SOUTH AFRICAN MUSEUM

daa

uta wi nuit

iu

HEAT

itl

i

ng

(I

TULL

nm

mn

HN

ul

mm

i

i

+

Fig. 15. Occlusal and buccal views of the Percrocuta australis mandible (L 13033) from
Langebaanweg.

quite well into the canine alveolus of L 6381.

Although both halves of the mandibular corpora of the holotype are
preserved, the posterior portions are detached, crushed and fused together by
the phosphatic matrix which is characteristic of the upper levels of Bed 2 in the
vicinity of East Stream. When articulated, the anterior parts of the corpora are
V-shaped in ventral view, rather than being arched and almost U-shaped as
in C. crocuta. The symphysis is similar in shape to that of C. crocuta and also
terminates below the posterior roct of P,. It is, however, somewhat longer since
the depth of the fossil corpora below P, is greater than in C. crocuta. The fossil
mandibular corpora, especially in L 13033, are very robust and the overall
length of the mandible is greater than those of the C. crocuta comparative series.
The masseteric fossa of L 13033 is very deep and there is a horizontal ridge
of bone along its inferior margin which is more or less parallel to the inferior
margin of the mandible in this region. In C. crocuta this ridge and the inferior
margin of the mandible towards the angle are inclined upwards, with the result
that the condyle is higher relative to the cheekteeth than is the case with the
fossil.

95

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

‘gh61 nry 2 Sunox ,

vGG6I IaMy z
‘oS61 UdTIIOT, ,

avs

gSaprouand ad
nuavdIaaT

grafsof”

synuavd py

puavdy nuavd py
Ulpopy

OOOO OS eee eee) ee | | | |

— — gor gc aL

L‘Q%

61

tuvdvyvu
puavdy vuavd yy

OOO rrr ——nsess ds ee eee ds | | | |

o'Gs ‘9

xafrofard
synuand yy

—_—_—_—_—e.?.?}?}__—— _e—_:.:.:.:.:. eee | | Ns ON

19‘
GG1 o%%
gS‘1
OFFI =z
€G‘1
1 bos
oS‘
gai = B61
0g‘I
okt a a ae 4p a
gS‘1
aE = 90%
bot
er 69'O1

Lint
o‘11 S‘6r
gS‘1
bor ‘QI
£951
L‘6 Q‘G1
GG‘1
v6 g‘b1
6L‘1
19 CFI °
19‘I

GOLF Ty *

q so1oedg
puavd Ey

DIUOAGD
puand py

SCS | | | | fa

oQ*I
og 6°41

PL‘1
Q‘II Z'0%

L‘S —
19 —
a9 1°6
oG:9| —
— G‘or
PG 96
2g —

SUDAISND
DINIOAIAIT

‘soroods pruovdy Joy} dws Jo asoy} YIM poreduioo ‘Somuveqosuey] wo sepruseAzy Jo yI99) saddn oy} jo SUOISUSULIICT

QI ATaV],

ANNALS OF THE SOUTH AFRICAN MUSEUM

96

‘9r61 n'y 2 BuNOZ

‘DGG61 IIMq z

‘oS61 UdTII0], _

,Saplouandany
puantaaaT

araflof
synuavdpy

puandy nuand yy
UIIPO/[

tuvdoyout
puandy puavd py

xaflo fog

syouavdyy

ot o'Gs
— Q‘Vz
GSI L‘vs
Gol 6‘0%
6‘6 050%
06 0°61 °9
I‘1I o'Ss

gh‘t
Ol OVE
691
691
Sol (Aural
611 che)
git
G11 Goa
gh‘t
6‘o1 o‘61
£651
OLOonn an
06‘1

q soroods
puavd ey

DIuU0GD
Duane py

SUDAISND
DINIOLIAIT

Pol 1°1d
Por 660%
G‘s1 °9 b6a ‘9
q I
W

961
Teche ORO cit
6S1 +9 Lbe

q I
"d

oL‘t
DEI gas
oS‘
vG‘1
eon 661
gsr L “61
Lost
Vit 061
99‘1
Homi Ni
19‘I
‘6 1°G1
26‘1
G‘o1 rca OY 6
oQ°I
9‘6 CL
Vet
OM < Ma he ok
q I
*d

Lo‘t
ofor L‘or
VO‘1
G‘9 6°E1
Cael
9‘L VSI
CL°1
Ti VS
€9°1
£°9 o‘G
LL*1
gL Q°E1
G91
Gor v¥‘61
l I
“d

— Sor Si
gt <= =
ov 9‘6 VSI
— VE1 = 1°QI
— eyes) WVhi
I al I
0)

Li aTavy,

‘sotoods prusvAy 19430 IUIOS JO 9soy} YIM poseduiod ‘Bamueeqosuey] wo sepruseAP] JO YI99} IOMOT DY} JO suoTsusMIG,

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE Q7

The lower incisors of the holotype are smaller than those of C. crocuta,
but are similar in their relative proportions. The I, is also lower crowned and
does not have such a prominent lateral projection.

The lower canines of the holotype are more transversely compressed than
those of C. crocuta and also appear to have larger roots. The holotype canines
are remarkable because they are worn in the way which is unusual in hyaenids.
They lack horizontal wear facets and instead have vertical wear facets extending
from the postero-external surfaces of the base of the crowns to the anterior
surfaces near the original apices of the crowns. This is probably an individual
peculiarity since the upper canines of L 13033 do have horizontally worn
crowns in addition to wear facets on their anterior surfaces.

Both halves of the mandible of the holotype lack P, and the diastema is
longer than in any of the C. crocuta comparative specimens. The anterior portion
of the mandible of L 13033 is damaged, so it is not known whether or not P,
was present in this individual.

The P, of the holotype is shorter and narrower than the P? of L 6381. It
also narrows anteriorly and lacks the slight inflation of the postero-internal
cingulum evident in the P?. It lacks an anterior accessory cusp. The P, of
C.. crocuta is shorter, relatively much broader and lower crowned.

The holotype P, is appreciably larger than P, and, except that it is more
or less rectangular in outline, it is otherwise similar to P, in morphology.

The P, resembles P, in that it also narrows anteriorly, but it differs from
both P, and P, in having a fairly prominent anterior accessory cusp and a more
pronounced postero-internal cingulum. This part of the cingulum also has aslight
vertical component, so that there is a small postero-internal cusp flanking the
posterior accessory cusp. This is less marked in L 13033 than it is in the holotype.

Although the P,, P; and P, of C. crocuta differ quite considerably from the
corresponding teeth of the fossil species, there are certain basic morphological
similarities. The lower premolars of the two species have the same cusps
represented, but there is a difference in their relative sizes. In both species
broadening is most evident in P3. The least dissimilar of the premolars is P,
and it is really only the more rectangular outline and reduced anterior accessory
cusp of the C. crocuta P, which differentiates it from the fossil.

Both lower carnassials of the holotype are crushed and incomplete. The
talonid is short, although not as short as that in C. crocuta, and the paraconid is
a little longer than the protoconid. The metaconid is present, but is very small.
The M, of L 13033 is similar, except that the metaconid is apparently absent.

The holotype lacks M, and it is likely that this tooth was absent in L 13033
as well.

Postcranial skeleton (Tables 25, 26)

The only postcranial remains which are positively identified as belonging
to this species are some vertebrae and parts of the right manus and left pes of
L 13033. The individual skeletal elements are most readily distinguished from

98 ANNALS OF THE SOUTH AFRICAN MUSEUM

those of available skeletons of modern C. crocuta (n = 1) and H. brunnea (n = 2)
by their larger size.

Of the seven vertebrae known, only one, the 7th cervical, is largely intact.
This specimen is only slightly larger than the 7th cervicals of the modern com-
parative specimens, but the centrum of the fossil 2nd cervical is appreciably
larger.

The manus is represented by a metacarpal I, the proximal end of meta-
carpal V and possibly some of the phalanges. The metacarpal I, a bone which is
vestigial in modern Hyaena and Crocuta, resembles that of Proteles cristatus in
morphology, but it is considerably larger. Its size relative to the remaining
part of the metacarpal V is, however, similar to the relative sizes of these bones
in Proteles. The metacarpal V is larger than those of the Hyaena and Crocuta
comparative specimens, but is morphologically similar, although in dorsal
view its proximal articular facet is slightly concave, rather than straight or
slightly convex as in the modern specimens.

The tarsal bones of L 13033 which were recovered are a navicular, a
cuboid and an internal cuneiform. They are much larger than those of the
comparative specimens and although there is a basic morphological similarity
between the navicular and cuboid of the modern and fossil specimens, there
are some differences in detail. The navicular has a transverse diameter com-
parable to that of the available C. crocuta specimen, but it is broader than those
of the two H. brunnea. Its antero-posterior diameter is greater than those of all
three modern specimens. It also differs in that it lacks the deep median indenta-
tion at the posterior end of the proximal articular facet, while the medial facet
of articulation with the cuboid is circular rather than antero-posteriorly
elongated. The facets on the distal surface which articulate with the cuneiforms
are more distinctly separated from one another than is the case in the modern
specimens. There are comparable differences in size and in the nature of the
articular facets in the cuboid. The internal cuneiform differs from those of
the moderns only in its larger size.

The metatarsals IJ and V, which are intact, are similar to those of the
comparative specimens, except again for their larger size. The metatarsal II
has an abnormal growth of bone near the distal end of the shaft and the trochlea
is inflected ventrally at a slight angle to the shaft. A similar pathological con-
dition is evident in one of the 1st phalanges, while another is barely recognizable
as a phalanx owing to a severe condition which has left the bone spongy,
deformed and incomplete. The cause of the pathology is not known, but it
might be the result of an old injury.

The phalanges are, in general, similar to those of the modern specimens,
except for their larger size and the fact that the terminal phalanges are more
strongly arched.

The relative sizes of fore- and hindlimbs of this species cannot be accurately
assessed on the basis of the available material, but the relative sizes of the
proximal ends of the metacarpal V and metatarsal V are not as disproportionate

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 99

as those of extant hyaenids.

The postcranial material indicates that the fossil species was far larger
than modern C’. crocuta and since the known skull parts are only a little larger
than corresponding elements in C. crocuta, it suggests that the fossil form was not
as curiously proportioned an animal as C. crocuta.

Discussion

The affinities of the large hyaenid from Langebaanweg clearly lie with
Percrocuta Kretzoi, 1938 rather than Crocuta Kaup, 1828. The former genus
has been the subject of a critical study by Kurtén (19575) who tentatively
regarded it as a subgenus of Crocuta. Both Kurtén and Pilgrim (1932) suggested
that there might be no direct phyletic connection between Percrocuta and
Crocuta and ‘Thenius (1966) subsequently indicated a major phyletic separation
between them and allowed Percrocuta full generic status. Ficcarelli & Torre
(1970) went a step further and recognized two genera within this group,
namely, Percrocuta and Adcrocuta Kretzoi, 1938. It is, however, the arrangement
of Thenius which is accepted here.

The Pliocene Percrocuta populations do fall into two groups, one culminating
in the very large P. gigantea, Schlosser 1903, and the other in the smaller
P. eximia of Eurasia. It is with the latter group that the Langebaanweg species
apparently has its closest connections. The graphic representation of ratios
devised by Simpson (1941) and used for cheektooth lengths by Kurtén (19576)
in his analysis of Percrocuta, gives a clear indication of the similarity between
P. eximia and the Langebaanweg species (Fig. 16).

Kurtén (19575: 399-400) recognized three subspecies of P. eximia and these
range in age from early to mid Pliocene. In view of the late Pliocene age of the
Langebaanweg species, it might be expected that it would be more advanced
than the recorded subspecies of P. eximia. This is indeed the case and it is on
these grounds that the specific separation is justified, while the wide geographic
separation between P. eximia and the Langebaanweg species is intended to be
suggested by the proposed new name for the latter, P. australis.

The most important observable differences between P. australis and
P. eximia are that the former lacks P, and anterior accessory cusps on P?, and Ps,
while its premolars are more elongated than those of P. eximia. P. australis
may well be a descendant of P. eximia, the ancestral form having been present
in Africa earlier in the Pliocene when this species was at its peak and widely
distributed in Eurasia as well.

Since the nature of the relationship, if any, between the late Tertiary
Percrocuta and Quaternary Crocuta is obscure, and since P. australis is apparently
the most recent of the smaller-sized Percrocuta group, it is of interest to consider
the possibility of a relationship between the Langebaanweg species and Crocuta.

Kurtén (19575) concluded that none of the Percrocuta species known at
that time could be ancestral to Crocuta (sensu stricto). He regarded C. sivalensis
Falconer & Cautley, 1868 as the earliest recorded direct ancestor of C. crocuta

100 ANNALS OF THE SOUTH AFRICAN MUSEUM

(ener mes) &
&

06 0 06 12 18 24
= +

Fig. 16. Ratio diagram comparing relative lengths of the cheekteeth of Percrocuta carnifex (2),
P. eximia eximia (3), P. gigantea (5) (Kurtén 19576) and P. australis (4), with modern Hyaena hyaena
(1) (Kurtén 1956) as a standard.

and visualized a hypothetical ancestor of the former species as having the
following characteristics:

) M, metaconid well-developed.
2) M, talonid long.

) Protocone of P4 large.

) Anterior premolars heterodont.

Both Kurtén (1957)) and Thenius (1966) thought it likely that Crocuta
was descended from Lycyaena chaeretts Gaudry, 1861. Since this species is a rela-
tively primitive and rather generalized hyaenid dating from the earlier part of
the Pliocene, there is little difficulty in regarding it as an ancestor of Crocuta.
It is however, perhaps significant that later species of Lycyaena (e.g. L. lunensis
from Europe and L. nitidula from South Africa) were ‘non-scavenging fully
predaceous forms in which the trend towards the development of heavy conical
hammer-like premolars suitable for bone-crushing was reversed’ (Ewer 1955):
851). While it is not impossible that L. chaeretis also gave rise to a lineage in
which the typical hyaenid trend was not reversed, it is curious that this is not
reflected in the fossil record.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE IOI

On the other hand, it is equally curious that the Crocuta-like Percrocuta
group are essentially late Tertiary in age, while Crocuta itself is confined to the
Quaternary, and that the apparently logical chronological succession has not
been supported by the evidence of developing morphological characters.
If it could be demonstrated that P. australis was more like early Crocuta than was
P. eximia, then it might be accepted that a direct phyletic connection between
Percrocuta and Crocuta did exist. However, this does not appear to be the case.

The four characteristics which were listed above and which were regarded
by Kurtén as being likely in an ancestor of C. szvalensis are not convincingly
evident in P. australis. In addition, there was apparently a trend towards the
elongation of the premolars in the P. eximia—P. australis lineage and this would
have had to be reversed if the latter was indeed ancestral to C. crocuta (Fig. 17).

Since Percrocuta has now been recorded from sub-Saharan Africa, there
may yet be evidence forthcoming from the region which could substantiate a

4 1 2 3
p2 +> s »
F 6
ae
a
4
4
p3 4 eCESTIMATE

,06 0 06 A2 18
- 4
Fig. 17. Ratio diagram comparing relative lengths of the cheekteeth of Percrocuta eximia eximia (2),

modern Crocuta crocuta (4.) (Kurtén) 1957, 1956) and P. australis (3), with modern Hyaena hyaena
(1) (Kurtén 1956) as a standard.

102 ANNALS OF THE SOUTH AFRICAN MUSEUM

direct relationship between Percrocuta and Crocuta.

Another opinion on the origins of Crocuta was recently expressed by
Ficcarelli & Torre (1970), who concluded that this genus had an independent
history dating back to the Miocene. This view is apparently at least partly
based on the belief that C. honanensis Zdansky, 1924 from China is early Pliocene
in age. No explanation is given of the otherwise complete absence of Crocuta
in the Miocene and Pliocene, times when the fossil record of hyaenids is by no
means poor.

Another African record of Percrocuta is P. algeriensis, which was described
by Arambourg (1959) as a species of Hyaena. Ewer (1967) also regarded it as an
Hyaena, and while Ficcarelli & Torre (1970) had doubts about its relationships,
they refer to it as Hyaena algeriensis in their text. Thenius’s (1966) grouping
of this species with Percrocuta is accepted here (Fig. 18), since this is its most

Lu
Zz
Lu
U
O
kK
WN
io =5
—4
QQ
P. gigantea
P. australis Asia
Africa
Lu
FL
Lu
UO
O P. eximia P. grandis
= Eurasia Asia
P. algeriensis P. carnifex P. tungurensis
Africa Asia Asia
Ww
= P. miocenica
U Europe
O P
=

PERCROCUTA

Fig. 18. Tentative phylogeny of the genus Percrocuta. (Adapted from Thenius 1966.)

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 103

likely phyletic position. P. algeriensis is apparently late Miocene in age and
since Hyaena was probably descended from Pliocene Ictitherium, it is most
unlikely that there can be any direct phyletic connection between P. algeriensis
and Hyaena. This species is most likely to be comparable to P. tungurensis, which
seems to have belonged in a monotypic and ‘highly precocious phylum’ that
became extinct without issue (Kurtén 19575: 401). The relationship between
P. australis and P. algeriensis is probably no closer than that between P. eximia
and P. tungurensis.

Family Hyaenidae
Subfamily Hyaeninae
Hyaena abronia n. sp.
(Figs 19, 20, 21)
Holotype

L 14186—A skull lacking only a few parts and an incomplete skeleton of which
only the thorax is not represented.

Referred material
L 13167—Right mandibular fragment with M,.

Locality and horizon

These specimens are from Bed 2, ‘E’ Quarry, Langebaanweg.

Diagnosis

A species of Hyaena comparable in size to H. hyaena Linnaeus. Braincase
larger relative to face than in H. hyaena; post-orbital region broad and frontals
not much inflated; premaxilla less protuberant than in H. hyaena; palate
longer and broader. P,, M, present; M? sometimes present; canines and anterior
premolars are, in general, smaller and lower crowned relative to those of
1. hyaena; P3 lack anterior accessory cusps and these cusps are small in P3;
posterior accessory cusps of P3 are small; P* protocone, parastyle and paracone
reduced relative to those of H. hyaena; M? relatively large and triangular in
outline. Fore- and hindlimbs more or less equally proportioned; metacarpal I
approximately half the length of metacarpal IT; tail long.

Etymology

From abronia meaning ‘graceful’, a reference to the body proportions of
this species, which contrast with the more clumsy appearance of modern
Hyaeninae.

Description

The holotype of this species is the most complete skeleton of any species
ever recovered at Langebaanweg. The skull, some parts of which have been

104 ANNALS OF THE SOUTH AFRICAN MUSEUM

reconstructed, one half of the mandible, a few vertebrae and parts of all four
limbs are known. The skeleton is that of a mature adult and the teeth are
only moderately worn.

Skull (Tables 16, 17, 18, 19)

The skull with the mandible still articulated was damaged by a mechanical
excavator, but relatively few parts were lost and it has been possible to almost
completely restore the specimen. The upper dentition lacks only the left I’. The
left half of the mandible is largely intact and the dentition is complete. tne
right half of the mandible is represented only by the ascending ramus.

The skull is comparable in size to that of H. hyaena, but it differs from
this species in certain details. The braincase of the fossil is larger relative to the
face than those of three available H. hyaena skulls (Table 18). This is at least
in part due to the greater development of the premolars in the modern species.
The sagittal crest of L 14186 is slightly less prominent than is normally the case
in H. hyaena and consequently the height of the fossil braincase is less. The
sagittal crest of H. hyaena apparently usually terminates anteriorly at the contact
between the parietals and frontals. Anteriorly from this point are two ridges
of bone (temporal ridges), which may be parallel to one another for a short
distance, but which then curve laterally terminating at the posterior edge of the
post-orbital processes. here is a similar arrangement of these features in the
H. abronia holotype, except that the bifurcation of the temporal ridges is already
quite marked at the parietals/frontals contact and their actual divergence
begins at this point. Related to this is the fact that the post-orbital region of the
fossil is much more expanded than that of H. hyaena. On the other hand, the
frontals on the dorsal surface between the orbits are less inflated. The post-
orbital processes of the zygomata are much less prominent in the fossil and the
rest of the zygomata have a lesser vertical development as well. The more
prominent sagittal crest and stronger zygomata in H. hyaena indicate that the
muscles of mastication are more powerfully developed in this species.

TABLE 18

Ratios of braincase to facial lengths in Hyaena abronia and Hyaena hyaena.

(1) Facial (2) Braincase (12)
length* length*
Hyaena abronia L14186 . . . 104 6. 135 if 35,310
Modern
Hyaena hyaena BMNH 39.440 . 110 120 if OO)
BMNH 23.3.4.9 . 115 DQG 1:1,10
SAM 36335 . . 103 122 IES IS)

* Point of division taken on midline of skull immediately posterior to the postorbital projections.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 105

Differences between the posterior and ventral parts of the braincases of
H. abronia and H. hyaena are more difficult to assess owing to the variation of
features in these regions in the latter species. One apparently constant difference
is that in the fossil the basi-occipital is broader between the anterior parts of
the bullae, whereas in H. hyaena the broadest part is more posteriorly situated.
There is also a greater development of the posterior part of the tympanic
region in H. abronia.

The palate of H. abronia is longer than those of the fourteen H. hyaena
skulls examined early on in this study (Table 19), and it is broader than those
of the three H. hyaena skulls examined during the latter stages of the present
investigation. On the other hand, the premaxilla of H. hyaena projects further
forward than that of H. abronia, owing to the longer pre-canine diastema of the
former. This longer diastema is directly related to the larger size of the lower
canine in H. hyaena.

In H. abronia the infra-orbital foramen is situated slightly more posteriorly
than in H. hyaena and the orbits themselves are smaller in the fossil species.

Taking into account the age disparity between the two species, there are
remarkably few major differences in skull morphology.

The same applies in the case of the mandibles of the two species. ‘They are
comparable in size, but there are a few differences which are significant. ‘The
symphysis of H. abronia is shorter than those of three H. hyaena specimens and
covers a smaller surface area. This suggests a weaker connection between the
two halves of the mandible. The most marked differences are, however, in that
region posterior to the cheekteeth, which is where the muscles of mastication
attach. The fossil mandible has a very prominent subangular lobe, posterior
to which the inferior margin at first inclines upwards and is then horizontal for a
short distance anterior to the angular process. The angular process itself is
bulbous in lateral view. In H. hAyaena the subangular lobe is a less prominent
feature and the inferior margin of the mandible below the ascending ramus
continues to incline upwards to the very end of the angular process. This
process is more elongated than that of H. abronia and in lateral view has a slight
dorsally directed termination. The result of these differences is that in H. hyaena
the condyle is more elevated relative to the cheekteeth. In H. hyaena specimens
with teeth in a similar state of wear to those of the H. abronia holotype, a line
drawn from the dorsal surface of the condyle to the top of the crown of I,
passes well above the cheek teeth, whereas in L 14186 it intersects the crowns of
M, and P,.

It is likely that all the significant differences between the H. abronia and
H. hyaena skulls and mandibles relate directly to the feeding habits of the two
species, a conclusion which is supported by the differences in their dentitions.

In general, the teeth of H. abronia are a little smaller than those of H.
hyaena, while the cheekteeth of the fossil are both more numerous and more
evenly spaced. They are also smaller and much less specialized than those of
H. brunnea and Crocuta crocuta.

106 ANNALS OF THE SOUTH AFRICAN MUSEUM

Width across occipital a) aS
condyles a Uke

Skull height Bs
(Occipital condyle— a oS
sagittal crest) = =
(Bulla —sagittal crest) 09 =
Skull height co a
a
ot
Om

78,1

Mastoid width

aS
~ ec)
S . . Ye) Ne
3 | Post-orbital width + ms
S y ca WN
8 S
g
SS)
8
a)
: : ~ eo) | Cex
8© | Inter-orbital width s | eS
BS a Rt |) Ser
q
a
3
Ta ° S| 6
aD se Te Co). Ile ee)
= & | Zygomatic width = co | te
fc
S| 4 S S
a o
2
Sl es
le) io} A ee
= | Width across canines Ss | Ss
Le) —
a S
oO
G
eo Yo)
Se Palate S | one
z) (Anterior incisors— - fo easy
posterior palatine incisure) 8 | is
c
B)
& °
Q n Oo =
Lateral length of nasals oS | IG
: i)
S
; 1S)
Median length of nasals af | iG
To) 4
2 2
Condylobasal length i= Se | 32
nN a On
a
ive)
D
S
* enn Lo
BS a =
ej S S =e rs)
eS | SS] Ss
Sel Sei ae oe
+) Ss] “0 *
— OOS} . g
ea seh tS se) he

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 107

As an exception to the previously stated generalization, the Ij and I? of
H. abronia are a little larger than those in the H. hyaena comparative series.
This applies also to I;, but the I? of H. Ayaena is larger, not in diameter, but in
crown height. Much the same applies in the case of the canines, although in
H. abronia these teeth are also a little shorter.

The P! of H. abronia differs from that of H. hyaena only in that it is slightly
smaller. The fossil P? is also smaller and, in addition, it lacks an anterior
accessory cusp. The P? is narrower, lower crowned and has a more prominent
posterior accessory cusp.

The upper carnassial of H. abronia differs from that of H. hyaena in several
respects. The protocone is smaller and more anteriorly situated and its posterior
edge meets the lingual margin of the shearing blade at an obtuse angle. In the
available comparative specimens this angle is nearer go°. Also in H. hyaena,
the parastyle and paracone are more or less equal in length, while the metastyle
is shorter, whereas in the H. abronia P* it is the paracone and metastyle which
are similar in length and the parastyle which is shorter. The H. Ayaena P* is thus
differentiated from that of H. abronia by a greater development of its three most
anterior cusps.

In the upper dentition the most marked differences between the two
species are in the post-carnassial teeth. The M! of H. abronia is larger than that of
H. hyaena and is more or less triangular in outline, whereas in H. Ayaena reduction
of the postero-external part of M? has left it nearly oval in shape. In the H.
abrona holotype there is an M? present on the right side. It is a small, single-
rooted and almost circular tooth. There are no indications that the left M?
was ever present in this individual. This tooth is apparently never present in
H. hyaena. In the fossil the buccal margins of M! and M? make an angle of
approximately 110° with the long axis of P* and both molars are partly visible
in lateral view. In H. hyaena the M? is hidden behind P* in lateral view and the
angle between its posterior margin and that of the long axis of P* is about go’°.
The longer palate of H. abronia is at least in part due to the nature of the post-
carnassial dentition.

The most striking feature of the lower dentition of H. abronia is that both
P, and M, are present, a characteristic which distinguishes it from the extant
hyaenid species.

The P, is a small, single-rooted and almost circular tooth, very much like a
reduced version of P?. It is situated a little forward from P, and the postcanine
diastema of H. abronia is consequently shorter than that of H. hyaena. The fossil
P, is similar to P?, the principal difference being that it lacks a prominent
internal cingulum. Unlike the P, of H. hyaena, it does not have an anterior
accessory cusp. Morphologically P, is essentially similar to P,, but differs in
being broader anteriorly than it is posteriorly. There is a pronounced bulging
of the cingulum at the most anterior end of the tooth which is almost distinct
enough to be regarded as an anterior accessory cusp. It is smaller and lower-
crowned than the P, of H. hyaena, a species which has a more distinct anterior

108 ANNALS OF THE SOUTH AFRICAN MUSEUM

a

il

mn

Huu

nil

i

II

lg

Mai

na

| TT

mM

UU

hi

TL

ha

mm

Itt

nit

tH

itll
6 z

HAI

il

Fig. 19. Dorsal and ventral views of the Hyaena abronia skull (L 14186) from Langebaanweg.

——

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 109

1

TL UL

° 2i1 212 2

a A

112

ni

uu
6 1

IU UN

nn

mM
3 1

Hy
4

mn

5

A
sit laa

mm
2 1

nm
0 WW !

mm
3 1

i

Mn
6 117 1

i
8 1

0c

9

i

OU

a

HU
8 1

nD

nl

mn

nr
0 Wi 1

HUI
2 1

mM
3 1

TL YULI/UNHN v(t

Fig. 20. Buccal, lingual and occlusal views of the Hyaena abronia mandible
(L 14186) from Langebaanweg.

IIo ANNALS OF THE SOUTH AFRICAN MUSEUM

accessory cusp and a posterior accessory cusp which is relatively less prominent.
In both species, the P, is essentially the same, although that of H. hyaena is
larger and has a slightly more prominent anterior accessory cusp. __

The lower carnassials of the two species are also similar, except that in
H. abronia the talonid of this tooth is slightly longer, while the metaconid is a
little more prominent.

The M, of H. abronia is small, single-rooted and slightly elongated. It is a
little larger than P, and has a more complex crown made up of four tiny cusps,
two situated lingually and two smaller and more closely set cusps situated
buccally.

Postcranial skeleton (Tables 20 to 26)

The holotype of H. abronia is remarkable in comparison to many other
early hyaenids since its postcranial skeleton is so well represented. Relatively
little attention has previously been paid to the postcranial skeletons of fossil
carnivores, probably because they are often not well represented and they
are certainly less useful for taxonomic purposes than skulls.

Modern Hyaenidae are perhaps the most aberrant group of carnivores in
respect of their body proportions and it is therefore appropriate that when
opportunities arise to examine these proportions in ancestral forms, this should
be done. In a classic study on animal locomotion, Howell (1944:51) referred
to the Hyaenidae as follows:

‘For carnivores of such size the hyaenas are rather slow, cumbersome beasts.
Nevertheless, they are of very real interest in any study of locomotion because
of the fact that the forequarters are much heavier than the hinder ones and the
former are employed chiefly in progression, while the hind limbs are used
chiefly to maintain balance, at least during the low speeds observed in cap-
tivity. The proportions of their limb segments are noteworthy among all the
carnivores.’

In the present study, the functional aspects of the postcranial skeleton of
H. abronia are touched upon, but attention is focused principally on the size
of individual skeletal components relative to those of the two available skeletons
of modern H. brunnea. In view of the conclusion on the relationships of H.
abronia, it would have been more appropriate to make comparisons with the
skeleton of H. hyaena, but such material was unfortunately not available. The
study did little more than demonstrate that H. abronia had a skeleton very much
as would be expected of an early form of Hyaena. However, the descriptions
which follow may ultimately prove useful in studies aimed specifically at
elucidating aspects of the evolution of the curious skeleton of modern hyaenids
and the functional implications of changes in elements of the postcranial
skeleton.

The postcranial material recovered in association with the skull L 14186
is as follows:

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE III

Vertebrae (Table 24)

Of the cervical vertebrae, only the 7th is known. It is similar in morphology
to those of the available H. brunnea skeletons, but it is appreciably smaller,
particularly in respect of the size of the neural arch. This is in keeping with the
much greater development of the skull and neck muscles in H. brunnea.

A single slightly crushed and incomplete lumbar vertebra was recovered.
Its position in the series is uncertain, but it was probably the 4th (second last).
As far as can be judged, it is similar in size to that of H. brunnea, except that
the transverse processes have a greater antero-posterior diameter.

The largely intact sacrum was also recovered. It is made up of three
completely fused sacral vertebrae, the same number present in the H. brunnea
comparative specimens, although Flower (1885) lists this species as having four
sacral vertebrae. The fossil specimen is narrower than that of H. brunnea and
the sacral foramina are smaller, but the total antero-posterior length of the
centra is greater.

Five complete caudal vertebrae and the posterior half of a sixth are known.
The incomplete specimen is probably the roth in the series, while the others are
the 11th to 15th. They are notable because they are about one-third longer than
the corresponding ones of H. brunnea. This species has a tail length of about
300 mm (Fitzsimons 1919), and consequently the tail of H. abronia is likely to
have been about 400 mm in length. This is considerably in excess of the tail
length of 240-280 mm in H. hyaena (Harrison 1968).

Since the skull of the H. abronia holotype is smaller than that of H. brunnea,
the size differences in cervical vertebrae are readily accounted for, but it is
notable that the length of the centrum of the fossil 7th cervical is little different
from that of H. brunnea. This apparently applies in the case of the lumbar
vertebra as well. Consequently, it appears that the length of the spine from skull
to pelvis was approximately the same in the two species. The fossil sacrum is
longer and narrower than that of H. brunnea, while the tail of H. abronia was also
appreciably longer. The latter is regarded as a primitive characteristic exhibited
by this species and so too is the lesser development of the neck.

Forelimbs (Fig. 21; Table 25)

A large part of the right scapula and some parts of the left are known. The
overall length of the fossil scapula is approximately the same as that of H.
brunnea, but it has a smaller glenoid fossa, narrower neck and, apparently, less
expanded blade.

The right humerus is largely intact. It is only a little shorter than those
of the H. brunnea comparative specimens, but it is appreciably more slender.
In spite of being more lightly built, the deltoid crest is very prominent and so is
the projection where the medial ligament of the elbow joint attaches. In other
respects the humeri of the two species are similar, although the articular surfaces
are much larger in H. brunnea.

The left radius is intact and the right lacks only the distal end. The

I12 ANNALS OF THE SOUTH AFRICAN MUSEUM

.
NS
,
,
¢
/
7
,
,
/
‘
¢
‘
(
\
\
\
N
\
Nee
Seo
NO
‘
Sh \
NY
Sos
RS
.
.
.
U
/
oe ee
a)
f \
t) )
‘
/
/

Fig. 21. Elements of the forelimb of the Hyaena abronia (L 14186) from Langebaanweg.

-

-
SSo on DOSS eee

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE II3

complete specimen is slightly shorter than that of H. brunnea, but it does have a
somewhat stouter shaft. There is a very considerable difference in the sizes of the
distal ends, that of H. brunnea being larger. There is a corresponding, but less
pronounced difference in the sizes of the proximal articular facets. The fossil
specimens are thus more evenly proportioned along their entire lengths.

The left ulna lacks only the most proximal part of the olecranon, while the
right ulna lacks most of the olecranon and the distal end. The latter specimen
apparently suffered an injury or fracture during life, approximately 45 mm
from the distal end. The now missing distal part of this bone does not appear to
have been properly knitted on to the shaft, where it is swollen and the bone is
porous: The fossil specimens have stouter shafts than those of the comparative
specimens, although the distal parts of the more complete specimen is actually
more slender. The semi-lunar notch is smaller in the fossil and in keeping with
the smaller size of the distal articular end of the humerus.

The manus of the fossil is represented by metacarpals II to V of both sides,
the right metacarpal I, left scapho-lunar, pisiform, unciform, magnum and
trapezoid, as well as several phalanges. In addition, a bone tentatively identified
as the left radial sesamoid is known. The fossil carpal bones are all smaller than
their counterparts in H. brunnea, but are otherwise similar in detail.

The radial sesamoids of the comparative specimens were lost, but in
extant hyaenid species it is apparently a small and rather featureless bone.
The articular facet for the radial sesamoid on the scapho-lunars of the H. brunnea
specimens are similar in size to that on the fossil specimen, in spite of the fact
that the latter is a much smaller bone. This suggests that the radial sesamoid
was a relatively more prominent bone in the manus of H. abronia. The fossil
bone tentatively identified as a radial sesamoid is approximately almond-shaped,
with a more or less flat articular facet situated parallel to the long axis at the
broadest end of the bone. It is actually not dissimilar to the vestigial meta-
carpal I of H. brunnea, although it is much larger. It is, however, clearly not a
metacarpal I, since the articular facet is situated laterally rather than proxi-
mally and, in addition, the metacarpal I of H. abronia is known.

The metacarpal I is a significant bone, since its retention in a recognizable
form is a primitive characteristic exhibited by H. abronia. It is a small bone,
about half the length of metacarpal II and resembles that of Proteles cristatus
in morphology. In H. brunnea this bone is much reduced and has no phalanges
associated with it, but in H. abronia at least the 1st phalanx must still have been
present as the metacarpal I has a distal articular facet.

The other metacarpals are shorter and their combined proximal articular
surfaces are smaller than in H. brunnea. The metacarpals II and V are only
slightly shorter than those of H. brunnea and are similarly proportioned, while
the metacarpals III and IV are relatively and absolutely still shorter. Conse-
quently, when the fossil metacarpals are articulated, III and IV do not project
as far forward from II and V as is the case in H. brunnea.

The phalanges of the manus and pes of H. brunnea can be readily differen-

114 ANNALS OF THE SOUTH AFRICAN MUSEUM

tiated on the basis of their size, those of the manus being larger. This size
differentiation is far less obvious in H. abronia, but the phalanges which were
recovered could be separated into two categories. Of the twenty-one phalanges
recovered, thirteen were identified as being from the manus (seven Ist, two
and and four 3rd). They are all smaller than their counterparts in H. brunnea
and the size difference is especially marked in the case of the 3rd phalanges.

Hindlimbs (Table 26)

The greater parts of both innominates were recovered, although the
symphyseal portions are missing and only the ilia and ischia are reasonably
intact. The pelvis is similar in size and general morphology to that of H. brunnea,
although the ischiatic spines are more prominently developed.

Both fossil femora were recovered, but both lack the greater trochanter
and distal ends. ‘They are similar in length to the femur of H. brunnea, they have
slightly stouter shafts, but the heads are a little smaller. The lesser trochanter of
the fossil and the ridges of bone on the shaft where muscles attach are more
prominent. ! |

The left and right tibiae were recovered and both lack their proximal
ends, but in spite of this they are longer or nearly as long as the comparative
specimens. Their estimated total lengths exceed that of the largest of the
HH. brunnea specimens by at least 5%. The shafts are appreciably stouter than
those of the H. brunnea tibiae, but the distal articular ends are similar in size.
This is in very marked contrast to the corresponding bone of the forelimb
(i.e. the radius), in which the distal articular end was much smaller than that
of H. brunnea.

Only the lateral malleoli and most distal parts of the shafts of the fibulae
are known. They are similar to corresponding parts of the fibula of H. brunnea,
but the fossil specimens are broader and what is visible of the shafts more
robust.

The left pes is represented by the calcaneum, astragalus, navicular,
external cuneiform, internal cuneiform and parts of metatarsals III, IV and V.
The right pes is represented by the same tarsal bones, with the cuboid in addi-
tion, and by the complete metatarsals II to V. In addition three Ist, two 2nd
and three 3rd phalanges were recovered.

Whereas the carpal bones of H. abronia were noticeably smaller than their
counterparts in H. brunnea, the tarsal bones are not as simply differentiated.
For example, the fossil and modern astragali are similar in size, the fossil
cuboid is smaller and the fossil internal cuneiform is larger. If any generalization
is warranted then it is that the proximal tarsal bones of the two species are
similar in size, while those tarsals which articulate with the metatarsals are
iarger in H. brunnea, except for the internal cuneiform which is reduced. There
are also some differences in the nature of the articular facets on the individual
bones. For example, that facet on the cuboid which articulates with the external
cuneiform has a figure-of-8 shape, whereas in the H. brunnea specimens it is more

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE LON Els

or less circular and, in addition, this facet is closer to the proximal end of the
bone, whereas in H. brunnea it is nearer to the distal end.

The differences between the fossil and modern metatarsals are much less
marked than those between the metacarpals. The metatarsals III and IV of
the two species are similar in length, but the metatarsals II and V of the fossil
are a little longer than those of H. brunnea. consequently, the articulated fossil
metatarsals take on the same aspect as the metacarpals in that the two central
ones do not project much further forward than those on either side.

The phalanges of the hindfeet of H. abronia are similar in all respects to
those of H. brunnea.

Discussion

Of the extant species of Hyaenidae, H. abronia is undoubtedly most similar
to H. hyaena. ‘The similarity between the latter and another fossil hyaenid from
South Africa, H. makapani Toerien, 1952, led Ewer (1967) to regard it simply
as a sub-species of H. hyaena. It might therefore be expected that H. abronia
and H. h. makapani are related and comparisons between these two fossil forms
are necessary.

Some of the characteristics of H. h. makapani mentioned by Toerien (1952:
294) and which are relevant in the present instance are:

(1) ‘the teeth ... are on the whole a little smaller than the average for (H. hyaena).’

(2) “The anterior cusp of the second premolar is practically absent. Only in one specimen is a
trace of it present. In P? the anterior cusp is not as well developed as in H. Ayaena.’

(3) ‘In P, the anterior cusp is almost absent and in P, it is very feebly developed.’

(4) ‘The anterior cusp of P, .. . (is) not well separated from the central cusp.’

(5) ‘In the lower carnassial the inner cusp (metaconid) is better developed and more distinct
than in H. hyaena.’

All these statements apply equally well to H. abronia and, consequently,
in respect of these characters the Langebaanweg species differs as much from
H. hyaena as H. h. makapani. The decision to distinguish the Langebaanweg form
from H. hyaena at the species level is based principally on the fact that it retains
P,, M, and sometimes also M?. The indications are that H. abronia is simply a
more primitive version of H. h. makapani, which is in keeping with the inferred
relative ages of the two forms, H. abronia dating from the Langebaanian and
H. h. makapani dating from the Makapanian.

H. abronia, H. h. makapani and H. hyaena are here regarded as successive
elements in a single lineage.

In his discussion of relationships within the genus Hyaena, Thenius (1966)
took H. pyrenaica Depéret, 1890 (= H. donnezani Viret, 1954) to be the species
from which all later forms of Hyaena arose. H. pyrenaica is an element of the
French ‘Perpignan fauna’, which is dated as ‘early middle Pliocene’ by Bonifay
(1969). Although the age of the Langebaanweg fauna relative to that from
Perpignan is uncertain, it does seem likely that it is somewhat younger. Conse-
quently, it might be expected that H. abronia was also a descendant of H.
pyrenaica, but this is clearly impossible since in respect of the presence of P,

116 ANNALS OF THE SOUTH AFRICAN MUSEUM

and M,, H. abronia is the more primitive of the two species. The only possible
explanation of this fact is that they are early members of two distinct lineages
(see Fig. 29).

It is now widely accepted that Hyaena stemmed from the Viverra-like
ictitheres of the Pliocene. Kurtén (1971: 143) referred to this ancestral group
as follows:

‘The small Ictithertum was still rather like an overgrown civet, while the
related Palhyaena comprises a number of species ranging in size from a fox
to a wolf. The dentitions of these hyaenids suggest that they were highly
predaceous.’

In view of the age of the Langebaanweg fauna, it might be expected that
H. abronia would hold an approximately intermediate position between the
ictitheres and H. hyaena in terms of morphology, and perhaps also habits. It
does in fact resemble the ictitheres in the number of teeth in its dentition as it
retains P,, M, and M?, but in tooth morphology there is a greater resemblance
to H. hyaena, especially in the development of P?,, P?, and P, into broad crushing
teeth. It is also more like H. hyaena in respect of its overall size.

H. abronia is also intermediate between the ictitheres and extant hyaenids
in respect of the development of its post-cranial skeleton. In his discussion on
European Pontian Ictithertum, Pilgrim (1931) contrasted the limb proportions
of the species he compared to Ictithertum by calculating the ratios between lengths
of metatarsals and metacarpals. The same ratio was calculated for H. abronia
and it was found that this species does indeed fall into an expected position in
relation to Ictitheriwm and advanced hyaenids (Table 20).

TABLE 20

Length ratios of metacarpal IV and metatarsal
IV in some carnivores.

Mc IV : MtIV
Viverra tangalunga . . . AB 2 7
Ictitherium orbignyit . . 1.@ S i
Ictithertum robustum. . . E,20)0 1
Ictithertum hipparionum . . 1,06 : I
Hyaenaabronia . . . . 1,03 : 1
Fijaena hyacna vy 25 0,96 : 1
CHOGHA GOGHIE 2 6 «6 < @,93) 5) 1

Howell (1944) devised a series of indices which are useful in comparing
limb proportions in animal species. The limb segment ratios of H. abronia are,
in general, intermediate between those of a primitive aeluroid carnivore,
represented here by Viverra tangalunga, and advanced hyaenids (Table 21).
There was evidently very little change in the relative sizes of the humerus and

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE Le ad

radius in the H. abronia—H. hyaena lineage, but there was a reduction in the size
of the tibia relative to the femur, as well a reduction in the size of both these
bones relative to those of the forelimb. Consequently, the most marked
differences between H. abronia and H. hyaena are in those ratios which contrast
fore- and hindlimb elements (i.e. the femoro-humeral, tibio-radial and inter-

membral indices).

TABLE 21

Limb segment ratios in some carnivores.

Humero- | Femoro- | Femoro- Tibio- Inter-

n radial humeral tibial radial membral
index index index index index
Viverra tangalunga’ . .. 5 go,I 81,3 96,2 76,1 78,8
Hyaenaabronia. . . I 106,7 88,2 c. 90,9 C. 10355 €. 9555
Hyaenahyaena®. . . 3 106,8 9557 88,9 115,0 104,8
Hyaena brunnea. . . 2 111,8 88,1 81,7 120,5 102,7

1 Davis 1964.
* Kurtén 1956.

Limb segment ratios have also been expressed in other ways and some of
the figures quoted by Pilgrim (1931: 8g) are useful in the present instance, since
Ictitherium robustum is included. This species probably represents, or at least
resembles the archetype of the H. abronia—H. hyaena lineage. Comparisons
between these three species (Table 22) show H. abronia to be more advanced
than J. robustum in the development of both fore- and hindlimbs, while the
H abronia forelimb longbone elements are proportioned in a similar fashion to
those of H. Ayaena and those of the hindlimb are not as advanced.

TABLE 22

Limb segment ratios in some carnivores.

Humerus : Femur :

radius tibia

Viverra tangalungat . . lt i73°1309 £50,907
Ictitherium robustum1 . . I. 1,00 2 €..0,96
Hyaena abronia. . .. ee 0 Ee. 0,01

Hyaena hyaena*® . . . 13, 1,07 I : 0,89

1 Pilgrim 1931.
2 Kurtén 1956.

118 ANNALS OF THE SOUTH AFRICAN MUSEUM

The limb segment ratios of H. brunnea differ somewhat from those of
H. hyaena (Table 21), and this has already been discussed by Kurtén (1956) in
relation to the limb proportions of Crocuta. In H. brunnea the radius is enlarged
relative to the humerus and the tibia reduced relative to the femur. Conse-
quently, while the femoro-humeral index in H. brunnea is similar to that of
H. abronia, the tibio-radial indices or these two species are appreciably different.
Although the intermembral indices of H. brunnea and H. hyaena are similar, the
indications from the other indices are that the two species achieved their
present limb proportions in different ways. On the assumption that the few
specimens whose limb segment ratios have been recorded here are indeed
representative of their species, it follows that H. brunnea and H. hyaena may well
have evolved along different lines for a long time, although ultimately they have
ended up with essentially similar body proportions.

The limb segment ratios already discussed have not taken the extremities
of the limbs into consideration, although some indication of relative sizes is
gained by the figures recorded for the metapodials (Table 20). When the
elements of the manus and pes of H. abronia are articulated, it is evident that
the forefeet were a little broader than the hindfeet. This is, however, largely
due to the presence of metacarpal I and since this digit was either non-functional
or not fully functional, the effective widths of the fore- and hindfeet of H.
abronia must have been much the same. This contrasts with the situation in
modern hyaenids in which the forefeet are broader than the hindfeet, even
though the pollex has been almost completely lost.

In respect of the development of its limbs, H. abronza is clearly much more
advanced than its ictithere ancestor and is closer to, although not quite as
advanced as H. hyaena. The same applies to its skull characters. With its more
or less equally proportioned fore- and hindlimbs and its long tail, it must have
been a less grotesque-looking animal than modern hyaenids, although its
relatively large skull must have made it unmistakably hyaenid in appearance.
It was perhaps more fleet-footed than modern hyaenids and may well have
been more actively predaceous.

Family Hyaenidae
Subfamily Hyaeninae
Hyaena Species B
(Figs 22, 23)

Comment

Sufficient is known of this species to suggest that it belongs to a taxon
distinct from other Langebaanweg Hyaenidae, although there is a possibility
that the two individuals which are represented are large or otherwise aberrant
forms of Hyaena abronia. Owing to this uncertainty the species is not formally

LATE CENOZOICG CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE I19

named, although the description of the material is based on the belief that the
first of the above alternatives is correct.

Material

L 12848— Parts of the skull and skeleton of a single individual, including the
following:
Right maxillary fragment with C, P? and P®; left maxillary fragment
with P3, P* and M!; right mandibular fragment with P, to M,;
left mandibular fragment with P, and P,.
One cervical and one caudal vertebra.
Parts of one ulna, one radius and one tibia.
Twelve carpal and tarsal bones.
Parts of at least seven metapodials.
Five ist, five 2nd and three 3rd phalanges.
Eight sesamoids.
L 11206—Incomplete left M,.

Locality and horizon

These specimens are from Bed 2, ‘E’ Quarry, Langebaanweg.

Description

Although this species is less well represented than H. abronia, sufficient is
known to allow for a fairly detailed analysis of its characteristics. The material
L 12848 represents the remains of a young adult individual in which the per-
manent teeth are all fully erupted and, at most, only slightly worn, but in which
the epiphyses of some elements of the postcranial skeleton are unfused.

Skull (Tables 16, 17)

The skull of L 12848 was badly damaged by a mechanical excavator and,
while some repairs have been possible, it is still largely fragmentary and much
is missing. Most of the teeth recovered were well preserved and intact. Of the
skull itself, only parts of the occipital, the frontals and one nasal are in a condi-
tion which allows for examination of their characteristics.

The occipital condyles, foramen magnum and immediately adjacent parts
are essentially similar to corresponding parts of the skulls of the H. abronia
holotype and the H. hyaena comparative series, although the condyles are more
elongated antero-posteriorly than in either of these species. The frontals are,
as far as can be seen, similar to those of H. hyaena, but they differ from those of
H. abronia in being more constricted in the post-orbital region. The preserved
nasal is similar to those of H. abronia and H. hyaena.

The right mandible has been largely restored, except for parts of the
ascending. ramus which are missing. It is very like that of H. abronia in all
observable respects. The subangular lobe is prominent and the curvature of
the inferior margin posterior to this feature appears to be comparable to that

120 ANNALS OF THE SOUTH AFRICAN MUSEUM

of H. abronia, although the shape of the detached angular process is like that
of H. hyaena. ‘This process is, however, shorter than in the modern species. ‘There
is also a foreshortening in the anterior part of the mandible which is not evident
in either H. abronia or H. hyaena.

This foreshortening of the anterior part of the snout is best illustrated by
reference to tooth row lengths. Although the C to P® of L 12848 are actually
longer than the corresponding teeth in H. abronia, the total length of this part
of the tooth row is actually less in L 12848 (55 mm as against 60 mm). There
is a similar disparity in the corresponding teeth of the mandible, although the
total lengths of the lower cheektooth rows of the two species are almost identical.
The crowding of the anterior cheekteeth of L 12848 is at least in part due to this
foreshortening, although the elongation of these teeth is another contributing
factor.

It is the elongation of the cheekteeth and the fact that they are higher
crowned that most readily distinguishes L 12848 from H. abronia. In general,
the teeth are also longer than those of H. hyaena, but they are similar in breadth
and crown height. 7

The upper canines of L 12848 and H. hyaena are similar in size. They differ
in that the C of the fossil does not have the distinct V-shaped projection from
the anterior cingulum. The C of H. abronia is similar to that of L 12848 in this
respect.

The P! of L 12848 is lost, but judging from the size of its alveolus, it was
similar in size to that of H. hyaena and larger than that of H. abronia. It was
tightly sandwiched between the C and P?.

Only the right P? of L 12848 was recovered and this tooth is damaged. It is
larger than that of H. abronia in all respects and apparently had a small but
distinct anterior ancessory cusp. It differs from the P? of both H. abronia and
H. hyaena in having the inflation of the internal cingulum more anteriorly
situated.

Morphologically the P? of L 12848 resembles that of H. abronia more closely
than that of H. hyaena, since the anterior accessory cusp is smaller and the
posterior accessory cusp is larger than in the modern species. It is, however,
more comparable in size to the P® of H. Ayaena.

As with P8, the P* is morphologically more like that of H. abronia, but is
closer to that of H. hyaena in size. Both protocone and parastyle are reduced as
in H. abronia. The metastyle is more strongly arched posteriorly than in H.
hyaena, so at least in early wear the posterior part did not function as a shearing
surface.

There are marked differences in the nature of the upper molars of the
three species. The M! of Species B is appreciably larger than that of the H.
abronia holotype and larger still than those of the H. hyaena comparative speci-
mens. The posterior part of the maxilla of L 12848 is known from the left side
only and although M? is lost, its alveolus is preserved. This tooth had at least
two roots and was much larger than the single M? of the H. abronia holotype,

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE a2

FULL AHLHL LI LU LFV LLL LIL
fe ils. 14 1S Ne. aly

|

ja UU LLULLU LULL
11 112 113 114 1

il
5

Fig. 22. Buccal and occlusal views of the Hyaena sp. B maxillary fragments (L 12848) from
Langebaanweg.

122 ANNALS OF THE SOUTH AFRICAN MUSEUM

UAC ulti mn ul

8

ALL ALLL Ann UD) ATA INTIAL U UU TU

Fig. 23. Occlusal and buccal views of the Hyaena sp. B mandible (L 12848) from Langebaanweg.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 123

which is only single-rooted. The upper molars of L 12848 also differ from those
of H. abronia in their positions relative to P+. In the latter species the buccal
surfaces of M! and M? make an angle of approximately 110° with the long axis
of P4, but in L 12848 this angle is about 100°, so that in lateral view the molars
are, or were just visible behind P*. Species B is intermediate between H. abronia
and H. hyaena in this respect.

The P, of Species B is a little larger than that of H. abronia, but is otherwise
similar. The other lower pre-molars are all more elongated and higher crowned
than those of H. abronia, while in the P, the anterior accessory cusp is more
clearly separated from the principal cusp.

Except for their larger size, the M, of L 12848 and the damaged specimen
L 11206 are similar to that of H. abronia. The M, of L 12848 is damaged, but it
also appears to have been similar to that of H. abronia.

Postcranial skeleton (Tables 23, 25, 26)

Little of the postcranial skeleton of L 12848 was recovered, but there is
sufficient to indicate that this individual was a little larger than the H. abronia
holotype, even though the animal itself was younger. There is also sufficient
to suggest that Species B, like H. abronia, did not have the enlarged forelimbs
characteristic of more advanced hyaenids.

Vertebrae

The preserved cervical vertebra, probably the 5th, is only a little smaller
than those of the two available H. brunnea skeletons, while the caudal vertebra,
probably the 1oth or 11th, is appreciably longer than those of the comparative
specimens. The latter is comparable to the 11th caudal vertebra of H. abronia,
which indicates that Species B also had a relatively long tail. A few isolated
epiphyses of vertebrae were also found.

Forelimb (Table 25)

The only significant observable feature in the fragmentary radius and ulna
which were recovered, is that in the former the distal end is transversely
expanded in a manner which is more comparable to the condition in the avail-
able H. brunnea specimens than in H. abronia. However, the distal epiphysis of
the radius is lost and the distal end of the diaphysis is not fully ossified, so the
character of this area of the radius may be due to the relative youth of the
individual. This region of the radius of L 12848 actually has a greater transverse
diameter than the distal end of the radius of the H. abronia holotype.

Corresponding to the enlarged distal end of the radius, the preserved
elements of the manus of L 12848 are also large compared to those of H. abronia.
The scapho-lunar and trapezoid are intermediate in size between those of
H. abronia and the H. brunnea comparative specimens. The cuneiform of H.
abronia is not known, but that of L 12848 actually has a greater transverse
diameter than those of the H. brunnea specimens and, as it is not as high, the

I24 ANNALS OF THE SOUTH AFRICAN MUSEUM

transverse elongation is very pronounced. ‘The transverse diameter of the
trapezium is also greater than in the H. brunnea specimens and the various
articular facets on this bone are larger as well. The size of the distal facet
indicates that the metacarpal I must have been a fairly substantial bone,
perhaps being proportionately as large as those of H. abronia and Percrocuta
australis.

The left metacarpal II and right metacarpal V are preserved intact,
while the right metacarpal IV is represented only by the proximal half. They
are comparable in size to those of H. brunnea, except that the two complete
specimens are slightly shorter.

It is clear from the transverse dimensions of the various elements of the
manus that the forefeet of Species B were broader than those of the H. brunnea
comparative specimens, although in overall size they were not as large.

Hindlimb (Table 26)

Too little of the tibia of L 12848 is preserved to allow useful comments on
its characteristics to be made.

Of the pes, the astragalus, navicular, all three cuneiforms, and parts of
metatarsals III, IV and V are known. These bones are all a little larger than
their counterparts in the H. abronia holotype and the two modern H. brunnea
skeletons.

Phalanges of both manus and pes of L 12848 are apparently represented.
The relative sizes differ to approximately the same extent evident in H. abronza.

Discussion

The overall impression gained from the comparisons between L 12848
and the holotype of H. abronia is that the two forms are little different, but that
such differences as do exist are probably greater than would be expected of two
individuals of the same species. They are here regarded as being specifically
distinct and as having had a common ancestor not far removed from them in
time. In respect of both size and the nature of the dentition, H. abronia is the
species likely to have borne a closer resemblance to the hypothetical common
ancestor, while Species B is the more specialized of the two. They are, however,
equally primitive in that both retain P, and M3.

The specimens L 12848 and L 14186 were found about 100 metres apart
at about the same level in Bed 2 and near each were the remains of a comparable
array of other vertebrate species, so that there can be little doubt that they were
contemporaries. The co-existence of two hyaenid species of comparable size
may have been made possible by their occupying slightly different ecological
niches. This possibility is lent some support by the nature of the dentition of
the two forms, since the more slender and high crowned teeth of Species B
suggest that it might have been an early member of a lineage in which the
trend was towards more highly predaceous forms.

Since the forefeet of Species B are a little smaller than those of H. brunnea

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE P25

and the hindfeet a little larger, there was clearly not the disproportionate
development of the forefeet that is evident in modern hyaenids and in this
respect Species B resembles H. abronia. ‘The general similarity in the size of
fore- and hindlimbs in Species B cannot, however, be demonstrated in the same
way as it was in H. abronia. Some additional indication of limb proportions was
obtained by comparing the sizes of the proximal facets of the scapho-lunar
and astragalus of the three species (Table 23). The size of these facets is a
reflection of the size of the radius and tibia and the fossil species are closer to
one another in this respect than either is to H. brunnea.

TABLE 23

The dimensions of the scapho-lunar and astragalus of some hyaenid species.

(1) (2)

Greatest trans. diam. | Greatest trans. diam.

of radial facet of of tibial facet of (1) : (2)

scapholunar astragalus
Hyaena brunna(n = 2) . . . 26,30 18,55 i: 0,71
Gijaraaoronia . . . ».» « « 23,10 18,00 1: 0,78
hgammrpectes B 2 -. «le 25,20 20,10 I : 0,80

The possibility does exist that L 12848, and the isolated carnassial L 11206,
belong to large individuals of the species H. abronia and that the elongation of
the teeth merely represent individual peculiarities. The situation would no
doubt be resolved by the recovery of more specimens and since material is still
being collected from Bed 2 in ‘E’ Quarry, there is a good chance that this might
happen. Consequently, although the opinion expressed here is that the present
material is specifically distinct from H. abronia, and apparently other previously
recorded hyaenid species, some additional comment on this matter will have to
be made in the future.

Family Hyaenidae
Subfamily Hyaeninae
Hyaenictis preforfex n. sp.
(Fig. 24)
Holotype
L 10055 and associated pieces—The skull and parts of the skeleton of a single
individual, including:
Skull lacking parts of the braincase, palate, left nasal, right I’ and
left: Pt.
Left and right mandibles lacking I,, P,, M, and I, I,, Py, Me
respectively.

126 ANNALS OF THE SOUTH AFRICAN MUSEUM

Several incomplete vertebrae.

Parts of both humeri, both radii, one ulna and both tibiae.
Five carpal and tarsal bones. |
Parts of at least seven metapodials.

Seven Ist, two 2nd and four grd phalanges.

Locality and horizon

These specimens are from Bed ga, ‘E’ Quarry, Langebaanweg.

Diagnosis

A small species of Hyaenictis; post-orbital processes long; nasals short,
terminating anterior to the orbits; mandible long and slender; P, and M,
present; P§ and P, lack anterior accessory cusps; P,; broader than P,; M!
large.

Limbs slender and relatively long; fore- and hindlimbs more or less equally
proportioned.

Etymology

From pre meaning ‘before’ and forfex, the specific name of an hyaenid
from Swartkrans in the Transvaal (Ewer 19552).

Description

The holotype of this species is the incomplete skeleton of a very aged
individual in which the teeth are extremely worn. The skull was largely intact
when discovered, but was badly broken with bone fragments being held
together in a sandy matrix. Attempts to restore the specimen have been made
by several persons and it has been adversely affected by this attention. It is now
in two parts, the face being detached from the braincase. Most of the post-
cranial remains have suffered some post-mortem damage.

Skull (Tables 16, 17)

Owing to distortion of the face and braincase, the two parts of the skull
can no longer be fitted together, but most of the skull characters can still be
observed. The braincase lacks most of the occiput, the basi-cranium and the
posterior portion of the sagittal crest. Apart from its smaller size, the braincase
resembles those of H. abronia and the H. hyaena comparative specimens. The
frontals show signs of disease or injury and are otherwise remarkable for the
very elongated post-orbital projections. The facial portion of the skull is more
complete than the braincase and the only really important part which is missing
is the posterior palatal region. It differs from corresponding parts of the skulls
of H. abronia and H. hyaena in its smaller size and the relatively shorter nasals,
which terminate anterior to the orbits.

The mandible is long and slender and although it is not quite as elongated
as those of available Canis lupus skulls, it is otherwise not dissimilar in proportions.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 127

The subangular lobe is not as prominent as that in H. abronia and the inferior
margin below the ascending ramus is more like that in H. hyaena, although the
angular process is not as long. The condyle is not as high relative to the cheek-
teeth as it is in H. hyaena, but it is higher than in H. abronia.

Although the remaining teeth of L 10055 are very worn and many of their
characteristics are no longer evident, it is quite clear that they differ significantly
from the teeth of other hyaenid species from Langebaanweg. As with H.
abronia and Species B, P, and M, were still present in this species and in this
respect it is equally primitive. On the other hand, P$, P3 and P, are relatively
broader than those of the two Hyaena species from ‘E’ Quarry. In this respect

in

Fig. 24. Occlusal and buccal views of the Ayaenictis preforfex mandible (L 10056) from
Langebaanweg.

128 ANNALS OF THE SOUTH AFRICAN MUSEUM

the present species is the more highly specialized. The P3 and P, lack anterior
accessory cusps and the anterior keels of their principal cusps are not as promi-
nent as in H. abronia and Species B. The broadening of the anterior cheekteeth is
particularly noticeable in P? and P, and the latter is actually a little broader
than P,. It is also relatively broader than the P, of H. hyaena.

The carnassials and post-carnassial teeth are either very worn, damaged
or lost. The P* protocone seems to have been fairly prominent and situated at
right angles to the parastyle. The M1 was apparently relatively large and at least
as broad as that of Species B. In view of the relative sizes of these two species,
the size of M! relative to the other cheekteeth must have been greater in the
present species than in the Hyaena species from Langebaanweg. Since the
posterior part of the palate is damaged, it is not known whether or not M?
was present. The lower carnassials are very worn, but the proportions of these
teeth appear to have been similar to those of the H. abronia lower carnassials.
It is not known whether or not the M, metaconid was present. The Mg, like
P,, has been lost from both halves of the mandible, but whereas P, was lost
during life, the loss of M, was post-mortem. Both these teeth were small and
single-rooted.

Postcranial skeleton (Tables 24, 25, 26)

The postcranial skeleton of this species is less well known than that of
H. abronia, but is better represented than that of Species B.

Vertebrae (Table 24)

The vertebrae recovered are, for the most part, very fragmentary. The
cervical, thoracic and lumbar regions of the spine are represented and the most
complete specimen is the 7th cervical. It is similar in morphology to that of
H. abronia, but is appreciably smaller and is thus in keeping with the smaller
skull size of the present species. It is a great deal smaller than the 7th cervical
vertebrae of the H. brunnea comparative specimens and, although the other
vertebral fragments are not identified as to their actual position in the spine,
they too are all smaller than the vertebrae of H. brunnea.

Forelimb (Table 25)

The left humerus is largely intact and although only a little shorter than
that of H. abronia, it is much more slender. ‘The supratrochlea foramen is,
however, larger than that of the H. abronia humerus.

The proximal end and shaft of the left radius and the distal end and part
of the shaft of the right radius are preserved. Once again the estimated
overall length of the bone is a little less than that of the radius of H. abronza,
but it is much more slender. The same apparently applies in the case of
the ulna, of which only the proximal part of the one from the right side is
preserved.

The pes is represented by a scapho-lunar, both unciforms, a trapezoid,

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 129

parts of both metacarpals I] and III and part of one metacarpal IV. The
carpal bones are similar in size to the corresponding bones in H. abronia and
the metacarpals differ only in that they are more slender.

One of the 1st phalanges is much reduced in size and almost certainly
belonged to the pollex, although a metacarpal I was not found. This bone
may well have been similar to the metacarpals I of Percrocuta australis and
Hyaena abrona and not reduced as in modern Crocuta and Hyaena.

The right metacarpals II and III and another of the 1st phalanges show
signs of a severe pathological condition, the cause of which is not known,
although it could well be connected to, or aggravated by the advanced age of
the individual.

Hindlimb (Table 26)

The hindlimbs are less well represented. The tibia and metatarsal IV are
approximately the same length as those of H. abronia. The only tarsal bones
recovered were an incomplete calcaneum and navicular and both are smaller
than the corresponding bones of H. abronia.

Discussion

In determining the affinities of this species, one of the most obvious possi-
bilities to be considered is that group of Hyaenidae which are referred to the
genus Hyaenictis Gaudry, 1861. Pilgrim (1931: 101) defined this genus as
follows:

‘Hyaenidae with moderately long, slender, rather shallow mandible and
(by inference) with somewhat elongated facial region; M? large, triangular,
almost at right angles to P*; M? absent; P* with long posterior lobe and large
protocone; M, short, hardly longer than P,, without metaconid; M, small;
P, present but with a tendency to be deciduous; P, with large posterior and
small anterior cusps, P,; with large posterior and small anterior cusps; P,
with large anterior and posterior cusps.’

This definition is based on the type species, H. graeca, and it could, there-
fore, be modified to accommodate more advanced, but phylogenetically directly
related forms, which postdate the Pikermi species. One such species is ‘Hyaena’
bosei, which is probably from the Pinjor stage of the Siwaliks, and which was
included in AHyaenictis by Pilgrim (1932). This later species differs from H. graeca
in that its lower premolars lack anterior accessory cusps and P, is absent.

The Langebaanweg deposits postdate those at Pikermi, but are earlier
than those of the Pinjor, and it might therefore be expected that if L 10055 is
an Hyaenictis, it would be intermediate in character between H. graeca and H.
bose. This does in fact appear to be the case, although some of the features which
characterize Hyaenictis, such as the absence of M? and presence of M, metaconid,
are not evident in L 10055. H. graeca has anterior accessory cusps on the lower
premolars, they are lacking in H. bosei, while in L 10055 they are absent in

130 ANNALS OF THE SOUTH AFRICAN MUSEUM

P, and P;. The first lower premolars of L 10055 were lost during life and this
is taken as an indication that they had a ‘tendency to be deciduous’ as in
H. graeca. ‘They were, however, completely absent in H. bose.

The Langebaanweg species also shares at least two striking characteristics
with H. bosez. namely, very long post-orbital processes and nasals terminating
anterior to the orbits (see Pilgrim 1932).

Although there is some justification for referring L 10055 to Hyaenictis,
there is at least one other possibility to be considered. In her description of the
Hyaenidae from Swartkrans, Ewer (1955a) assigned one of the species which had
previously been referred to Hyaenictis to the genus Leecyaena Young & Liu, 1948.
The reasons she dismissed an association between the Transvaal species and
Hyaenictis were as follows:

(1) The Swartkrans species has a mandible which is heavy and fairly deep below Mj.

) The M, has a large metaconid. (It is, however, relatively small compared to that of H.
hyaena and other species).
(3) The Mt? is broad from side to side, but short antero-posteriorly.
(4) ‘The P* protocone is small.

) The skull does not have long post-orbital processes or nasals terminating anterior to the
orbits.

Since the Swartkrans species is much more recent than the: Pontian
H. graeca, most of these factors could be dismissed on the grounds that the former
is a more advanced species, but one which could nevertheless have derived
from H. graeca. The most telling point against this interpretation is the fact
that the Swartkrans species has an M, metaconid whereas H. graeca does not.
However, even this does not totally preclude a relationship since Kurtén
(1963a) has demonstrated that a ‘lost’ feature such as an M, metaconid can be
regained in the course of the evolution of a single lineage.

One possible interpretation of the facts is that the Swartkrans ‘Leecyaena’
forfex and the small Langebaanweg hyaenid are members of an African Hyaenictis
lineage, or else were members of a lineage which paralleled but was indepen-
dent of the Eurasian Hyaenictis. Before examining this possibility, it is necessary
to examine the decision to refer the Swartkrans species to Leecyaena.

Subsequent to the description of ‘L.’ forfex, Ewer (1967) expressed the
opinion that it and the only other recorded species of Leecyaena (i.e. L. lycyaenoides
from the late Pliocene of China) were probably not closely related. It seems
likely that in view of their relative ages and the nature of their specialized
characters (Ewer 1955a) the two species belong in separate lineages, so
there is a real doubt that the Swartkrans species is a Leecyaena.

‘L.’ forfex differs from the small Langebaanweg hyaenid in the following
respects:

) ‘L. forfex is a little larger.

) The post-orbital processes are smaller and the nasals terminate more posteriorly.
) The premaxilla projects further forward.

) The premolars are broader.

) The mandible is more heavily built.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 131

Since the Swartkrans fauna is younger than that from Langebaanweg, it
follows that ‘L.’ forfex must be the more advanced of the two species if they are
indeed on the same lineage. In fact, all the observed differences between the two
species could be accounted for in this manner. There was a trend in hyaenids
towards increasing size; the nature of the post-orbital processes and nasals in
the Langebaanweg species is shared by at least one species of Jctztherium (Pilgrim
1932: 123), and they could therefore be regarded as primitive characteristics;

p2 2 ie

p3

,06 0 ,06
Fig. 25. Ratio diagram comparing relative lengths of the cheekteeth of Hyaenictis preforfex (2)
and H. forfex (3) (Ewer 1955a), with modern Hyaena hyaena (1) (Kurtén 1956) as a standard.

132 ANNALS OF THE SOUTH AFRICAN MUSEUM

the more prominent premaxilla of ‘L.’ forfex is coupled with a longer pre-canine
diastema and this in turn is determined by the larger size of the canines relative
to those of L 10055; there was a trend in most hyaenid lineages for the pre-
molars to broaden into crushing teeth; and the larger mandible of ‘L.’ forfex
is in keeping with the larger overall size of the species, coupled with the general
enlargement of the cheekteeth (Fig. 25). The fact that the premolars of L 10055
are relatively broader than those of other contemporary hyaenids from Lange-
baanweg, indicates that it did indeed belong in a lineage where there was a
trend for broadening of these teeth.

Perhaps even more significant than accounting for the differences between
these two species are the characters which they have in common. In both
species P§ and P, lack anterior accessory cusps, Ps is broader than P, and the
premolars are generally comparabie in morphology. Both species retain M,
and in both the M!? is large. The wear on the teeth indicates that they shared at
least one important functional adaptation. The P, of L 10055 is extensively
worn on its postero-lateral surface and is thus similar to that of ‘L.’ forfex which
‘is not a fully specialized exclusively crushing tooth, but still retains the shearing
action typical of normal carnivore premolar occlusion’ (Ewer 19554: 820).
In this respect the two species differ from H. hyaena and H. brunnea and parallel
Crocuta. j

The geographical proximity of the two species increases the likelihood that
they are phyletically connected.

It is concluded that the Langebaanweg and Swartkrans species were
members of the same lineage and are, therefore, congeneric. On balance it
seems probable that their affinities lie with Hyaenictis and they are here referred
to this genus. The specific name of the Langebaanweg Hyaenictis preforfex is given
in the belief that it was ancestral to Hyaenictis forfex.

Since these two species are now included in Ayaenictis, the following
revised statement concerning the genus is made:

Genus Hyaenictis Gaudry, 1861
Type species: Hyaenictis graeca Gaudry, 1861.

Diagnosis: Hyaenidae of small to medium size; M! large, almost at right
angles to P*; M? absent; P, sometimes present, but with a tendency
to be shed; anterior accessory cusps of P} and P3 present only
in the earliest species; M, metaconid small or absent; M, small;
mandible long and slender in early species, but more robust in later
forms.

Stratigraphic range: Mid Pliocene to early Pleistocene of Eurasia and
Africa.

Referred species: Hyaenictis bosei Matthew, 1929; Hyaenictis forfex Ewer, 1955;
Hyaenictis preforfex, new species.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 133

The postcranial skeleton of H. preforfex is of interest since comparisons
with that of the contemporaneous Hyaena abronia show that these two hyaenids
were very different in their body proportions. The indications are that H.
preforfex was only a little shorter at the shoulder and that the two had hindlimbs
of comparable length. The limbs of the Hyaenictis were, however, much more
slender and the head and neck less enlarged relative to the rest of the body. The
implications are that it was more cursorial than the contemporary species of
Hyaena. This is of interest because of the suggestion that the long-limbed
‘hunting hyaena’, Euryboas, was descended from Hyaenictis graeca (‘Thenius
1966). Pliocene Hyaenictis may thus have been a group of long-limbed forms
which evolved in two directions, one branch becoming increasingly cursorial
and actively predaceous (Euryboas), and the other paralleling Hyaena and
Crocuta (H. boser, H. forfex, H. preforfex).

TABLE 24

Dimensions of the vertebrae of Hyaena abronia and Hyaenictis preforfex from Langebaanweg,
compared with those of modern Hyaena brunnea

Hyaena brunnea
Hyaena Hyaenictis. -—_——_—_—_-
7th cervical vertebra abronia preforfex SAM SAM

L14186 L10055 36150 17238

Length of centrum. . 3555 31,0 37,7 40,9
Length from anterior eanaeanae to penieaioe

zygopophysis. . . 4353 36,4 4555 46,0

Transverse diameter of eae ates aS, 24,3 20,0 27,8 27,8

Width across anterior zygopophyses . . . 49,8 42,9 60,2 60,1

Hyaena brunnea

Sacrum Hyaena = |———
abronia SAM SAM
L14186 36150 17238
Antero-posterior diameter of centrum . 55,0 5455 52,5

13th 14th 15th
21,1! 19,3 17,7
16,1 14,7 13,5
31 31 31

Caudal vertebrae (lengths)

Hyaena abronia L14186
Hyaena brunnea SAM vias
Difference in %

Q‘II 6‘o1 — O01 ‘9 — — : Be ral * - JaJQUIVIP 1O1I19}sod-o19}uUe WINUIIXey

oL1 L‘gt — 0°61 —- — : : : 3 : IO}OUILIP ISIDASULI] WINWIxe/y wioyiauND
L‘&o Q°ss G‘Q1 — 6°L1 — : ; : * JOJOUIVIP 10119}Ssod-o19}Ue WINUIIXe|Y

9‘z& 9‘s& £°gz 19a o‘Q& — cae ok j ? ; JO}OUIVIP SIDASUeI] UINUIIxePY reuny-oydvog

SIvduvo

oS BGS = com 3°ES oo ; yo}OU JeUN]-TUWs Je I9}9UIVIP 1O1I19}Ssod-o19j7e WINUITUITY
GLE L‘g& 062 °9 — L‘GE — : ; ; sso001d pIlouo109 }e JOJ9UIeIP 1OT19}sod-o19}uy
o‘L 0g es =e 6‘o1 — ; ; ; ; ; ; ; JeYS JO IOJOUILIP ISIOASULI}] UINUITUITY
g‘ol 9°61 = = o‘OI = ; ; : : : ‘ —-«4yeys Jo Joj}0UIeIp 1O1I9}sod-o19}Ue WINUITUTTY
26 i) — — 36 — paar Neer ssoo0id proyAjs Jo JOJDWILIP ISIOASUI] WINUIIXe/]
ov oVI — = 6°11 — ° : : " — ssa00id aspieess ye AOI CUIELD JO1I9}SsOd-019}Ue WINUIIXe|Y
oSbz 0*Egz — — o°L&s °9 — ; ; ; ; : : * YSUI] [[e12AOC

3 VN1N

A Son Lit G‘g =e Le = ; ; 2 ; : * -44geys JO Jo}0WILIP 1O119}3sod-o19} Ue WUNUITUTY\Y

~ 1‘QI V‘LI 6°S1 — g‘9I — : : : : ; YeYS JO IOJDUIIP ISIOASULI] WUNUITUTIAY

= Per Gee EVI — o‘GI — : : : * -puo yeurrxoid 3e soJ9WIeIp 1O119}sod-019}Ue WUINUIxeyy

4 Qvz Gz 1°13 ‘9 — Gas = ; ; : : puo jewrxoid je JOJOUIVIP ISIOASULI] UINUIIXeyy

S Vso 6°Zs = — £°0% — 2 : : ‘ pud [eISIP 1 JOJOWIeIP IOTIO}sod-o19}Ue WINUIIXeY/[

me L‘9& 6°96 — — S1E —- ; F : : pue ae ye aoe? ISIDASULI] UINUITXRIY

< o°Gia 0°2&s o°G61 9 — o‘Loz — ° ; p : ; ° YISUg] [[VIIAO

im snlava

=) L‘o% 6S VG — 6‘g1 — ; ; * -yyeys Jo JoJOWeIp 1O1I9}sod-o19}Ue WINUWITUIYY

g ‘V1 Vol 9°61 — 6‘91 — : : ; : : : YeYS JO IOJOWILIP ISIZASULIY WUNUTTUTPAY

6‘9& g‘6E = — L‘G& — : : ; * - pud [eISIP 78 IOJOUIVIP IO1I9}sOd-o19}Ue WINWIIXe]]

= 6°LY o‘6V — — o‘Qv -—- 3 ; 3 : ; pUd [e}SIP Je IOJOWILIP ISIOASULI] WINUWIXe]]

fx o‘F61 0°90% o*Ggr °9 —- oV61 — : SoorJIne Te|noae [ane Puy jewrxoid us9mj0q YISUST

S 0‘90% 0°0%% — — 0‘Z0% — : ; ; i : ; YASUI] [[V19ACE

4 snuawnH

s — EL — — E°VG — : ; ; ; : * ¥BSSOJ PIOUI]S JO JOJOUIVIP IsIDASULL TY,

4 L‘9é ILE — — G°1E — ; ; * eSSO} PIOUITS JO JajOWIeIP 10119}sod-019]uy
£96 g‘gé — — a‘GE — ee hss a a RS yoou jo JaJaWIeIp Io110}sod-o19}0V
fey QGP — -— v‘ov — : puo yewrxoid je 19j9UIvIpP 10119}sod O19}Ue 3s91¥915)

VINdvos
oS19& goat CSoo1'y Qbgsiy 9givi'y Soe 17
‘WV'S WY'S xaftofasg | g sotsadg DIuUoLGD SsUp4sgsnD
——___—_—_'"——__—_|_ syaiuavdyy puavdpy puavd yy DINIOLILIT

pauunig Duavd pz]

re

“pauUunsg DUaDdFT JO ISOY} YIIM poreduiod ‘oepruseAPT SOMULEqISUL'T JO S}USUII[O QUIT[II10F JO SUOISUSTAIC

Gz aIav

135

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

puo

puo

puo

puo

pus

puo

pue [PISIp Je JOJDUILIP 9s19ASURI] WINUITXe YY
* pua [eIsIp ye JOJOUIeIp JOL19}sod-o19}ue UUNUUIXe Py
pue jeurxoid je JaJ9UIvIp ds19ASURI) WUNUUIXe Py
Lee 7e DIS wETP Jo1i9}sod-o19}ue WNUWIxepy
YIsuy]| [[V19AG

pUd [P}SIP 1¥ JOJ9UILIp dsIDASUPI} UNUTXe Yl

* pUd [e}SIp }v Jo}OWIeIp 10119}s0d-o1a}UL LUNUIIxe Ay
pus jeutxoid je Ja}9WIeIpP asIdAsURI) UINUIXe Py
et eee. ye si i Jo1i9}sod-o19}ue WMNUIxXe yy
YIBug] [[V19AO

pue [eIsIp }e JoJOWILIP osIOASURI} LUNUIXe YY

[eisip ye JoJOUIvIP AOLIO}sod-o19jUe UMUIIxey
pus jeutxoid je J9j9WIeIp asI9ASURI} UINUIIXe Ay

a a sisiaahsinl' Jo1i1ajsod-o19jue WNUITxe yy
YIBUI] [[V19AO

pUd [PISIP }e JOJIUILIP 9sIDASUBIY LUNWIxe yy

pue [eIsIp Je JOJOUNIvIP 1O1I9}sod-o19;Ue WINUIXe Py
pus jeurxoid je Ja}9UIeIp os19ASURI) UMUITxe yy
Nad aga 1 eee Jo1i94sod-o19}ue WNUWIxeyy
YIBuy] [[V19AO

pus [vISIP Je J919WIVIpP osI9ASURI] LUNWTxe py

pus [eIsIp Je J9}9UIeIp JO119}sod-o19}ue WNUIIxey
pus jeurxoid je JoJoWIeIp ss19AsUeI) UMUIXey
et ca his aslaaia td Joli9jsod-o19}ue WMNUIxe yy
YISUI] [[e1IVAG

pie Pea }@ IBJOWUIVIP ISIOASUL J,
: YISUg]| [[e19AO

pe Monnens }@ JOJDUIVIP IsIDASULIT,
: YIBUy] [[e1I9AO

JIIUILIP IsIZASUBIY WINUIIXe yy
Ja}oUWIvIp 1o119}sod-o19}ue WNUIxe py
JOJIUILIP ISIOASUBI} LUNWIIXe I
JaJ2UIvIp 1O1I9}sod-o19}ue WINUITxe Py
JI}IUIVIP ISIDASULI] WUNWIXe
Jg}9UIvIp JO119}sod-o19}ue WINWIxe Py
JO}DUILIP ISIOASULI} WLINWIxePy

AW

AI °W

TIT eW

II °W

I°W

SIVduvOv.Lay
prouresas

[PrP eid

Weer
winizode. J,

prozodeiy,
wnuseryyy

ULIOJIOUA)
(panuzjuo2) SIVauvo

ANNALS OF THE SOUTH AFRICAN MUSEUM

136

ee ey ee

ocL1 — ULI 1‘QI 1% hee cal ang: IQ}DUILIP ISIOASULI] WINUITXe yy
00% == L‘0% 30S 6‘0& : : ; * - JQJOUIVIP IOTI9}sod-o19]Ue WINUIIXL{y Je[NIIACNY
90% = G°ss °9 3°OS So : : S : : JOIUIVIP ISIDASULI] LUNUITXe{Y
oS == — v‘QI oa : g : 5 BEL OUEED JO11I9}sod-o19]Uv UNUIIXeyy
76% — ra 26% — ee te : So bees * YSU [[212AO snjeseljsy
O'S == == SIS aa : : ; : IQJIUILIP ISIDASULI] WINUITXeIY
61% = aaa CO aan ae TS 6 poe JO119}sod-o019]Ue WINUIIxeyy
o‘LV — — 067 — : : ; ; : : * y)8ug] [[V190A—C unsure
STVSUVL
g‘6 = — S01 = : g : ; g PpUd [eISIP JO IoJOWIVIp IsIDASULI} UINUWIXe/y
o‘VI — — 6°E1 — pud [eISIp Jo JoJOWIVIp Io110}sod-o19}Ue WINUIIxey]
vi1nla
6°G1 L‘S1 — o‘QI — Se ee ere ey es YeyYs JO IO}DUILIP IsIOASULI LINUTTUTYY
g‘VI avi — QV = i ; ; : : " —-:YyeYS JO 1OJOUIVIP IOTIO}sOd-o19}Ue WUNUITUT]Y
v6 o‘LZ — o ‘6 =e toll «ony Sl, PpUd [eISIP JO 19N}9UILIP IsIOASURI] LUNUWUIXe/\]
G°Gs 6‘gI — L°Ss os pue (ane ue zOLOUIENP IOII9}SsOd-019}Ue WINUWITXe J]
0261 0‘00% °9 — 0‘00G ‘2 — : : : ; : ; : * YSUIT [[V19AO
VIGIL
o‘gI — — oS == Cl yap le a er yeYs JO IoJOWILIP osIZASURI] LMUUTUTY
EG — — o‘QI aa ; q ° 3 ; ‘ —-« 4yyeys Jo JoJOUILIpP JoTIo}sod-o19}Ue WINUIIUITY
ove — — Qs — yt peay JO JojOWIeIp 1o0110}sod-o19]Ue LUNUWIIxe jy
G‘1z — — G1z — Fe Gis wats go ee ee Bey i) fou IEP) [eUdA-OsIOp WUNWITxeyy
o°&&% — — 0°0ZG °9 — ; parties's 6 rae is : YISUIT [[V19AO
wows
oLa — — 9‘Gz — ys te uNge}99e Jo JOJOWIVIP JOII9}sod-o19}UYy
Q'S — —— ov1 — wINn[Nge}I9e 0} JOIIN}SOd UIMIYISI JO JOJOUILIP [VIJUDA-OSIOP UUINUWITUIT]
5‘Q& — — o‘Lz -— * + UUNNGe}99e 0} IOTIOJUL UINI][I JO JOJOUIVIP [eIJUDA-OSIOp UINUTUTT
ALVNIWONNI

gtzLi GGSoor1'T gbgzi'y QogItiy ES0E1'T

WVvs xaflofarg | g sotoadg DIU0LGD SYDAASND

—| siaiuandpy puavapy puavdpey DINIOLILIT

DaUUNLG DUADAET

“pauunsg DuavA fF] JO 2sOYy} YM poreduiod ‘ovepruseAP] SamuLveqosuv’y] JO S}UIUIZ[9 QUIT[pUTY 24} Jo SUOTSUSWIC]

Q% A1av J,

LJ)

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

PUd [VISIP JO JOJIUILIP ISIOASULAY LUNUIXe IY
* pud [ISIp Jo Ja]9WeIp 10L19}s0d-o19}Ue WINUWIIXe |

pua jeurxoid jo 19} 9WUIvIpP ISIDASURI} LUNWIXL

ae a a i ED st -O19}UR WINUIXR |]
yisug] [[219AQ

pUd [BISIP JO 19] 9UIVIP ISADASUBAY WINWITXV YY

pus [ejsip jo sa}0WIvIp sOl9}sod-o19}7UR WINUIxe py
pus jeurxo.id jo IIJIWIPIP ISIOASUPBI} WUNUITXP TAT

ide ord p SSASEREP Jo1i9sod-o19jue UWINUIXeyyy
YISUy] [[P19AG)

pUd [BISIP JO J9}OUILIP Is1DASUBIY LUNUUIXe Ay

pua |eISIp Jo JoJOWUIeIp s10119}sod-o19}Ue WUNWIxe yy

pus jewrxoid jo 19}9WIeIp IsIDASURI) WUUNWUIXe yy

pa oi cataae Ja spel 3 Jo1iojsod-o19jue WNUWIxepy
yisuyg] [[e19AO

puod [eISIp JO 19}9UIeIp IsIDASURIY WINUWIxe yy

pu? [vISIp JO 19}9WIvIpP 10119}sod-o19]Ue WINUIIxe py

pus [eutxoid jo 1J9}9UIvIp dsI9ASURIy WNUUTxe yy

se eae jo ciahia Jo1iaysod-o19jue WNUIXe)y
YIBUy] [[V19AO

saieue ty Jo1i9}sod-o19}Ue WNUIIXe yy
YIBUY] [[V1IAO

JIIUIVIP ISIDASUBI} LINUITXL IAT
zaigtae yy Jo1193sod-o19}ue WNUWIxeyy
; Yyi3ug] [[219AO

JQJIUILIP ISIDASULIY WNUWIIXe IY
mune y Jo1i9}sod-o19jue WNUIIXe yy
YyIBuy] [[219AOC

I9JIUIVIP ISIDASULI} WINUWITXeYJ
TSIgInEIP Jo1io}sod-o19}Ue WNUIXe fy
ys8ug] [[219AO

A IW

AIAN

Ill IW

II WN

STVSUVLV.LAW
uI1OJIaUNd
jeuso}uyT

uLIOJIOUND

S1PPIN

wWOFIOUNS
[euro XY

peers ba)
(panu1juor) STVSAVL

138 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Hyaenidae
Subfamily Hyaeninae
Species E
(Fig. 26)
Material
L 2673—Right mandibular fragment with P, and P3.

Locality and horizon
This specimen is from ‘E’ Quarry, Langebaanweg, and is probably from
Bed ga.

Description

This fragmentary specimen belongs to an immature individual of a species
which is comparable in size to H. abronia. The permanent canine is unerupted
and is visible through damaged portions of the alveoli of the third incisor and

wut

Sa me
AKA se SS

int PUTA LLUA LLP LAE LL

Fig. 26. Occlusal and buccal views of the hyaenid species E
(L 2673) from Langebaanweg.

LATE CENOZOICGC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

139
13 P3
O
12 Hyaena_ hyaena
OO
<=
=
o 11 @ Species E
{=e}
e
Species B
10

@ Hyaena abronia

@Hyaenictis preforfex

Length

18 19 20 21 22 23 24
Length

Fig. 27a. Dimensions of the P, and M, of Hyaenidae from Langebaanweg, compared with those
of a series of modern Hyaena hyaena.

140 ANNALS OF THE SOUTH AFRICAN MUSEUM

18 1
m @
Species B
7
16

@
Hyaena abronia

5 5 7 8 9 10 11
Length

Fig. 27b. Dimensions of the M! of Hyaenidae from Langebaanweg, compared with those of a
series of modern Hyaena hyaena.

deciduous canine. There are no indications that P, is, or was ever present. The
P, and P, are only partly erupted. The mandible is broken posterior to P,
and the nature of the other lower cheekteeth is unknown.

The P,, which measures 12,9 by 8,8 mm, differs from those of other
Langebaanweg Hyaenidae in that there is no trace of an anterior accessory cusp
and no swelling of the cingulum at the anterior end of the tooth. In addition,
although a posterior accessory cusp is present, it is much smaller than those of
the other species. The tooth has prominent anterior and posterior keels and
it is relatively broader than those of other Langebaanweg hyaenids. Both buccal
and lingual cingular margins are markedly convex, in contrast to the other
species in which these margins are more or less straight.

The P, (c 16,5 x 11,0) is morphologically very similar to P,, but is larger
and relatively broader. The posterior accessory cusp is almost identical in size

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE I4I

6
32 4 2 ii :
ge
7
Fk
4
p3 +CESTIMATE
> ~~
SS
pane
>
we
06 0 06 12
~- +

Fig. 28. Ratio diagram comparing relative lengths of the cheekteeth of Hyaenidae from
Langebaanweg (Hyaena abronia (2), Species B (3), Hyaenictis preforfex (4), Species E (5), Per-
crocuta australis (6)), with modern Hyaena hAyaena (1) (Kurtén 1956) as a standard.

to that of P, and is thus a relatively smaller and less significant feature of the
tooth.

Discussion

The lack of P, and the broader, almost conical P, and P3, which lack
anterior accessory cusps and have very small posterior accessory cusps, readily
distinguishes L 2673 from the other Langebaanweg hyaenids as well as other
species previously recorded from South Africa. Its characteristics cannot be
attributed to the ontogenetic age of the individual, since other specimens from
Langebaanweg which belong to individuals of comparable age (e.g. 5/1966/1,
vide infra) have unerupted cheekteeth which are morphologically comparable
to those of adults.

142 ANNALS OF THE SOUTH AFRICAN MUSEUM

Since this species is so poorly represented, and since its affinities were not
determined, it is not classified below the subfamily level. For the purposes of
convenience it is designated Langebaanweg hyaenid ‘Species E’.

Family Hyaenidae
Subfamily Hyaeninae
Incertae sedis
(Hendey, 1970a: pl. 2A, B)
Material

(1) The following specimens are from Bed 2, ‘E’ Quarry:
Lg139— Right premaxilla with 1! to I and associated left C.
L 12499/500—Right I? and M?.
L 12850— Mandibular fragments with right C and left P,.
L 12868— Mandibular fragments with incomplete P,.
L 13042—Left P,.
(2) The following specimens are from Bed ga, ‘E’ Quarry:
(i) Excavation LBW 1966/1:
5/1966/1 and associated pieces—Parts of the skull and postcranial
skeleton of a single individual, including:
Right and left maxillary fragments with Q@ and P? to Ml.
Right mandibular fragment with P, and M,; left mandi-
bular fragment with C and P3.
Parts of one humerus and one ulna.
Two tarsal bones.
Parts of four metapodials.
One ist phalanx.
(i) Excavation LBW 1966/2:
L 10566 to L 10569—P*, M}, P! and I?.
L 10802 to L 10804 and L 10806 to L 10808 — M1, M,, Ps, P,, P, and I?.
(ii) Excavations LBW 1969/1 and LBW 1970/1:
A series of teeth probably belonging to a single individual:
L 15742—Right dp*, P? and P*.
L 15797A, L 15592, L 15896A, L 15897—Right C, P3, Py, M,.
I 15610, 2 1580608; L 15808) 15715 — Lett dp, Ps aren
Other isolated teeth:
L 16055K—Right M!.
L 15824 —Left P*.
L 15588B/A — Left I°.
J, TRG, 1h, meee, Ib nSayOlB, 1b nGAGO—IRiclet (C, Jos, Pa, (Pn
L 16055B, L 15854, L 16246—Left P, (two), M,.
(3) The following specimens are from ‘E’ Quarry, provenance unknown:
L 2055 —Left mandibular fragment with P,.
L 5731—Right mandibular fragment with ?P,.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 143

L 6378—Right mandibular fragment with parts of P, to M,.

L 6379—Right mandibular fragment with P, to M,.

L 6380—Left mandibular fragment with M, and M,.

L 10746 to L 10748—Premolars.

L 11822, L 11891, L 12101, L 12423B—Canines.

L 2054—Part of M,.

Seven premolar fragments, one maxillary fragment and two mandibular
fragments.

Discussion

Owing to certain similarities between the material assigned to Hyaena
abronia, Species B and Hyaenictis preforfex, and to the poor definition of the
characters of Species E, difficulty was experienced in identifying the material
listed above, although it almost certainly does belong to one or other of these
species.

Bed 2 material (Tables 27, 28)

Those specimens from Bed 2 could belong to either H. abronia or Species B.
For example, although the P,, L 13042, is very similar to that of the H. abronia
holotype, it is an unerupted tooth belonging to a very young individual and,
when fully developed, might conceivably have taken on the characters of the
P, of Species B. It would serve no useful purpose at present to make tentative
identifications of the Bed 2 specimens.

Bed 3a material (Tables 27, 28)

The material recovered from excavations in Bed ga is more important
since a large number of specimens, some relatively well represented, are
involved. Probably most of the specimens should be referred to H. abronia
(a Bed 2 species), but at least one exception is the P,, L 16055B, which is
virtually indistinguishable from the P, of Hyaenictis preforfex (a Bed 3a species).
Many of the unclassified Bed 3a specimens belong to immature individuals
and although in size such teeth are equally close to those of the H. abronia and
HM. preforfex holotypes, they are morphologically more similar to the former.
Some of the isolated teeth other than P, and P, could perhaps belong
to Species E.

The best represented of the unclassified Bed 3a specimens is the series
from the excavation LBW 1966/1. These represent the remains of an immature
individual of about the same ontogenetic age as the specimen referred to
Species E. The deciduous dentition is not represented, but only the molars
are fully erupted. The epiphyses of the preserved elements of the postcranial
skeleton were not yet fused. The teeth of this individual are similar enough in
size and morphology to those of the H. abronia holotype to suggest that the two
are conspecific, although there is a somewhat ambiguous situation in respect
of the M? of these two individuals.

144 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 27

Dimensions of unclassified hyaenid upper teeth from Langebaanweg.

Ee pe [pe M!
] b ] b ] b ] b
Fy MOO Se Geen. Ilias G20 TSU RISING OQ. T5.e leone 13,0
Dv5624 = 8 —}|— — | 27,7. 15,6 | — —
[eNO507 5 = — — — — — — | 7,3 12,7
Lrosee . se | —}— — — — | 7,6 1257
Li6o55K. . .| — = haa — — — | 7,6 14,2
ILMARGCY 5 a. is — — — — a — | 85 15,6
TABLE 28

Dimensions of unclassified hyaenid lower teeth from Langebaanweg.

1 iP M,

] b ] b l b
EyCCOMM i sae « 17 /e Q,2 17,9 9,4. 20,2 Q,2
L10803/6/4 oe 16,2 953 17,8 9,6 19,1 955
E6970) eee ee a iGatOss OSS Gar 7-3 c.057 19,8 9,8
TSG 025. 1 4 17.0 Q,1 — — — —
TEE 7G ase Ges ae NOS 2.) mali — — — —
IeVSO42 eee FST, MOO — — — —
n6o46/60° > > = — — GUOD Oe 21,9 10,6
LvGS5450 3 ee — a 19,0 ion a= ==
Lir6055Bs 4s 9 = = 1755 9.5 ri oe
J FARES SIO\ SYA 3) be — — 18,4 9,3 —_ —
boone hia er: — — 18,6 10,6 — =
TET5GOT =. oa Mx = = = ist 21,6 935

TEQU Seat) nt eek ee — — — — 21,8 -LOs5

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 145

The M!?! of 5/1966/1 is virtually identical to two other specimens from
Bed ga (L 10567, L 10802), while another (L 16055K) is more or less inter-
mediate in size and morphology between that of 5/1966/1 and the M! of the
H. abronia holotype. The M? of the H. abronia holotype is present on the right
side only and is a small, single-rooted tooth. In 5/1966/1 it was present on
both sides and these teeth, which are unfortunately lost, had at least two, and
possibly three roots. Since Bed ga is later than Bed 2, the situation in respect
of the M? of 5/1966/1 and L 14186 is the reverse of what might have been
expected. If these two specimens are indeed conspecific, the explanation may
simply be that the M?, although becoming vestigial, was still very variably
developed in the species during that period in time involved in the deposition
of Bed 2 and Bed ga. The significance attached to the differences in the M? of
the Viverra leakeyi from these beds (vide supra) might therefore be unwarranted.

It is tentatively concluded that the unclassified Bed 3a hyaenid material
represents at least two species and that these are probably Hyaena abronia and
Hyaenictis preforfex.

Unprovenanced material (Tables 27, 28)

The unprovenanced material from ‘E’ Quarry apparently also includes
specimens belonging to more than one species.

Judging from the preservation of the mandibular fragments and the dates
of their discovery, these specimens are most likely to have come from Bed ga.
Consequently, they too might be expected to represent H. abronia and H. pre-
Sorfex, while in some cases at least, Species E is another possibility.

An example of the difficulties encountered in identifying this material is
offered by the mandibular fragments L 2055, L 6378 and L 6380. These speci-
mens belonged to adult individuals and are, therefore, comparable to the
holotypes of H. abronia and H. preforfex in this respect. L 6380 is much more
robustly developed than the mandible of the former, but the differences are
such that they might be accounted for by individual intraspecific variation.
This would be still easier to accept if the Bed 3a H. abronia was a larger variety
of the species than that represented in Bed 2. The M, and M, of L 6380 are very
worn, but they are clearly appreciably smaller than the corresponding teeth of
Hyaena Species B. Comparisons with Species E are completely inconclusive
owing to the nature of the single specimen referred to this species.

The problem becomes more complex when L 6378 is taken into considera-
tion. This specimen is still larger than L 6380 and it seems unlikely to be
conspecific with H. abronia because of its size, although on the basis of the
fragmentary teeth of L 6378, this possibility cannot be excluded. This would be
the most reasonable identification if the Bed 3a variety of H. abronia was larger
than that from Bed 2. In the case of L 6378, the nature of the cheekteeth pre-
cludes the possibility of it being identified with Species B or Species E.

The mandibular fragment L 2055 compounds the uncertainty because in
this instance the corpus is actually a little more slender than that of the H.

146 ANNALS OF THE SOUTH AFRICAN MUSEUM

abronia holotype, although it is still larger than that of the H. preforfex holotype.
It is almost inconceivable that L 2055 and L 6378 could be conspecific, unless
there was in this species a far greater size range of variation than those observed
in modern hyaenid species in the course of the present study.

This appears to be yet another situation which will only be satisfactorily
resolved by the recovery of additional material and there seems little point
in making provisional identifications of the unprovenanced material at present.

GENERAL DISCUSSION ON THE LANGEBAANWEG HYAENIDAE

On the basis of the specimens presently available, the following grouping
of the Langebaanweg Hyaenidae is proposed:

SPECIES SIZE PROVENANCE
Percrocuta australis Large Bed 2
Hyaena abronia Medium Bed 2
Hyaena Species B Medium Bed 2
Hyaena cf. abronia Medium Bed 3a
Species E Medium ?Bed ga
Hyaenictis preforfex Small Bed 3a

The unclassified material probably belongs to one or other of the listed species.

Although there are obvious differences between the various species (e.g. see
Figs 27, 28), the presence of unclassified material raises the question of whether
or not the various species which have been named are adequately defined. There
is no real problem with Percrocuta australis as the large size and specialized
dentition of this species makes it easy to recognize. The difficulties le with
the medium- and small-sized species, which have rather generalized, primitive
hyaenid dentitions.

On purely theoretical grounds it might have been predicted that it would
be difficult to classify the Langebaanweg hyaenids, since this family was
diversifying during the late Pliocene and early members of new lineages would
exhibit only slight differences from the archetypes. By the Pleistocene those
hyaenids which are known from South Africa were well advanced on their
particular lineages and, consequently, they can be more readily distinguished
from one another.

The evolutionary history of Hyaena and Hyaenictis as it relates to species
recorded from South Africa (Fig. 29) is visualized as follows:

(1) The late Pliocene H. abronia from Langebaanweg and the mid Pliocene H. pyrenaica from
Europe are regarded as the earliest recorded representatives of the H. hyaena and H. brunnea
lineages respectively.

The H. hyaena lineage, which includes H. h. makapani from the Transvaal, was a conservative
one which underwent comparatively little change from the late Pliocene onwards. The
differentiation of this lineage might well have taken place in Africa.

An ancestor of H. brunnea probably entered Africa from the north during the Pliocene. It is
first-recorded in South Africa during the Makapanian (Swartkrans) and it apparently
replaced H. hyaena, which is last recorded earlier in the age (Makapansgat). AH. bellax,
another Makapanian species, is probably an off-shoot of the H. brunnea lineage.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 147

Species B probably arose from an immediate ancestor of H. abronia. The problematical
H. namaquensis from Kleinzee (see Ewer 1967) may be related to Species B.

Another lineage which must have had its beginnings during the Pliocene is that which
includes Hyaenictis preforfex and H. forfex. It is not known to have survived the Makapanian.

— —
IS<) i)
wa pe)

(4) Species E is a problematical form whose affinities are not known.
Lu
Zz
O
fe) Hyaena hyaena Hyaena brunnea
O
ac
Hyaena hyaena

Fy prisca
Ze
UO Hyaena
O brevirostris
ES Hyaenictis Hyaena
io forfex bellax
=

Hyaenictis Hyaena hyaena Hyaena

bosei makabani perrieri

Hyaenictis Hyaena Hyaena
preforfex Sp. B abronia

Lu Hyaena
z pyrenaica
U
0
x Hyaenictis Ictitherium, Palhyaena

graeca

Fig. 29. Tentative phylogeny of some Hyaenidae.

The recorded late Pliocene Hyaenidae of South Africa thus include one
species belonging to a lineage which is still extant (H. abronia), one species which
has a Pleistocene descendant (fH. preforfex), two which apparently became
extinct without issue (Hyaena Species B and Percrocuta australis), and two of
uncertain affinities (7. namaquensis and Species E).

This apparently complex association of hyaenids is not unique. Kurtén
(1953) has investigated Chinese Pontian hyaenids and at a single occurrence
(Loc. 49) the following species were recorded:

Ictithertum gaudryi, I. ‘sinense’, I. wongii, I. hyaenoides, ? Lycyaena dubia and Crocuta
varvabilis.

Kurtén’s (1953) reassessment of this material led to the conclusion -that
I. ‘sinense’ is probably a slightly aberrant J. gaudryi and ? L. dubia is an aberrant
I. hyaenoides. C. variabilis was subsequently referred to Percrocuta (Kurtén 19570).
The Loc. 49 hyaenids thus comprise a Percrocuta and three species of Ictitherium,

148 ANNALS OF THE SOUTH AFRICAN MUSEUM

the latter including two aberrant specimens previously referred to other species.
This is a remarkable match for the Percrocuta, three smaller hyaenid species and
two problematical specimens from the South African Langebaanian. It is
probably expecting too much to suppose that H. namaquensis and Species E
can be disposed of as satisfactorily as the Loc. 49 I. ‘sinense’ and ? L. dubia, but
the important point is that there are precedents for an association of hyaenids
such as that recorded for the Langebaanian and at Langebaanweg in particular.
While this does not in itself validate the conclusions reached here, it does mean
that the variety of species represented at Langebaanweg is not unexpected.

Although it is usually accepted that Hyaena arose from the ictitheres during
the Pliocene, there is as yet no unanimity as to which, if any, of the recorded
species is likely to have been the actual ancestor. Thenius (1966) concluded that
Hyaena was derived from I. robustum, while Kurtén (1971: 144) states that, “The
modern genus (Hyaena) presumably evolved from a Palhyaena ancestor’. By
Palhyaena he presumably means ‘J.’ hipparionum of the European Pontian and
‘I. wongi of the Chinese Pontian.

In an earlier study on the ictitheres, Kurtén (1954) tentatively concluded
that the recorded species could be grouped as follows:
(1) JI. wongi, I. hipparionum
(2) J. robustum, I. gaudryi, I. tauricum
(3) JI. sivalense, I. hyaenoides, ?I. indicum

(Not mentioned: J. orbignyi, a small species occupying a somewhat isolated position.)

Kurtén has thus apparently favoured derivation of Hyaena from group (1),
while Thenius thought the genus derived from group (2). Perhaps both were
right and Hyaena is polyphyletic rather than monophyletic in origin. It has
already been suggested that the early ancestors of H. hyaena and H. brunnea
were differentiated by the mid Pliocene and that the former may have arisen
in Africa and the latter in Europe, or at least Eurasia. The mid Pliocene was
also the time when the ictitheres were at the peak of their radiation and,
although they have yet to be recorded in Africa, it is probably safe to assume
that they were present on this continent as well. Consequently, there does
seem to be a reasonable possibility that H. hyaena and H. brunnea did evolve
from two different ictithere species. The common ancestor of the two species
may thus date back to the Miocene, and the characteristics which they share
might be the result of parallel evolution rather than a close phyletic relationship.

In recognition of their long independent history, it is proposed that the
H. hyaena and H. brunnea lineages be differentiated by giving their members
separate subgeneric status.

Subgenus Hyaena
Type species: Hyaena hyaena Linnaeus, 1758.

Diagnosis: WHyaenidae usually of medium size; P, and M$ retained in late
Pliocene forms; anterior premolars only moderately enlarged even
the most advanced forms; M, metaconid large.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 149

Stratigraphic range: Mid Pliocene to Holocene of Africa and Eurasia.

Referred species: Hyaena abronia (H. makapan Toerien and H. prisca De Serres
included as subspecies of H. hyaena)

Subgenus Parahyaena

Type species: Hyaena brunnea 'Thunberg, 1820.

Diagnosis: Wyaenidae of medium to large size; P,; and M$ lost by the late
Pliocene; anterior premolars much enlarged in the Pleistocene
species; M, metaconid often small or absent.

Stratigraphic range: Mid Pliocene to Holocene of Eurasia and Africa.

Referred species: Hyaena bellax Ewer; H. brevirostris Aymard; H. perrieri Croizet &
Jobert; H. pyrenaica Depéret.

Family Felidae
Subfamily Machairodontinae
Machairodus sp.
(Figs 30, 31, 32, 38)
Material
L 20505—Parts of the skull of a single individual, including: Isolated ? I’
or 2: right maxillary fragment with C and P® to M}; left maxillary
fragment with P? and P*. Right mandibular fragment with part of
M,; left mandibular fragment with M,.

Tentatively referred material

L 11890—? Left I°.
L 6386 —Right mandibular fragment.
L 12641—Right mandibular fragment with incomplete P, and M,.

Locality and horizon

These specimens are from Bed 2, ‘E’ Quarry, Langebaanweg.

Description

The fragmentary skull (L 20505) belongs to a relatively primitive and
moderately large machairodont, the primitive character being suggested
principally by the presence of P?. Both the left and the right P? of this specimen
have been lost, but their alveoli are preserved. The left alveolus (4 x 4 mm)
indicates that the tooth was small, circular and single-rooted, while the right
P? was evidently a little larger, antero-posteriorly elongated and double-rooted,
the alveolus measuring 6 x 4,5 mm.

The isolated incisor of L 20505 is a worn and rather nondescript tooth
and only because of its direct association with the remainder of the skull is it
identified with this species.

150 ANNALS OF THE SOUTH AFRICAN MUSEUM

The right C is largely intact, except for some slight damage to the root.
It is a relatively large tooth (Table 29), the overall length along the anterior
curve behind about 120 mm. The root and crown make up approximately
equal parts of the tooth. The root is somewhat bulbous and its contact with the
neck is therefore clearly defined. The crown is transversely compressed and
has a diametrical index of only 0,46. There are prominent anterior and posterior
keels and these are finely serrated. The anterior serrations are less distinct and,
unlike those on the posterior keel, they are absent for about 3 mm from the
apex of the tooth. The posterior keel is rectilinear, but the anterior one curves
lingually towards the base of the crown.

TABLE 29
Dimensions of the upper teeth of Machairodontinae from Langebaanweg, compared

with those of a specimen from Makapansgat.

Langebaanweg

Makapansgat
Machairodus cf. Machairodus | cf. Homotherium | Homotherium
L20505 L118g90 M8280*
1 c. 6,2 — —

ae: or 2

trans. diam.
Mt!
Ant-post-diam.

C-P4 |
* Collings 1972.
The right P? is damaged, but the left is intact, except for the apex of the

principal cusp which is lost. It has prominent anterior and posterior accessory
cusps, the latter being the larger of the two, while there is an additional small

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE I5I!

|

5
4

qn HUTT MULVEY LL LL LL LL
11 112 113 114 1S 116 117 118 119 210

Sit Z\l gil Sit r\I £\I raul Tl oll 6
PATA TTT in nH utl

Fig. 30a. Buccal and occlusal views of the right maxilla (L 20505) of the Machairodus
sp. from Langebaanweg.

152 ANNALS OF THE SOUTH AFRICAN MUSEUM

AUT rm TULLE HLUHLL LIL LLL nt

Fig. 30b. Buccal and occlusal views of the left maxilla (L 20505) of the
Machairodus sp. from Langebaanweg.

cusp situated at the posterior end of the tooth. This cusp is in close contact with
P4 and the pressure brought about by this contact has resulted in damage to its
enamel surface. The same applies in the case of the right P®. These teeth are
relatively long and narrow.

Both the left and the right P* are complete. They are also long and narrow,
with the protocones much reduced in size. They have prominent ectoparastyles,
which are, however, much smaller than the parastyles. The ectoparastyle,
parastyle and paracone of the left P* are about 20,5 mm in length, while the
metastyle, which is the longest of the cusps, measures 16,5 mm. The roots

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 153

|

(HH

z
mT

I

un

i

nn

8
unui

1

L

1

a

ni nt

i

Mm

nu
8 113

yn

II

NHI

mM

ll

qa HIN iil

nt

HU

nm

mm
2 1

nm
3 1

INU
4 1

mm
5 1

Fig. 31. Buccal, occlusal and posterior views of the Machairodus sp. mandible (L 20505) from
Langebaanweg.

supporting the protocones are anteriorly directed, their dorsoventral axes
being in line with the apices of the paracones. Although the C and P3 of this
specimen show no obvious signs of wear, the shearing surfaces of the P? para-
cones and metastyles are heavily worn, while the protocones have been all but
worn away. The lingual surfaces of the parastyles and ectoparastyles are also
slightly worn.

The preserved right M! is also worn and little of the crown remains. It is
single-rooted, but has a slight transverse elongation. The alveolus of the missing
left M! shows vestiges of the more primitive double-rooted condition.

Little remains of the skull itself and the dorsal edges of the preserved parts
are heavily abraded. The largest piece of the skull which is preserved consists
of that part of the basi-cranium which includes the glenoid regions and the
intervening basi-sphenoid. The glenoid fossae are approximately 13 mm below
the level of the basi-sphenoid, the relative positions of these features being
characteristic of the machairodonts. In the modern feline skulls used for com-
parative purposes during the present study the glenoid fossae and basi-sphenoids

154 ANNALS OF THE SOUTH AFRICAN MUSEUM

were all more or less in the same planes. This characteristic of L 20505 is
also illustrated by the fact that the inferior margin of the left post-glenoid
process is about 25 mm below the level of the inferior margin of the external
auditory meatus. In a young adult lioness skull of similar overall size (SAM
35115), the corresponding figure is about 15 mm.

Both halves of the mandible of L 20505 lack the dorsal margins of the
coronoid processes and the corpora anterior to M,. The characteristics of the
fossil are clearly illustrated by comparisons with corresponding parts of the
mandible of the modern lioness referred to above. The two specimens are
similar in overall size (Table 30), but there are marked differences in individual
characteristics. In the fossil the angular region is directed buccally, whereas in

TABLE 30

Dimensions of the lower teeth and mandibles of Machairodontinae from Langebaanweg,
compared with those of a young adult Panthera leo.

cf. Machairodus Panthera

Machairodus_ |———————_—_,—____— leo 2
L20505 L12641 L6386 SAM35115
l — 12,3 6, 1250 —
P, alveolus
2 ai 595 = _
l = 20,5 os is
P, alveolus
b is 935 ae ay
] — c. 28,0 — —
M, alveolus
b — II,O — —
] Cc. 29,0 — — —
M,
b 11,2 — — —
C-M, ] — c. 104,0 — 98,0
C-P, diastema . -- 627.0 C. 30,0 15,0
Height of corpus anterior to P; : — G2 33,0 C5 37,0 40,4
Breadth of corpus anterior to P,;_ . —— 20,0 22,0 18,4
Height of corpus below M,. . . 36,5 3559 — 3553
Breadth of corpus below M, . . 17,0 18,6 — 19,4
Trans. diameter of condyle. . . 40,3 — — 43,0

DO el

Distance between posterior ends of
condvicyandeNijy im eee 66,0 -— — 73,0

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 155

SAM 35115 it is directed posteriorly. The most striking differences are in the
coronoid processes. Although both these processes are damaged in the fossil,
it is evident from the configuration of the masseteric fossae that little of the most
dorsal parts is in fact missing and, as is characteristic of all machairodonts,
the processes are low relative to those of the lioness and, indeed, other
felines as well. In addition, the anterior margin of the ascending ramus
is more steeply inclined and the coronoid processes are broader antero-
posteriorly.

The left M, is intact and the tooth is relatively long and narrow. Super-
ficially it appears to consist only of a protoconid and paraconid, the former
being about 20% longer than the latter. The cingulum of the most posterior
part of the tooth is slightly bulbous, its outline in occlusal view not conforming
with those parts anterior to it, while there is a barely discernable notch on the
posterior keel dorsal to this region. The keel posterior to this notch and dorsal
to the bulbous cingulum is very finely serrated, the serrations being smaller
than those of the C. This region of the tooth may in fact be the talonid which is
in the phyletic process of being incorporated with the protoconid. Both the
protoconid and the paraconid have well-developed shearing facets worn on
them and there is an additional shear facet resulting from occlusion with M1
worn on the buccal surface along the contact between the ‘talonid’ and proto-
conid. The carnassial notch is fairly prominent and there is a slight groove on
the buccal surface ventral to it which extends almost to the base of the
crown.

The right M, has lost the paraconid and supporting root. In this tooth the
‘talonid’ is not readily demarcated since the serrations and notch separating it
from the protoconid are absent. The transverse fracturing of this tooth at the
carnassial notch has revealed the presence of an additional root beneath the
protoconid. This root is relatively small and is situated on the buccal side
towards the midpoint of the tooth. It has a slight antero-buccal inclination and
serves to anchor the M, very firmly in the mandible.

The isolated ? I® (L 11890) and the mandibular fragments (L 6386,
L 12641) are only tentatively grouped with L 20505, since the nature of the
specimens precludes conclusive comparisons.

The crown of the incisor consists of a high, rather conical cusp, the sides
of which are keeled and serrated, the latter characteristic suggesting its machairo-
dont affinities. ‘There are two small projections from the cingulum situated just
posterior to the keels. It is a slightly larger tooth than the isolated incisor of
L 20505.

The more complete of the two mandibular fragments (L 12641) lacks the
ascending ramus and angle and all the teeth are lost or damaged. The corpus
is long and low, but is relatively broad. The inferior margin below the cheekteeth
is rectilinear. The symphyseal region is high and broad and there is a small
mental flange (crest), which arises from below the larger and most anterior of
the two mental foramina.

ANNALS OF THE SOUTH AFRICAN MUSEUM

156

4

|

oy

mm

‘somuveqosury] wo (1Fgs1 J) s[qrpueur snpcuvysvpy °Jo 9y} Jo MaIA Jeoong ‘ZF “S1q

iii

4S

I van

nl

mn

eI

nil

cll

mui

im

Ul

itt

8

i

i

Z

ut

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE r5/

The dorsal surface of the symphyseal region is damaged, but parts of the
alveoli of I, and C are preserved. These two teeth were antero-posteriorly
elongated and although their dimensions cannot be accurately measured, it is
evident that the C was appreciably larger than I. It is also a little more pos-
teriorly situated. Since their alveoli are not visible, it is probable that the I,
and IJ, were much smaller than I,. The alveolar margins of the symphyseal teeth
must have been well above the level of those of the cheekteeth.

The postcanine diastema is long and at about its midpoint is a small
alveolus (2,0 x 1,5 mm), which contains the root of a vestigial premolar. The
P, has been broken off at the roots, but it was clearly a much smaller tooth than
P,. The P, itself is badly damaged, but it was apparently made up of a principal
cusp, anterior and posterior accessory cusps and a smaller cusp projecting from
the posterior cingulum. Of the M, only the anterior root and part of the crown
which it supports is preserved. In addition to the post-mortem damage, this
tooth was reduced to its present state by heavy wear during life. It lacks the
additional small root beneath the protoconid which is visible in the right M,
of L 20505.

In as far as comparisons between L 12641 and the mandibular fragments
of L 20505 are possible, it appears that the specimens are essentially similar,
although L 12641 has a broader corpus. This may, however, be accounted for
by the fact that it belonged to a more aged individual, a judgment which is
based on the more advanced wear on its M,. A further indication that L 12641
belongs to a machairodont which was similar in size to L 20505 is the fact that
the right maxilla of the latter specimen matches the referred mandible
reasonably well. If the two specimens are indeed conspecific, then the C of
this form must have projected outside the buccal cavity when the mouth was
closed.

The second mandible fragment (L 6386) lacks part of the symphyseal
region and those parts posterior to P,. The damaged roots of the C and P,
are still in place, while the lingual surface of the P, alveolus is still visible. This
specimen is generally similar to corresponding parts of L 12641, although it is
more robust, has larger mental foramina and does not have a vestigial premolar
anterior to P,. There is a very marked contrast in the relative robustness of the
mandibular corpora of L 6386 and L 20505 and, although this does not neces-
sarily preclude their being conspecific, it is possible that L 6386 and perhaps
also L 12641 belong to a larger species. Such a species is recorded from Bed ga

(vide infra).

Discussion

There has been a considerable amount of confusion in the past concerning
the taxonomy of the Machairodontinae and only five genera are recognized
here. They are Homotherium, Dinobastis, Megantereon and Smilodon, which were
all essentially confined to the Pleistocene, and the Pliocene genus Machairodus
(see Kurtén 1963); Thenius 1967).

158 ANNALS OF THE SOUTH AFRICAN MUSEUM

The following characteristics were included in a diagnosis of Machairodus
Kaup, 1833 by Pilgrim (1931: 128): |

‘Machairodontinae of large size . . . ; upper canine very large, stout, long,
flattened; . . . mandible with deep symphysis; canines and incisors elevated
considerably above the level of the back teeth, without mental process but |
with a prominent mental crest; ... P* with protocone weak; paracone and
parastyle well developed; with an extra cusp anterior to the parastyle
(ectoparastyle) ;... M, with vestigial metaconid;...’

If the cusp referred to by Pilgrim as the ‘metaconid’ is the same as that
tentatively identified here as the ‘talonid’, then the Bed 2 machairodont
material is accommodated quite wel! by the above diagnosis.

In addition, the retention of P? in L 20505, and the relatively large size of
P,, the vestigial anterior premolar and the inferred relative sizes of the symphy-
seal teeth in L 12641, are all primitive characters in the Machairodontinae
which are probably sufficient to distinguish the species from those referred to
the four essentially Pleistocene machairodont genera.

The Bed 2 machairodont is therefore referred to the genus Machairodus,
but its specific identity is uncertain. The C of L 20505 (24,2 x 11,1 mm) 1s
similar in size and other respects to that of the Makapanian Machairodus
transvaalensis from Bolt’s Workings at Sterkfontein (24,5 x 12,1 mm) (Broom
1939). However, the isolated upper carnassial referred to this species by Broom
has a much larger parastyle and shorter metastyle than those from Langebaan-
weg. There is no certainty that the Bolt’s Workings P* and C do belong to the
same species, but if this is so then the Langebaanweg Machairodus is clearly not
M. transvaalensis, although it might be an ancestor of this species.

There are no other named machairodonts from South Africa to which the
present material can be referred and comparisons to previously recorded
Eurasian species were inconclusive. Since it may prove to be conspecific with
one of the latter, it is for the present not identified at the species level.

Family Felidae
Subfamily Machairodontinae

cf. Homotherium sp.
(Fig. 38)
Material
L 11846—Incomplete left C

Locality and horizon

This specimen is from Bed ga, ‘E’ Quarry, Langebaanweg.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 159

Description

This canine, which belongs to a machairodont of large size, has lost part
of the crown, but it has an estimated length along the anterior curve of about
200 mm, which is 80 mm longer than the C of the Bed 2 Machairodus. Even if the
length of the restored part of the crown has been exaggerated, the overall
length is most unlikely to have been less than 180 mm and the tooth is still
appreciably larger than that of the Machairodus (Table 29). It is also very
narrow and has a diametrical index of only 0,39. The anterior and posterior
keels of the crown are serrated, the posterior serrations extending about 35 mm
closer towards the root than those on the anterior keel. It differs from the
canine of the Bed 2 species in this respect, since the anterior and posterior
serrations of the C of L 20505 terminate in about the same horizontal plane. The
canines of the two species also differ in that both keels of L 11846 are rectilinear,
whereas the anterior keel of L 20505 curves lingually towards the base. As a
result the anterior serrations of L 11846 are all visible in buccal view, while in the
Machairodus they are all visible only in anterior and lingual view. There is no
appreciable thickening of the root of L 11846 as is the case with L 20505.

Discussion

This specimen was previously tentatively referred to Machairodus (Hendey
1970a), but this identification is almost certainly incorrect.

Recently Collings (1972) described a machairodont from Makapansgat
under the name of ‘Megantereon problematicus’ and the right C of the holotype is
similar in size to L 11846 (Table 29), and also has serrated keels. The Lange-
baanweg and Makapansgat species are probably congeneric and perhaps even
conspecific, although the nature of the relationship cannot be concluded on the
basis of the material presently available. In addition, the identity of ‘Megantereon
problematicus’ is open to question. The relatively short, serrated canines and the
size of the teeth suggest that its affinities lie with Homotherium rather than
Megantereon. Collings made no comparisons with previously recorded species
of Homotherium and the specific identity of the Makapansgat specimen is there-
fore also not settled. It is concluded that it should for the present be listed as
Homotherium sp. indet., while L 11846 is tentatively identified as belonging to a
Homotherium as well.

The fact that the two ‘E’ Quarry machairodonts are recorded from different
units in the stratigraphic succession suggests that they might not have occupied
the area contemporaneously. These large predators are, however, very poorly
represented at the site and this supposition is thus very insecurely based. Since
the two species evidently differed considerably in size, it is likely that their
preferred prey also differed so that their co-existence need not necessarily have
been a problem. Taking this into account, together with the inferred broad
contemporaneity of Beds 2 and ga, it may yet prove that both species occurred
in the area at the time of deposition of both beds.

160 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Felidae
Subfamily Felinae

Felis aff. isstodorensis Croizet & Jobert, 1828
(Fig. 33)

Material

L 16055C—Left mandibular fragment with or parts of P, and P,, and M,.
L 3199 —Right mandibular fragment with M,.

L 15608 —Right C.

L 15788 —Left P#.

Locality and horizon

All these specimens are from ‘E’ Quarry, Langebaanweg. L 3199 is of
unknown provenance and the others are from Bed ga.

Description
This material belongs to a felid which was similar in size to the modern

lynxes.
The CG is comparable in size to those of large individuals in the available

series of modern Felis caracal (n = 19) (Table 31), but differs in that the groove
on the buccal surface is longer and deeper, while the antero-internal keel is

more prominent.

TABLE 31

Dimensions of teeth of the Langebaanweg Felis aff. isstodorensis, compared with those of some
other lynxes.

C p4 M, CHM P,-M,
l b l b ] b ] Nt
= 15608, Lrv5 738) ¢.8,8 70 | 17,6 854 — — — —
b o |
Pe L16055C —- — — — 15,3, ¢. 6,0 c. 56,0 c. 36,0
38 Pe |
io)
L3199 sae tae le eg a | ES Oie) 559 =: =
F’.. isstodorensis (Means) * — — | 20,2 9,5 14,7. — —- 36,8
Modern Mean 7) 5,5) || tO 97-3 | 122) 5,4 44,3 31,1
aes Range 6,5- 4,8- | 14,8- 6,4- | I1,0- 4,9- 41,4— 29,0—
9,0 6,7 | 17,8 8,7 | 14,0 6,1 5055 33,9
n 17 19 19 19 7}

* Saint-Vallier specimens (Viret 1954).

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 161

The P* is similar in all observable respects to the P* of large individuals
in the F. caracal comparative series.

The more complete of the mandibular fragments (L 16055C) has a longer
post-canine diastema than any specimen in the comparative series and, as a
result, the C to M, length is also greater. The other specimen (L 3199) may
belong to a female, since the M, is much smaller than that of L 16055C.
However, even though this tooth is also smaller than those of males in the
comparative series, that part of the mandibular corpus which remains is as
large as corresponding parts of the mandibles of male F. caracal. This suggests
that the fossil species was somewhat larger than the extant South African F.
caracal.

The C has a larger root than any specimen in the comparative series and
the groove on the buccal surface of the crown is more pronounced.

Although the P, and P, of L 16055C are damaged, they appear to be
similar to the corresponding teeth of F. caracal.

In addition to the difference in their sizes, the two known lower carnassials
also differ in that the smaller (L 3199) has a less prominent talonid. The M, of
L 16055C has a very small metaconid, while in L 3199 there is just a trace of
this cusp still visible. The former also has a slightly more prominently developed
buccal cingulum. Both specimens have a wide and deep carnassial notch,
which contrasts with F. caracal in which the notch is closed and terminates
further from the cingulum.

| 112 13 114 WS 116 117 1

Fig. 33. Buccal view of the Felis aff. issiodorensis mandible
(L 16055) from Langebaanweg.

Although the provenance of L 3199 is not recorded, the preservation and
adhering matrix suggests that it also came from Bed ga. The observable
differences between the lower carnassials of these two specimens are probably
no greater than might be expected in a single species, while the size difference is

162 ANNALS OF THE SOUTH AFRICAN MUSEUM

certainly within reasonable limits (Table 31). There are, therefore, no grounds
for believing that more than one species is represented by the material listed
above.

Discussion

The modern felids of southern Africa range in size from the small Felis
nigripes, through FP. libyca, F. serval, F. caracal, Panthera pardus, Acinonyx jubatus,
to the large Panthera leo. A similar range of species of differing size is recorded
elsewhere in the world. Dental and osteological characters of certain species
in any given area may be such that they can be most readily distinguished by
their size. However, overlaps in the size range of variation in species can lead to
great, or even insuperable problems !n the classification of fragmentary remains.
A further complication with fossil populations is that these may comprise
individuals whose average size 1s greater or less than those of extant populations.

It is here assumed that the small feline from Langebaanweg is most closely
related to the modern species which are closest to it in size, i.e. the lynxes. While
it is here regarded as a lynx, it might alternatively or additionally have close
phyletic links with the wild cat and serval groups. Lynxes are widely distributed
in the Old and New Worlds and a distinction is made between the essentially
Holarctic Felis (Lynx) and the essentially Ethiopian and Oriental Felis (Caracal).
In view of the age of the Langebaanweg species, its possible relationships to
both these groups must be considered.

F. caracal still occurs in the south-western Cape today, but in view of the
temporal and observed morphological differences between it and the Lange-
baanweg form, it is very unlikely that they are conspecific. In the case of the
extant species of Felis (Lynx) there is the additional factor of wide geographical
separation which is applicable.

Kurtén (1968: 80) regards the European Villafranchian F. issiodorensis
Croizet & Jobert and the possibly conspecific Chinese F. shansius Teilhard as
being ‘close to the starting point of the evolution of lynxes’. He also regards the
earlier North American species, FP’. rexroadensis Stephens, as the possible ancestor
of F. tssiodorensis, and stated that, ‘The Issoire lynx may well be ancestral to all
the living Old World lynxes’. Since the Langebaanweg species is probably older
by at least 0,5 million years than the earliest European record of F. issiodorensis
(i.e. that from Etouaires), and is comparable in age to F. rexroadensis, it must be
considered in relation to both these species.

One of the characteristics of the European F. lynx lineage is the regeneration
in M, of a small metaconid and an increase in the size of the talonid (Kurtén
1963a). The Etouaires form of F. isstodorensis lacks a metaconid and has a rudi-
mentary talonid and in these respects it resembles the M, of L 3199. On the
other hand, the characters of the M, of L 16055C are more comparable to
advanced F. issiodorensis. In view of the relative ages of these forms, that from
Langebaanweg might be regarded as having retained in M, some of the
characters evident in the ancestor of felines, namely, Pseudaelurus, but at the

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 163

same time showing the trend towards reduction of metaconid and talonid,
which in the F. (Lynx) lineage was climaxed in the Etouaires F. issiodorensis.
Kurtén (1963) states that in only 30% of modern F. caracal (n = 23) is the M,
metaconid and talonid present. In 8 recent specimens from the Cape Province,
the metaconid was not present at all, while the talonid was always very small.

The implications are that the Langebaanweg species could be ancestral
to both F. issiodorensis, and therefore also F. lynx, as well as F. caracal. In the
European F. lynx lineage there was a reversal of the trend toward reduction
of the M, metaconid and talonid, but in the African F. caracal lineage this trend
was apparently continued.

In general, the teeth and mandible of the Langebaanweg lynx are quite
similar to those of F. issiodorensis. For example, the Saint-Vallier form of this
species (Viret 1954) is comparable in size to the Langebaanweg lynx (Table 31),
it also has the buccal surfaces of the canines deeply grooved, while the post-
canine diastema of the mandible is equally long. However, the illustrated Saint-
Vallier specimens, like modern lynxes, differ from the Langebaanweg form in
that the M, carnassial notch is closed.

Unfortunately, the lower dentition of F. rexroadensis is unknown (Bjork,
1970), but the Langebaanweg P* differs from that of the American species in
that the protocone is not as posteriorly situated (see Stephens 1959). It is more
like F. issiodorensis in this respect. Another North American species, F. lacustris
Gazin, is closely related to F. rexroadensis and is of comparable age (Bjork 1970).
In this species the P* protocone is similarly situated to that of the Langebaanweg
form. However, the M, of F. lacustris differs from the Langebaanweg specimens
in that it lacks both metaconid and talonid (Gazin 1933). F. lacustris is also a
larger species.

Although they all date from the late Pliocene, it is unlikely that there is any
close connection between the Langebaanweg species and the two from North
America. Apart from the wide geographical separation which must lessen the
chances of direct phyletic links, Bjork (1970: 43, 44) concluded that although
it is possible that the affinities of F. rexroadensis and F. lacustris lie with the ‘Lynx
group’, this ‘must be considered with caution’, and he ‘would not assign [them]
to Lynx’.

On the basis of the material presently available, the Langebaanweg species
cannot be differentiated from F. isstodorensis and it probably represents an
early stage in the development of this species. Since it is not certain that the
two are conspecific, the Langebaanweg lynx can only be said to have affinities
with F. isstodorensis.

The possible phyletic relationships of fossil and modern lynxes will be
considered again following the description of the second feline from Langebaan-
weg (vide infra).

164 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Felidae
Subfamily Felinae
Felis obscura n. sp.
(Fig. 34)
Holotype

L ro10o0o—Left maxillary fragment with C to M!, with parts of premaxilla and
jugal attached.

Locality and horizon

This specimen is from Bed ga, ‘E’ Quarry, Langebaanweg.

Diagnosis

A medium-sized feline with a short face; ascending ramus of premaxilla
broad; P? absent; P® lacking anterior accessory cusp and with the posterior
accessory cusp flanked posteriorly by a smaller cusp projecting from the cingu-
lum; P* narrow, but with a prominent protocone linked to the paracone by
a ridge of enamel; P* parastyle relatively small and a very small ‘ectoparastyle’
also present; P* paracone and metastyle approximately equal in length; M!
double-rooted and nearly triangular in shape.

Etymology

The specific name indicates the doubt which exists about the relationships
of the species.

Description

This specimen belongs to a felid which in some respects is comparable to
small forms of modern Panthera pardus. It was initially compared and contrasted
with a series of seven skulls of leopards from the south-western Cape and
adjacent mountains, which represent a small variety of the species. Subsequently
it was compared to other felines, including the Felis caracal series already referred
to.

The pre-canine diastema is shorter than those of the available P. pardus
specimens and the premaxilla does not project as far forward. The ascending
ramus of the premaxilla is much broader in lateral view than in P. pardus.

The C of L 10100 is damaged, but it was evidently a fairly high crowned
tooth. It is more transversely compressed and has a somewhat flatter lingual
surface than the upper canines of the P. pardus series. The posterior keel was
prominent, although its enamel has been lost either through wear or post-
mortem damage.

The post-canine diastema is short, measuring only 5,5 mm, and P! and P?
are absent.

LATE CENOZOIG CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 165

The P® is slightly smaller than those of the P. pardus series (Table 32),
but otherwise differs only in that the posterior accessory cusp is flanked by a
more prominently developed posterior cingulum, the median part of which is in
the form of an additional small cusp. In this respect it is more like the P® of
available Acinonyx jubatus specimens, but it differs from this species in that it
lacks an anterior accessory cusp.

TABLE 32

Dimensions of the teeth of the Langebaanweg Felis obscura, compared with
those of Sivafelis potens from the Siwaliks and a series of modern Panthera

pardus.

r: Fe Mi}
] b ] b ] b
Lio100 14,0 6.3 |¢.29.8 Swe he. 5.6 734
Sivafelis potens* = a 24.0: 11,5 a irae
Modern Mean 15,1 735 23,57 12,4 ms. ie
Panthera —— 14,8- 6,7- | 20,7- 10,4- = rm
1557 8,2 24,9 12,9 == oe

pardus n 7 7 cae

* Pilgrim 1932.

The P?# is similar in size to the smaller of the P. pardus specimens, but the
shearing blade is more slender, the parastyle less prominent, while the protocone
is more prominent. The apex of the protocone is situated opposite the notch
between the parastyle and paracone and there is a ridge of enamel between it
it and the apex of the paracone. There is a small hollow in the enamel surface
between this ridge and the parastyle. The paracone is slightly longer than the
metastyle, and both are longer than the parastyle. The anterior and posterior
keels of the paracone are more or less mirror images of one another and this
cusp resembles that of Acinonyx. In P. pardus and other extant South African
felids, the anterior keel is more rectilinear than the other. There is a tiny cusp
situated antero-externally from the parastyle. It would perhaps be more correct
to regard it as a prominently developed part of the cingulum rather than as a
true ectoparastyle such as that of the machairodonts. There is a similar but more
obviously developed feature in the P* of the Acinonyx comparative series.

The M? has two roots and since the metastyle is still comparatively well
developed, it is almost triangular in outline.

The distance between the P! alveolar margin and the inferior margin of the
orbit is appreciably less than in P. pardus, while the infra-orbital foramen is
slightly elongated dorso-ventrally rather than being circular in shape. In the

ANNALS OF THE SOUTH AFRICAN MUSEUM

166

‘Somuveqoesury] wos (O10! J) eyIxeu
DANISGO St}aq IY} JO SMITA [enSuI] pue jesnjo00 ‘qeoong ‘bE “Bry

bea

I iin mm mm nm ii a ni

I

L\1 git S| b\l £{l c | (
NIHON

AUNLAVIUIUULLAIIUT wt

i

.

I I

LATE CENOZOIC CARNIVORA OF SOUTH=-WESTERN CAPE PROVINCE 167

fossil there is an additional very tiny foramen situated on the inferior margin of
the infra-orbital foramen. A similar feature was observed in the available felid
comparative material only in a skull of a North American F. rufus.

Discussion

Since this species lacks P? its affinities clearly do not lie with either Panthera
or Acinonyx. In general, it probably resembles Felis caracal more closely than
any other extant southern African felid.

It is similar to F. caracal in the following respects:

(1) Both species have a relatively broad ascending ramus of the premaxilla.

(2) The shape of the infra-orbital foramen and its position relative to the cheekteeth are similar
in the two species.

(3) Both species lack P?.

(4) Both have relatively short pre- and post-canine diastemas.

(5) The P? morphology of the two species is similar, although F. caracal may sometimes have a
small anterior accessory cusp and the posterior cingulum cusp may be relatively less promi-
nent.

(6) In general, the P4 of the two species are similar, although the protocone of F. caracal is
relatively reduced and in this species there is only occasionally a slight thickening of the
cingulum antero-externally to the parastyle (‘ectoparastyle’).

The fossil and F. caracal definitely differ in the following respects:

(1) F. caracal is a much smaller species.

(2) In lateral view the jugal of F. caracal extends as far forward as the infra-orbital foramen,
whereas in L roroo it does not. In this respect the fossil resembles P. pardus and Acinonyx.

(3) The M! of F. caracal is relatively more reduced.

There is nothing which would preclude L 10100 belonging to a species
which was directly ancestral to F. caracal and there is a good deal which suggests
that such a relationship might be possible.

Of recorded fossil species, those referred to Sivafelis Pilgrim, 1932 are
perhaps most likely to have affinities with L 1o1oo. Pilgrim (1932) included
three species in this genus, namely, S. potens Pilgrim and S. brachygnathus Lydek-
ker, both of which are probably from the Pinjor stage of the Siwaliks, and
S. pleistocaenicus Zdansky from China. The Langebaanweg species apparently
predates the three Asiatic species and this reduces the likelihood of it being
conspecific with any of them.

Unfortunately, the holotypes and most of the referred material of the
Asiatic species are mandibles and comparisons with L ro1oo are therefore
unsatisfactory. Pilgrim (1932) has, however, referred a maxillary fragment
with P# to S. potens and this tooth is similar to the P* of L 10100 in some respects.
They are comparable in overall size, their protocones are similar in size and
position and in both the protocones are linked to the paracones by a ridge of
enamel. This ridge is separated from the parastyle by a small hollow in both
species, while they also have a similarly shaped paracone. They differ in that
the P* of S. potens has a longer metastyle, which may simply be a more advanced
characteristic in this species. The relatively short mandible of Sivafelis is matched
by the short maxilla of the Langebaanweg species.

168 ANNALS OF THE SOUTH AFRICAN MUSEUM

There are, therefore, grounds for believing that L 1o1oo belongs to a
species of Sivafelis, but one which is more primitive than the early Pleistocene
S. potens.

In his discussion of Szvafelzs, Pilgrim (1932) did not consider the possibility
of its relationship to F. caracal. On the basis of size alone, such a relationship
does at first sight seem rather improbable. There have, however, been some
very marked size changes in other felid lineages and the size factor is not
necessarily significant. It is thus possible that F. caracal stems from Sivafelis
rather than from F. zsstodorensis as was suggested earlier. ‘The phyletic relation-
ships of the lynxes, Szvafelis and L 10100 are therefore not altogether unequivocal
and the Langebaanweg species is named accordingly.

A possible explanation of the available record is that the dichotomy of the
Felis (Caracal) and Felis (Lynx) lineages took place earlier than the late Pliocene
and the Szvafelis group was ancestral to F. (Caracal), while the true lynxes arose
from F. issiodorensis, which is here taken to include the smallest of the Langebaan-
weg felines (Fig. 35).

Lu
Za
O Felis caracal Felis lynx Felis (Lynx) spp.
fe) Africa, Asia Eurasia North America
O
x=
Lu
FL
Lu
U
oO CARACAL LYNX
wn
wi F. potens F. issiodoreénsis
= 5 >
oa Asia Eurasia
SIVAFELIS
F. obscura F. aff. issiodorensis +
Lu Africa Africa
FL
Lu
OU
fe) T6 North American
re F. rexroadenis &
F. lacustris
e ie
ZA
Lu
6 Metailurus
=

Fig. 35. Tentative phylogeny of some small- to medium-sized Felidae.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 169

In the same way that the late Pliocene/early Pleistocene Vishnuictis was
earlier suggested to be a subgenus of Viverra, so Sivafelis is here regarded as a
contemporaneous subgenus of that group of felines commonly termed the
lynxes. L roroo is thus classified as Felis (Sivafelis) obscura.

Family Felidae
Subfamily Felinae
Dinofelis diastemata Astre, 1929
(Figs 36, 37, 38; Hendey 1970a: pl. 2D)

Material from Bed 2, ‘E” Quarry, Langebaanweg
L 20284 — Isolated right C; left and right mandibles with C to M,.

L 20685 —Isolated left and right C, right C and M,; left mandible lacking
incisors and P,; associated with both clavicles, several ribs and
vertebrae, some of which are intact, a few fragmentary long bones
and a number of carpal and tarsal bones, metapodials and phalanges.

L 20702 — Left C.

Material from Bed 3a, ‘E’ Quarry, Langebaanweg

L 12237—Left mandibular fragment with P, and M,.

Material from ‘E” Quarry, but of unknown provenance

L 2674—Parts of a skull, including left maxillary fragment with C and right
maxillary fragment with P’ to M!.

Description

This material belongs to a felid which is approximately the same size as
the Langebaanweg Machairodus, and one which has some machairodont
characteristics, although it is undoubtedly a feline.

Skull (Tables 33, 34)

The Bed 2 specimens L 20284, represent parts of a single skull. The isolated
C is far larger than the canines of the mandible and superficially it bears a
remarkable resemblance to the C of the Langebaanweg Machairodus (L 20505),
differing only in that it is slightly broader and in lacking serrations on the
prominently developed keels. It has a crown height of about 50 mm and the
overall length of the tooth along its anterior curve is about 120 mm. By contrast,
the crown height of the C is only 25 mm. Thus in respect of the development of
its canines this species is comparable to machairodonts, although the diametrical
index of the G (0,63), while being less than that of modern felines, is greater
than that of most machairodonts (see Ewer 1955c: Table 3). The enamel of this
tooth in the vicinity of the keels and on the buccal surface is finely rugose.

The GC, L 20702, is similar in all observable respects to that of L 20284,
although it is less well preserved. The anterior and posterior keels were worn

57o ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 33

Dimensions of the upper teeth of the Langebaanweg Dinofelis, compared with those of Dinofelis
from other localities.

Langebaanweg
Dinofelis Dinofelis Dinofelis
diastemata! | L2674 L20685 L20284 L20702 barlowi? piveteaui?
l = c. 19,0 19,6 ZOO). G5 21-5) C2480) ones 20,5
CG b — — 10,6 I@ok, $6.51 3-2) 1 Gp5-O) eine OIE
I:b 1 :0,54 = 1-0554, *020,03" 170,01 | 170,02) 1 -o.0n 1 :0,59
Postcanine
diastema — 455 — — — 9 — as
] 21 19,5 — — — 21,5 ¢.20,0 19,8
Ps
b -- c. 8,5 — — — Gales —- 10,0
l 29 €. 29,0 mi a 5 36,2 3754 41,0
P4
b — GUA a — — 16,5. | 16,6 iO
alveolus
= 5c, aia = ze — 4,5
M!
b —- 9g,0+ == — _— Tate -— 457

1 Piveteau 1948.
2 Ewer 1955.

during life, the apex of the crown is lost and the tooth is cracked longitudinally.

The right half of the mandible of L 20284 is the more complete and it
lacks only the most posterior parts, including the angular process and condyle.
The coronoid process is very high and typically feline in this respect, while in
marked contrast to the condition in machairodonts. The symphyseal region
of the mandible is, however, similar in structure to corresponding parts of the
mandible of the Bed 2 machairodont. The post-canine diastema is relatively
long and a small metal flange is present. The symphyseal regions of the mandi-
bles of the two species differ mainly in that in the machairodont it is broader,
higher and more stoutly developed.

The narrower mandibular symphysis in L 20284 is due to the fact that the
incisors of this species were much smaller than those of the machairodont.
Although these teeth have been lost, parts of the three alveoli are visible in the
left half of the mandible and all were clearly far smaller than the C. This
species differs markedly from machairodonts in this respect.

The C of L 20284 is less transversely compressed than the C and, while
it does have a prominent posterior keel, the anterior edge of the tooth is rounded.
The ‘anterior’ keel is rotated on to the lingual surface, where it terminates in a
bulge on the cingulum at about midway along the length of the tooth.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE E71

TABLE 34

Dimensions of the mandibles and lower teeth of the Langebaanweg Dinofelis, compared with
those of Dinofelis from other localities.

Dinofelis Langebaanweg
diaste- Dinofelis Dinofelis
mata\»? | L20284 L20685 L12237 barlowi* piveteaui®
= 15,0 14,6 12,9 —
C
b 10,5 =
Postcanine diastema 27,0 : —
] 16,0 Sax
Ps
b Se.
| 2357
Py
b 11,4
2757
M,
b 16,9
P,-M, | ==
Max. height of ramus
(coronoid process to inferior ae
margin)
Mandible height below M, 3552
Mandible breadth below M, 18,7

1 Astre 1929.
2 Piveteau 1948.
3 Ewer 1955.

The cheekteeth are typically feline in character. The P, is much smaller
than P, and has a prominent principal cusp, a posterior accessory cusp flanked
by a well developed posterior cingulum, while there is a shelving of the anterior
end of the tooth where the anterior accessory cusp would have been situated
if present. The P, is relatively long and narrow and, except for the presence of a
prominent anterior accessory cusp, its structure is similar to that of P;. The M,
is the largest of the cheekteeth and is made up only of the paraconid and
protoconid, although vestiges of a talonid are still evident. The paraconid
blade (6,5 mm) is far shorter than the protoconid blade (12,3 mm).

The specimens L 20685 belong to an individual which was appreciably
smaller than that represented by L 20284, the size difference perhaps reflecting
sexual dimorphism in the species. Otherwise the material is essentially similar

172 ANNALS OF THE SOUTH AFRICAN MUSEUM

6 7 8 S) ¢

mn HLL Hi Av mi HA

nn Hu a AA A LL III na il

Fig. 36. Buccal and occlusal views of the left mandible and buccal view of the right
mandible of the Dinofelis diastemata (L 20284) from Langebaanweg.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 173

HII i HUTA LULL LLU LL LL
110 11 112 113 114 115 116

Fig 37. Buccal and occlusal views of the Dinofelis diastemata
maxilla (L 2674) from Langebaanweg.

in all observable respects to that already described, although the C has a lower
diametrical index than either of the other two specimens.

The Bed 3a mandibular fragment (L 12237) is similar to corresponding
parts of the mandibles from Bed 2 and compares most closely in size with
L 20685. This specimen lacks P,, but the bone surface anterior to P, 1s scarred,
which suggests that P, was lost during life. The P, of the preserved left half of
the mandible of L 20685 is also missing, although in this instance the alveolus
is still open. There are, however, indications of periodontal disease at the
site of P,; with alveolar recession and an overgrowth of bone on the buccal
surface of the mandible. Consequently, this tooth might also have been lost
during life even though there are no indications of alveolar resorption.

The unprovenanced material (L 2674) belongs to an aged individual and

174 ANNALS OF THE SOUTH AFRICAN MUSEUM

witli!

S$

9

nan

ra

unlit

8

nln

6

unt

1

unt

|

mM

if

uni

{

uni

1

unt

1

9
mun

1

wn

if

un

i

una

A

ait

Fig. 38. Buccal view of the upper canines of Dinofelis diastemata (L 20284), Machairodus sp.
L 20505) and cf. Homotherium sp. (L 11846) from Langebaanweg.

-—

is very fragmentary. The preserved C is incomplete and is crushed so that its
dimensions cannot be accurately measured. The post-canine diastema is very
short and both P! and P? are absent. The P® is similar in morphology to the
P, of the Bed 2 and Bed 3a mandibles.

Detailed observations on the characteristics of the P* of L 2674 are ham-
pered by the fact that this tooth is very worn. The apices of all cusps have been
worn away and the buccal surface of the paracone is damaged. The paracone
is the most prominent of the cusps and is a little longer than the metastyle,
which in turn is a little longer than the parastyle. A small ‘ectoparastyle’ is
present and in relative size and position it resembles that of the Felzs obscura
holotype. The protocone is largely worn away, but it was evidently a fairly
prominent cusp situated adjacent to the notch between the parastyle and
paracone.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 175

The crown of M! is completely worn away, the wear facet being oval in
shape with its long axis at right angles to that of P*. The single, transversely
elongated root bifurcates near its base.

Postcranial skeleton (Tables 35 to 38)

The postcranial remains of L 20685 were compared with corresponding
parts of the skeletons of two modern leopards (Panthera pardus) and two cheetahs
(Acinonyx jubatus). In general the fossil bones were larger than those of the
leopards, although of similar proportions and therefore readily distinguishable
from the bones of the cheetahs.

The preserved vertebrae include three thoracics and five caudals. The
former are unremarkable being most similar to those of the cheetahs in size,
but the caudal vertebrae are appreciably shorter than those of the comparative
specimens. It was not possible to identify the positions of the fossil caudal
vertebrae, but three of them correspond morphologically most closely to the
6th to roth caudals of the comparative specimens. A comparison of the lengths
of the available fossil and modern specimens shows that the former are only
about 60% of the length of the leopard caudals and about 50% of those of the
cheetah (Table 35).

TABLE 35

Lengths of the caudal vertebrae of the Langebaanweg Dinofelis compared with those of modern
Panthera pardus and Acinonyx jubatus.

Dinofelis L20685

Modern SAM 36051
Panthera
pardus SAM 36324
Modern SAM 36703
Acinonyx
jubatus SAM 36704

Parts of at least nine ribs were recovered and of these three are largely
intact, while both clavicles, one of which is complete, are preserved. Damage to
ribs and vertebrae is largely of recent origin and resulted from plant growth in
the area where the skeleton was buried. The completeness of these relatively
delicate bones and of elements of the manus and pes is in marked contrast
with the fragmentary condition of the long bones, some of which have been
burnt.

176 ANNALS OF THE SOUTH AFRICAN MUSEUM

Only the left ulna was recovered reasonably intact, although it is very
poorly preserved, while the proximal end and part of the shaft of the right tibia
and the complete right fibula were restored from a number of fragments found
over an area of several square metres. These bones, and parts of both radii,
are appreciably larger than corresponding elements in the two modern leopard
skeletons. The articular surfaces of the fossil bones are, in some instances, similar
in size to those of the cheetahs, although they are otherwise more stoutly pro-
portioned. The few measurements possible on these bones are listed in Table 36.

TABLE 36

Dimensions of some elements of the postcranial skeleton of the Langebaanweg Dinofelis compared
with those of modern Panthera pardus and Acinonyx jubatus.

Radius Tibia Fibula

Ant.- | Ant.- Ant.-

Post. Trans. | Trans. post. post, || #linans:

diam. diam. diam. | Overall | diam. diam. diam.

prox. prox. prox. length prox. dist. dist.

end end end end end end
Dinofelis L20685 20,0 AVA || Gs RAK || Go QUO 24,6 16,8 Any,
Modern SAM 36051 42,9 198,0 16,6 8,1
Panthera ——— ——|— ————|— eg
pardus SAM 36324 39,8 191,0 15,0 ASS)
Modern SAM 36703 16,7 21,7 51,8 280,0 19,9 17,5 9,2
Acinonyx =———_——— |---|] — —— | —__—_—.
jubatus SAM 36704 1(5)7) 22,3 51,5 270,0 20,5 19,0 9,6

Parts of all four feet were recovered and the various elements are mostly
complete and well preserved. A few exhibit a pathological condition (osteitis).
Apart from the fact that they are appreciably larger than the corresponding
bones of the leopards, they are otherwise similar in both morphology and
proportions, although perhaps somewhat more stoutly built. In respect of their
proportions they are clearly distinguishable from the generally elongated foot
bones of the cheetah which have been adapted for cursorial locomotion. ‘The
dimensions of some of the elements of the manus and pes are given in Table 37.

The fossil species differs from both the leopard and cheetah in respect of the
relative size of its fore- and hindfeet. This is illustrated by a comparison of the
relative lengths of the metapodials II of the three species (Table 38).

The Langebaanweg Dinofelis was evidently a heavily built animal, with
the fore- and hindfeet, and perhaps the limbs in general, being more equally
proportioned than in either the leopard or cheetah, and possessing a relatively
short tail. Its locomotion is likely to have been ambulatory. The indications
are thus that it had paralleled the developments in the Machairodontinae in
its postcranial skeleton as well as the skull.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE eG,

TABLE 37

Dimensions of elements of the manus and pes of the Langebaanweg Dinofelis (L20685).

Meta- Meta- Meta- Meta- Meta- Meta- Meta-

carpal | carpal | carpal | carpal tarsal tarsal tarsal
I II III V JRE IV Vv

Overall length 25,0 70,5 78,4 60,2 80,9 90,8 80,8
Max. ant. post. diam. at
prox. end 13,2 18,4 16,3 14,9 18,0 19,3 12,2
Max. trans. diam. at
prox. end 16,9 14,0 16,0 12,5 10,5 15,8 16,0
Max. ant. post. diam. at
distal end 13,7 14,3 14,3 13,8 14,3 14,3 13,1
Max. trans. diam. at
distal end 1554 12,6 13,4 12,8 Ne 12,7 12,0

Calcaneum | Astragalus Navicular Cuboid
Overall length 70,9 37,2 — 19,6
Max. dorso-ventral diam. 30,8 21,3 26,5 18,3
Max. trans. diam. 33,2 33,4 19,3 17,8
TABLE 38

Length ratios of metacarpal II and metatarsal II of the Langebaanweg Dinofelis
compared with those of modern Panthera pardus and Acinonyx jubatus.

Mc II length | Mt II length | McII: Mt II

Dinofelis L20685 ie a
Modern SAM 36051 Eyck Lege
Panthera a

pardus SAM 36324 be, 1529
Modern SAM 36703 22 Tas
Acinonyx | -

jubatus SAM 36704 76,4 102,6 PP WGs4

178 ANNALS OF THE SOUTH AFRICAN MUSEUM

Discussion

These specimens clearly belong to a ‘false sabre-toothed cat’ in that group
of felids referred to Dinofelis Zdansky by Hemmer (1965). Included in this
group are three species which were previously referred to the genus Therailurus
Piveteau, namely, the European D. diastemata Astre, 1929, D. barlowi Broom,
1937 from Sterkfontein and D. piveteaut Ewer, 1955 from Kromdraai.

In the study of this group, Hemmer (1965: 75) arrived at the following
conclusions:

“The most primitive species concerning the specialization of the teeth is
Dinofelis diastemata (Astre 1929), the most evolved is Dinofelis piveteauz (Ewer
1955) and by parallel evolution Dinofelis abeli Zdansky 1924. D. diastemata—
D. barlowi—D. piveteaur form a connected succession whilst D. abel: takes a
more isolated place.’

The Langebaanweg Dinofelis evidently has its affinities with the D. diaste-
mata— D. piveteaut series. ,

The European species is Astian in age and is, therefore, broadly con-
temporaneous with the Langebaanweg material, while the two recorded South
African species date from the Makapanian. Consequently, because of geo-
graphical factors, it might be expected that the Langebaanweg species would
have close phyletic links with D. barlowi and D. piveteaut, but because of its age
it would be at a stage in evolutionary development comparable to D. diastemata.
This hypothesis was found to conform quite well with the nature of the various
specimens available.

Ewer (1955¢: 598) expressed doubts as to whether D. piveteau: could have
been directly descended from D. diastemata because of the low CG diametrical
index of the European species, although in other respects ‘(Dinofelis) piveteaut
is considerably more advanced than (D.) diastemata.’ As Ewer was dealing with
only one specimen from each population no account could be taken of the
possible range of variation in this particular character. In fact the range of the
diametrical indices of the three specimens from Langebaanweg (0,54 to 0,63)
actually exceeds that observed in the four specimens of the three previously
recorded species (0,54 to 0,62). Possibly when larger samples of the three species
become available, it will be found that their mean diametrical indices do differ,
but this is clearly not a reliable differentiating characteristic at present and it
cannot be used as a basis for determining phyletic relationships.

In all other observable respects the Langebaanweg Dinofelis is apparently
more similar to D. diastemata than to the Transvaal species. Probably the most
obvious character which differentiates the Pliocene specimens from those of the
Pleistocene is the relative sizes of their carnassials. These teeth are longest in
D. pwweteaui, the most recent of the species, a little shorter in D. barlowi, and
shortest in the Langebaanweg Dinofelis and D. diastemata from Europe.

There are actually no grounds for believing that the Langebaanweg
Dinefelis and the European D. diastemata are not conspecific and consequently

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 179

the former is identified accordingly. ‘The suggested phyletic relationships of the
various species are illustrated in Figure 39.

The characteristics of Dinofelis are yet another indication of the felid pro-
pensity for developing the ‘sabre-tooth’ condition. Although the Dzunofelis
experiment in this direction was not carried to the extreme evident in the three
sabre-toothed cat subfamilies (Hoplophoneinae, Nimravinae, Machairo-
dontinae—see Thenius 1967), it too was ultimately unsuccessful. The develop-
ment of ‘sabre’ canines by a member of the subfamily Felinae is not confined to
Dinofelis, since a similar development is evident in the extant clouded leopard

of Asia (Neofelis nebulosa).

Lu

Z

Lu

UO

oO

al

O

aE

Lu

&

U Dinofelis piveteaui
O Africa

Bi Dinofelis abeli

a Asia

oo Dinofelis barlowi

Africa

Dinofelis diastema Dinofelis diastemata
Europe ———————————Affrica
Lu
Zz
Lu
U
je)
= +
Metailurus
Fig. 39. Tentative phylogeny of the genus Dinofelis.
Suborder FISSIPEDIA
Incertae sedis
(Fig. 40)
Material

L 11752—Right maxillary fragment with C and two premolars (? P! and P?).
L 12273—Left maxillary fragment belonging almost certainly to the same
individual as L 11752.

180 ANNALS OF THE SOUTH AFRICAN MUSEUM

Locality and horizon

These specimens are from Bed 2, ‘E’ Quarry, Langebaanweg.

Description

The specimen L 11752 comprises the most anterior part of a right maxilla
to which part of the premaxilla is fused. The alveoli of I? and I® are present,
the C is damaged and the first two premolars are preserved intact. The anterior
rim of the orbit is damaged. The second specimen is only a small part of the left
maxilla in which the anterior part of the orbit and immediately adjacent areas
are preserved. The preservation of the two specimens is identical and they are
essentially similar in the features which are observable. Although they were
not found at the same time, it is probable that they do belong to the same
individual. ;

The species concerned is of large size, the canine being comparable to that
of modern Crocuta, but the snout is extremely short and the two premolars much
reduced in size (Table 39). Although only two incisor alveoli are preserved,
this species almost certainly had I? as well. Nothing is known of the nature of
the posterior cheekteeth.

TABLE 39

Dimensions of the upper teeth and facial region of the unclassified
fissiped carnivore from Langebaanweg (L 11752).

Is C ee PP
(alveolus)
l b l b ] b ] b

EX134G) 69760) |cx2050) 254 6,2 6,0 6,0 6.553

Distance between I? anterior alveolar margin and infra-orbital
foramen—c. 50,0

The infra-orbital foramina are unusual in that both are divided into two
more or less equally sized parts. In the right maxilla there are two distinct
foramina, but in the left there is a single external opening which divides inter-
nally. In the fairly comprehensive collection of skulls of modern Carnivora
examined in the course of the present study, the only species in which a com-
parable development of the infra-orbital foramen was observed was Proteles
cristatus. In a series of six Proteles skulls, two had both infra-orbital foramina as
single openings, two had one single and one double opening, and two had both
as double openings.

Discussion

The large canine and reduced anterior premolars of this species are
reminiscent of the condition in many ursids, but the snout of the fossil is even
shorter than that of such short-faced forms as Helarctos malayanus. Furthermore,

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 181

the P4 of the fossil must have been situated posterior to the infra-orbital foramen,
whereas in the ursids this tooth is always situated anterior to the foramen. The
fossil is almost certainly not an ursid and the specimens definitely do not belong
to the Langebaanweg Agriotherium africanum.

The nature of the infra-orbital foramina and reduced cheekteeth suggested
that the fossil might represent a primitive form of Proteles. L 11752 is, however,
far larger than corresponding parts of the skull of modern P. cristatus and the
premolars are more complex than the simple peg-like cheekteeth of P. cristatus,
being more circular in shape with a more prominent internal cingulum flanking
a lower and blunter principal cusp. In addition, the ? P? is a little smaller than
the ? P!, whereas in P. cristatus the second postcanine is larger than the first.
On the other hand, the fossil premolars are not dissimilar to the P! of Hyaeninae
and since Proteles is evidently descended from an early hyaenid, it was thought
possible that the premolars of L 11752 might represent an early stage in the
development of Proteles cheekteeth. Had L 11752 been found without the
second reduced premolar, and if the greatly foreshortened snout had not been
evident, then it might well have been identified as belonging to a hyaenid.

Although the structure of the infra-orbital foramina of Proteles and the
fossil is similar, in the former this foramen is situated above or slightly posterior
to the third postcanine, whereas in the fossil it is a little posterior to the ? P?.
The snout of the fossil is, therefore, relatively much shorter than that of Proteles.

Apart from the obvious size and morphological differences between the
fossil and Proteles, a relationship between the two might also be doubted on
purely theoretical grounds. Proteles has an extremely poor fossil record, but it is
known from the Makapanian, being represented by a single mandibular frag-
ment from Kromdraai, which differs from the moderns in only a few respects.
The implication is that for much of the Pleistocene at least, Proteles was repre-
sented by forms which were not very different from that which is still extant
(Hendey 1973a). It would therefore be surprising if the Makapanian Proteles
was descended from so large a Langebaanian species as that represented by
L 11752. Its derivation from a small form such as the Pontian Jctitherium orbignyt
(see Thenius 1966) is far more likely.

Another carnivore family with which the fossil was compared was the
Mustelidae. A shortened face and reduced anterior cheekteeth are characteristics
of the Lutrinae and since a large otter, Enhydriodon africanus, is included in the
‘E’ Quarry assemblage, L 11752 was first compared with this species. The
E. africanus mandible (L 9138) has very crowded anterior cheekteeth, whereas
L 11752 has a postcanine diastema of 8 mm and gaps of 2 mm and 3 mm
between the first three cheekteeth. The two specimens are clearly not con-
specific and judging from the size of the C, L 11752 belonged to a species even
larger than E. africanus. The modern Lutrinae, Aonyx, Enhydra and Lutra, also
have crowded anterior cheekteeth and are very much smaller even than
E. africanus, so that there is clearly not a close relationship between them and

Ett 752 either.

182 ANNALS OF THE SOUTH AFRICAN MUSEUM

hi LITA UT mn

fi IQOUID ALA UU LTUUATC U

Fig. 40. Occlusal and buccal views of the unidentified
fissiped carnivore maxilla (L 11752) from Langebaanweg.

Although it seemed unlikely that the fossil species was an otter, Pilgrim
(1932: 80, 81; pl. 17) has described a maxillary fragment from the Dhok
Pathan stage of the Siwaliks (GSI D 207) which is similar to L 11752 in some
respects, and which he believed belonged ‘to an animal of Lutrine lineage
which had specialized earlier than Enhydriodon, but along somewhat similar
lines’. According to Pilgrim it had ‘obviously remained closer to Potamotherium’.

Unfortunately the premaxilla, orbit and infra-orbital foramen of the
Indian specimen are lost, but it is similar to L 11752 in that it has ‘an excep-
tionally large’ canine and its ‘P? has a circular crown due to the presence of a

LATE CENOZOICG CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 183

marked internal cingulum’. It differs from L 11752 in that it lacks P! and has a
very short postcanine diastema. The C of the Indian specimen is smaller
(16,8 x 10,5 mm) than that from Langebaanweg, but the P? (6,2 5,7) is
similar in size. The P? of D 207 is a fairly large tooth (13,4 7,7 mm). It is
possible that the two premolars of L 11752 are in fact P? and P? and, if so, it
cculd represent a more advanced form of the unnamed Dhok Pathan species
in which P? was much reduced in size. Alternatively, it might be a representa-
tive of an African lineage which had an ancestor in common with that which
includes the Dhok Pathan species.

It is unlikely that L 11752 and D 207 are conspecific, but they could be
congeneric. Certainly of all the recorded late Cenozoic Carnivora, D 207
appears to be the one which is most likely to be related to the problematical
Langebaanweg species.

Particular interest is attached to L 11752 and the companion specimen,
since if they are lutrine, it means that there are two large otters represented
in the Langebaanweg assemblage and not just Enhydriodon. In addition, these
specimens probably belong to a genus which is as yet unnamed and if more
specimens can be found which would determine conclusively the relationships
of this animal, it would be another significant addition to the late Cenozoic
fossil record.

All that can be said of the specimens at present is that they belong to a
poorly known species, which may belong in a group of otters descended from
Potamotherium.

Suborder FISSIPEDIA

Unclassified specimens

The Langebaanweg assemblage, including that from ‘C’ Quarry, includes
carnivore specimens in addition to those already listed. Most of this material,
which is made up of relatively few specimens, is at least classified to the family
level and it is likely that most, if not all the specimens belong to species already
listed. Cranial remains make up only a small proportion of the unclassified
material and consist mostly of tooth fragments. Carnivore postcranial remains
from Langebaanweg are usually found dissociated from more readily identifiable
skull parts. In the case of the pinniped no problem in identification has arisen
as there is evidently only one species represented, but the same does not apply
to the fissiped postcranial remains.

The only fissiped species which has had elements of the postcranial skeleton
positively identified even though they were not found associated with skull parts
in Agriotherium africanum. This species is readily distinguished from others by its
very large size. Viverrid, hyaenid and felid postcranial bones are not uncommon
and although some have been identified with particular species, they were
excluded from the relevant specimen lists because such identifications were not
positive. There is no point at this stage in possibly confusing the issue on the

184 ANNALS OF THE SOUTH AFRICAN MUSEUM

nature of the postcranial skeletons of certain species. As more definite associa-
tions of skull and postcranial bones are discovered, it should become possible to
positively identify unassociated postcranial remains and their description can
be left to some future date. The same attitude was adopted in the case of the
Quaternary Carnivora.

Reference has already been made to pathological conditions evident in
some of the carnivore skeletal remains from Langebaanweg and there are other
such examples amongst the unclassified material. Although past palaeopatholo-
gical studies have been largely confined to hominid remains, there is a growing
interest in this subject as it relates to other mammals as well. For this reason,
and also because bone pathology is apparently more common amongst the
Carnivora than it is in other mammals from Langebaanweg, some account was
taken of those bones which show obvious indications of abnormal conditions.
Only two such conditions were recognized, namely, osteitis and osteo-arthritis,
the latter indicating diseases of the joints and the former indicating all other
bone inflammations (see Brothwell 1963). Specimens not already mentioned
include:

L 6388 —Right tibia (? Machairodus). Osteitis, mainly near the proximal end.
Osteo-arthritis at the proximal end, with severe eburnation of the
lateral articular facet and lipping of the bone (Fig. 41).

L 6422 & L 6425—Left and right calcanea (? Machairodus). Osteitis on lateral
surfaces anterior to distal extremity.

L 6430 —Metapodial (? Machairodus). Osteitis on dorsal surface. Osteo-arthritis
(eburnation) of distal articular facet.

The above four specimens may belong to a single individual.

L 9868 & L 9869—Proximal end of left ulna and distal end of left humerus
(? Machairodus). Severe osteitis on lateral and medial sides of ulna
and a less extensive but similar condition on the humerus, particu-
larly the arch enclosing the entepicondylar foramen (Fig. 41).

L 6391 —Distal end of right tibia (? Machairodus). Severe osteitis on the shaft.

L 6409 —Proximal end of left humerus (? Machairodus). Extensive osteitis.

L 6403 —Distal end of left humerus (Hyaenidae). Osteitis of the supra-
trochlear fossa.

L 9884 —Left radius lacking distal epiphysis (? Machairodus, immature).
Severe osteitis on the shaft.

L 12642—Metapodial (? Agriothertum). Osteitis, possibly resulting from a frac-
ture. The shaft is bowed and thickened, the inflammation being
largely confined to the thickened part.

Pathological conditions in the postcranial skeletons of the Hyaenictis
preforfex holotype and the Percrocuta australis specimen, L 13033, have already
been mentioned and in both these instances the conditions may relate to the
advanced age of the individuals concerned.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 185

ili

bg

nl

i

ii

6

man

i

i

ml

i

mn

I

ih

t

mn

git

antl

Fig. 41. Pathological conditions in carnivore postcranial bones from Langebaanweg. Tibia
(L 6388) showing eburnation and bony lipping (osteo-arthritis) of the proximal end. Ulna
(L 9868) showing osteitis at the proximal end and on the shaft.

There are two points worth noting about the specimens listed above.
Firstly, although relatively few individuals are represented, most of the speci-
mens are tentatively identified as belonging to a machairodont. Secondly,
both immature individuals and adults of this species are afflicted by osteitis,
indicating that it was not necessarily an age-related condition. On the other
hand, the osteo-arthritis may well have been related to advanced age.

186 ANNALS OF THE SOUTH AFRICAN MUSEUM

CARNIVORA OF THE PLEISTOCENE AND HOLOCENE

Although there are more carnivore species recorded from the Quaternary
of the south-western Cape than from the Pliocene, none of the individual
fossil assemblages matches that from ‘E’ Quarry in the variety of species repre-
sented. ‘The most prolific of the local Quaternary occurrences is Elandsfontein
and it is also this site which has yielded the largest number of carnivore species,
namely, fifteen, as against twenty from Langebaanweg. The list of Quaternary
species is lengthened by those which have been recorded during the historic
period, but which are not known locally as fossils.

The Quaternary carnivores are less problematical than those from Lange-
baanweg since, with few exceptions, they are closely related to species which are
still extant, and their identification was facilitated by comparisons with available
osteological material belonging to modern species. In most instances, however,
the modern comparative series were small and some of the observations made
in respect of the characteristics of certain species may be incorrect.

The local record of Pleistocene mammals is far from complete. Those few
species from Baard’s Quarry which are regarded as Pleistocene in age, probably
date from the earlier part of this epoch. Otherwise it is the Elandsfontein fauna
which includes the earliest of the local Pleistocene mammal fossils. The record
for the later Pleistocene and Holocene is comparatively good. In other words,
the Makapanian fauna is known only from the almost insignificant Baard’s
Quarry occurrence, the Cornelian is known from one reasonably large assem-
blage, while only the Florisian and Holocene faunas are quite well known.

Since the early element in the Elandsfontein fauna may date back no more
than 0,5 million years (vide supra), and the Plio/Pleistocene boundary is here
taken at 3,5 million B.P., it means that only about one-seventh of the Pleistocene
is represented in the local fossil record by assemblages including appreciable
numbers of specimens. It also means that the Pliocene species from Langebaan-
weg predate the next oldest of the larger local assemblages by perhaps 3,5
million years and it is, therefore, not surprising that the Langebaanweg Car-
nivora have so little in common with those from the local Pleistocene.

Suborder PINNIPEDIA
Family Phocidae
Subfamily Monachinae
Hydrurga leptonyx Blainville, 1820

The leopard seal is an Antarctic species which normally lives on the outer
fringes of the pack-ice, but which occasionally strays to the more southerly
parts of South America, Africa and Australasia (King 1964). It is not known as a
fossil in South Africa, but Roberts (1951) records the landing of a female on
the coast about 40 miles north of East London in September, 1946. The only
other South African record of this species is from the south-western Cape.

LATE CENOZOICGC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE roy

An immature female was found alive on the beach at Hout Bay near Cape
Town in October 1969. It subsequently died in a local zoo and its skeleton is
preserved in the South African Museum (SAM 35796).

It may be significant that these two specimens reached South Africa during
the same season (i.e. the southern Spring). Scheffer (1958) has reported that
leopard seals migrate northwards to ice-free islands during Winter and it is
possible that in the return migration the following Spring some individuals
become disorientated and land as strays far from their normal habitat.

Family Phocidae
Subfamily Monachinae

Lobodon carcinophagus Hombron & Jacquinot, 1842

Present status

The crabeater seal is an Antarctic species, but occasional stray individuals
have been recorded on other southern continents in recent times.

Material
Sea Harvest, Saldanha
S 1004—Fragment of a right tympanic bulla.

Description

This specimen consists of the greater part of the ectotympanic and lacks
only parts of the lateral, posterior and postero-medial margins. It is thick-walled
and inflated in a manner which is characteristic of the phocid seals and obviously
belongs to a member of this family.

It was compared with the bullae of several extant phocine and all extant
monachine genera and in size and its general configuration it is most similar
to that of Lobodon. The bulla of this, and other southern monachines, does
exhibit individual variations :n detail and although the fossil differs slightly
from the bullae of all the available L. carcinophagus specimens (n = 7), these
differences are no greater than might be expected within a single species.

Discussion

The tympanic bullae of the various phocid genera and, in the present
instance, those of the Antarctic monachines in particular, are sufficiently
distinct from one another to allow S 1004 to be confidently identified with
L. carcinophagus. This species, which is the most commonly occurring seal in the
Antarctic, has been recorded on the South African coast in recent years. The
only two records from the south-western Cape are that of an immature male
which came ashore at Gordon’s Bay near Cape Town in December 1971 and
another which was found nearby at Kalk Bay a month later. The skins and
skeletons of these individuals are preserved in the South African Museum

(SAM 36357, SAM 36358). Such vagrants are probably more common than

188 ANNALS OF THE SOUTH AFRICAN MUSEUM

the record suggests, since there is little careful reporting of seals landing on the
South African coast. Nevertheless, it may be only in certain years that vagrants
move as far north as South Africa and the number of individuals concerned
is probably never very great.

The Sea Harvest specimen may represent the remains of such a stray, but
since the chances of the natural preservation of the modern vagrants 1s extremely
remote, it is possible that the species was more common locally at times in the
past when sea temperatures were lower. This possibility is supported by the
South African fossil records of two sub-Antarctic seals (vide infra).

The inferred early Holocene age of the Sea Harvest fossils suggests that
S 1004 might indeed date from a period in time when conditions were colder
than at present, or, if not actually colder, when influences of the preceding
colder phase (latest Pleistocene) were still in operation. Since relative sea level
in the very late Pleistocene was lower than at present, the coastal environments
of that time are now all submerged so that there is no record of the locally
occurring pinnipeds and the relative numbers of the different species which
might have been present cannot be determined. Consequently, it is only the
early Holocene record which may give an indication of what pinnipeds did
occur locally during the more recent colder phases of the Pleistocene. The Sea
Harvest Lobodon does suggest that this southern monachine was more commonly
represented in the region than it is today and it was possibly still more common
during the colder phases of the Pleistocene.

Family Phocidae
Subfamily Monachinae

Mirounga leonina Linnaeus, 1758

Present status

The southern elephant seal is a sub-Antarctic species, but occasional stray
individuals have been recorded on the South African coast in recent times.

Material
Coastal midden at Ysterfonteen (Approximately 33° 21'S, 18° 9’E)
Q 1801A—Edentulous right maxilla.

Description

This specimen is indistinguishable in all observable respects from the
maxilla of adult male Mirounga leonina from sub-Antarctic regions.

Discussion

Since this species has a more northerly distribution than either the leopard
and crabeater seals, it has been more frequently recorded on the South African
coast than either of the Antarctic species (see Ross 1969).

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 189

Although the age of the Ysterfontein maxilla is not known, it has Late
Stone Age associations and because of the nature of the local coastal middens
it is most likely to date from the latter half of the Holocene (vide supra). Remains
of elephant seals are also known from an early Holocene context on the southern
Cape coast (Klein 1972), and this occurrence is more important because it is
securely dated and the material is more abundant. M. leonina still has breeding
colonies on the South American mainland (King 1964), and it is possible that
there were also permanent colonies on the South African coast during the late
Pleistocene and early Holocene. It has already been suggested elsewhere that
the decline in the range of M. leonina during the Holocene may have at least
in part been due to the warming of conditions during this epoch (Hendey

1972a).

Family Otariidae

Arctocephalus pusillus Schreber, 1776
(Fig. 42)
Present status
The Cape fur seal is still common along the coast of the south-western
Cape Province, although its numbers have declined appreciably during the
historic period.

Material

(1) Lime Quarry, Saldanha

5293A—Left mandible lacking only I,.
Isolated C and three postcanines.

5293B —Right maxilla with C and pc°.

5294 —Left humerus lacking proximal epiphysis.
Right femur lacking both epiphyses.

(2) Sea Harvest, Saldanha

S 19, S 259 to S 263, S 1009—Isolated postcanines.

S 214, S 1007, S 1008—Isolated canines.

S 1005—Skull fragment.

S 257 —Left femur lacking both epiphyses.

S 258 —Distal epiphysis of femur (not belonging to S 257).
S 1006—Metapodial lacking proximal epiphysis.

Tentatiwely referred material
Melkbos
Mb 546—Incomplete distal end of a humerus.

Comment
Reference has already been made to the Melkbos specimen (Hendey
1968), and nothing further can be added to these comments. Cooke (1955: 166)

190 ANNALS OF THE SOUTH AFRICAN MUSEUM

mentioned the Lime Quarry specimens as apparently being ‘the first record of
the Cape sea lion in the fossil state’, but gave no detailed descriptions of this
material.

Description

All the specimens listed, with the exception of the Lime Quarry mandible
(5293A), are indistinguishable from corresponding parts of the skeleton of
extant Arctocephalus pusillus.

Apart from the fact that the Lime Quarry mandible differs from that of
modern A. pusillus in certain respects, it is remarkable because it is so complete.
The missing incisor was apparently lost only recently and when it was first
discovered this specimen must have been complete in all respects.

Judging from the condition of the teeth and the character of the mandible
in general, it probably belonged to a mature adult male. For this reason com-
parisons were confined to a series of mandibles of adult males of the modern
species. he comparative series was made up of 37 specimens, all of which
belonged to individuals which had been grouped by Rand (1956) into his
categories G, H, J, K, and L (i.e. ‘bulls’). These individuals ranged in age from
young to very aged adults.

In size and morphology the teeth of the fossil fell within the range of
variation observed in the moderns, although the root of the canine was rather
large. The same applies in the case of the isolated canine and postcanines from
this locality.

The mandibular corpus, although robust, was also within the size range of
variation observed in the modern species. There was, however, a consistent
difference in the nature of the symphyseal region of the fossil and modern
specimens. The former has a prominent ‘chin’, whereas in the moderns the
anterior margin of the symphysis was generally receding. In this respect the
fossil most resembled a few aged individuals in the comparative series, but even
these specimens differed in that the margin of the symphysis posterior to the
‘chin’ was longer than in the fossil. The symphysis of 5293A is rather rectangular
in outline, but in the moderns it tends to be oval in shape. The presence of a
‘chin’ in the fossil also has the effect of increasing the surface area covered by
the symphysis, so that it is relatively and absolutely greater than in any of the
modern specimens. These differences are not clearly reflected by linear measure-
ments, although the fossil symphysis is a little longer than any observed in the
moderns (Table 40).

A feature of the mandible of modern A. pusillus which is apparently related
to the symphysis is an area of rugosity in the bone towards the inferior margin of
the corpus immediately posterior to the symphysis. This is presumably the
insertional area of a ligament attachment between the two mandibular corpora.
It is absent in the fossil. Possibly the larger surface area of the fossil symphysis
meant that the additional post-symphyseal attachment which is apparently
present in modern A. pusillus was not required.

IQgI

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

L& L& of L& LE LE LE Qz ul
G‘9S gb& ‘LP Q°6E o'h6 goz Gps of1 ofL1 sngjasnd
—G‘9E —9'S% —G*6E -9‘1f ~S‘EZ —gli -o'9b -0'6 9 -S‘a! aur snpoydas0j 247
| — |] | | |] |} TH |] ULIPOW
&‘9P 06% giab 6°bS 6°ZQ L*161 b‘o0S bor oF uvoy
06h o'be 6°0S G‘gf S‘96 10% b‘oS Cor Sp v2 662% Aven) Sunt]
ajApuoo jo G od 0} 2|qripuew 2[qipueul MOL q |
1YsIOY yysu9| I9}9UIeIp 1o1aysod jo VYysIOY jo yisuo] aurue9}sod
stsAyduihg sisAyduihg | osaoAsuety, sndio9 [[P12AQ [[R19AQ jo yasue%5 e)
Jo WYBIOH *

‘suouttoads usapour JO satis v YIM posedwioo ‘snppisnd snpvydasoja4p Aasen¢) QUI] 9y) JO aIqQrpueW pue oUTULD dy) Jo sUOIsUDUUTICT

ob alavy

192 ANNALS OF THE SOUTH AFRICAN MUSEUM

mT mg HAE AUT Hn mT ll “cc | All

§ ? & 213 24 2

]

H
|

il nn TTT I nT nl I mt nN

ml ni

An

HAN nn

mil

Fig. 42. Buccal views of the Arctocephalus pusillus mandible from Lime Quarry, Saldanha
(5293A) and a modern specimen (SAM 34640).

The most striking difference between the fossil and modern specimens is
in the greater development of the ascending ramus of the former. This is
reflected in the greater overall height of the fossil mandible (Fig. 43) and the
larger size of the condyle (Fig. 44). The general configuration of the ascending
ramus of the fossil is similar to that of A. puszllus and the coronoid process is
relatively broad and low. However, it resembles A. australis in having a deep
pterygoid process (see Repenning ef al. 1971: Fig. 7).

The degree of difference between the fossil mandible and that of modern

LATE CENOZOICG CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 193

Fossil * 5293

94

Modern Series
90

mandible

82

of

Height

74

180 184 188 192 196 200 204 208
Length of mandible

Fig. 43. The dimensions of the mandible of the Arctocephalus pusillus from Lime Quarry, Saldanha,

compared with those of a series of modern specimens.

A. pusillus is actually greater than is reflected in the metrical data presented
here, since the modern specimens to which the fossil is most similar in size are
the oldest individuals in the comparative series. Individuals from this series
which were judged on the basis of tooth wear and the thickening of tooth roots
to be ontogenetically comparable to the fossil specimen are all appreciably
smaller (Fig. 42).

The Lime Quarry maxilla (5293B) is evidently that of a female, since it is
comparable in size to corresponding parts of the skulls of modern A. pusillus
females.

15
| Modern Series I Fossil

—_
[—]

NO. OF
INDIVIDUALS

oa

TRANS. DIAM. OF CONDYLE

Fig. 44. The transverse diameter of the mandibular condyle of the Arctocephalus pusillus from
Lime Quarry, Saldanha, compared with those of a series of modern specimens.

194 ANNALS OF THE SOUTH AFRICAN MUSEUM

The postcranial material belongs to juveniles and so gives no indication
of whether or not the individuals were exceptional in size.
The Sea Harvest specimens are similar in all observable respects to modern

A. pusillus.

Discussion

There can be little doubt that the Sea Harvest and Lime Quarry specimens
should be referred to Arctocephalus pusillus, although only in the case of the
mandible from the latter occurrence need the identification be qualified in any
way.

This specimen belonged either to an unusually large individual or to a
more or less typical bull of a population in which the average size was greater
than that of the moderns. The latter alternative is favoured since it also applied
in the case of many other species investigated in the course of the present study.

Although relative size is one of the characters used in distinguishing the
modern species of Arctocephalus (Repenning et al. 1971: 31), it is not a particu-
larly useful criterion when applied to individuals owing to the considerable
range of variation in size within any one species. A. puszllus is in fact the largest
of the modern species and the Lime Quarry Arctocephalus is likely to be simply
a large temporal variant of the extant species. As such it would at most warrant
distinction at the subspecies level.

The problem of whether or not new subspecific names should be applied
in the case of large variants of extant species was one which recurred in the
present study and it will be discussed in greater detail later. The final decision
in this respect is anticipated here and the Lime Quarry material is simply
referred to A. pusillus without a subspecific distinction being made. The Lime
Quarry fauna is regarded as being late Pleistocene in age (vide supra), and the
seal is distinguished in a provisional manner by being referred to as the
‘Florisian A. pusillus’.

Arctocephalus is very abundantly represented in local Holocene coastal
middens. Those specimens which are in the collections of the South African
Museum were acquired as a result of unsystematic collecting and they represent
but a small part of what has been, or for that matter, still is available. The
specimens which have already been collected were not studied in detail and
they are, therefore, not listed.

Family Otariidae
Arctocephalus gazella Peters, 1875
&
Arctocephalus tropicalis Gray, 1872
The taxonomy of the fur seals from islands near the Antarctic Convergence

has long been problematical and there is still a diversity of opinion as to whether
one or two species should be recognized (Repenning ef al. 1971; Nel 1971).

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 195

The importance of these seals in as far as the present study is concerned is that,
like other Antarctic and sub-Antarctic species, they have been recorded in
South Africa in recent times (Nel 1971) and as post-Pleistocene fossils (Klein
1972). It is largely immaterial in the present instance whether these records
are of one species or the other, since they are all from regions beyond the
confines of the south-western Cape. However, since sub-Antarctic Arciocephalus
has been recorded on the South African coast, it is most unlikely that repre-
sentatives of the group have never landed in the south-western Cape itself.

The fact that they have not been reported locally in recent times may be
due to the fact that they would be easily confused with the indigenous A. pusillus
by casual observation. The Arctocephalus remains from local coastal middens
have not been critically examined, so it is possible that this material may
include species other than A. pusillus. It is on these grounds that A. gazella and
A. tropicalis are tentatrvely listed with the pinnipeds which did, and perhaps
still do occur in the south-western Cape.

Suborder FISSIPEDIA
Family Canidae
Subfamily Caninae
? Canis sp.
Material

Baard’s Quarry, Langebaanweg
L 1478—Left mandibular fragment with P,.

Description

This very incomplete specimen includes the intact P; and the alveoli,
some with roots of P,, P, and P,. Only those parts of the mandibular corpus
immediately adjacent to the roots of these teeth are preserved. The P, to P,
alveolar length is about 37,5 mm. The P, is made up only of the principal cusp
and measures 9,9 by 3,9 mm. It falls within the size range of variation of
modern Canis mesomelas (Table 42), and is essentially similar to the P of this
species in other respects as well.

Discussion

Since this specimen is virtually indistinguishable from corresponding parts
of the mandible of modern C. mesomelas, its identification with the Canidae is
reasonably secure and it may in fact be conspecific with this species. However,
the positive identification of fragmentary canid remains of this kind is difficult,
if not impossible.

Although this specimen may be conspecific with the poorly known
canid from ‘E’ Quarry, it is considered more likely to belong with the
Pleistocene element of the Baard’s Quarry fauna and it is listed accordingly.

196 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Canidae
Subfamily Caninae

Canis mesomelas Schreber, 1775

(Figs 47, 49)
Present status

The black-backed jackal is now extinct in the south-western Cape, but it
was apparently still common in the region during the nineteenth century.

Material
(1) Elandsfontein
(a) Previously described specimens (Ewer & Singer 1956):

EC 18 to 20, 24 to 27 and 30—Incomplete mandibular fragments.
EC 31—M,.

EC 32—P,.

EC 21— Maxillary fragment with M! and M?.

(b) Additional specimens:

(i) Skull and postcranial remains found in association and including:

17026— Part of a braincase and maxillary fragments with right I?, 1°, P* to M?,
and leit le 10) 12 and) Pio Me:

17022—Left and right maxillary fragments with P? to M?.

17000—Left and right mandibles lacking only incisors, left P, and right M3.

17001 —Right mandible lacking incisors, P; and Ms.

17021—Left mandibular fragment with P, to Mg.

16867 — Parts of a right mandible lacking only I,, and Mg, and associated with
isolated left and right P*, M? and M?.

(ii) A number of partial dentitions and isolated teeth found in association
and including:
15833 —Left maxillary fragment with P! to P®.
Five right mandibular fragments with teeth as follows:
C, P, to P,; P, to M,; P, and P,; P, and P,; P, and P,.
Four left mandibular fragments with teeth as follows:
C, P, to M,; P;, M, and M,; P, to M,; M, and M,.
Isolated carnassials and molars as follows:
P4—two; M!—four; M?—eight; M,—two; M,—one.
(i) Parts of the skull of a single individual, including:
17213—Braincase; left maxillary fragment with P! to M?.
17214—Right mandible with CG and P, to Mg.
(iv) Maxillary fragments with teeth as follows:

3361 —Left M! and M?.
5051 —Left M! and M?.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

5305 —Right P? and P* (both incomplete) and M!; left P* to M?.
5353 —Right P! to M?; left I? and P? to M?.

5448/g—Right P* to M?.

5467 —Right M! and M?.

6890 —Right M? and M?.

6891 —Right P* to M?; left P* and M!; left M1 and Mz”.

6892 —Right P* to M?; left P4.

oe “ae me * Possibly one individual.
8067 —Left M1 and M?.

8077. —Left M?.

8093 —Left P* to M?.

8110 —Right P? to M?.

9994 —Right P4 and M!.

15609 —Right P? to M1}.

(v) Mandibular fragments with teeth as follows:

3363 —Left M, and M,.

5305 —Right P, to M,; left M, and M,.
5478 —Right P, to M,.

5481 —Left M,.

5483 —Right P3.

6217 —Left P, and M,.

6863 —Right P, to M,.

6865 —Left M, and M,.

6871 —Right P, to M,.

6878 —Right P, to Mg.

7027 —Left M,.

7506 —Left P, to M,.

7525 —Left M,.

9491 —Right P, to M, (P, incomplete).
9507 —Right P,, part of P,, and M,.
11159—Left P, to Mg.

11450—Right M, and Mg.
15613—Right M, and M,.
15614—Left P,, P, and M,.
16622—Left M,.

20511—Right P, to P,.

20817—Right M,.

20999—Right P, (incomplete) and M,.
21005—Right P3.

(vi) Isolated carnassials and molars as follows:

P*—three; M!—seven; M?—five; M,—four; M,—one.

197

198 ANNALS OF THE SOUTH AFRICAN MUSEUM

(2) Melkbos
Mb 556—Right M,.

(3) Swartklip

ZW 1952—An almost complete skull, with mandible and first four cervical
vertebrae found in articulation; lacking part of the left side, but
with right side intact.

ZW 1g999—Parts of a skull with upper dentition lacking only left and right TP?
and J?, and with canines damaged.

ZW 2650—Part of a skull with right P! to M? and left P? to M?.

ZW 1998—Part of a skull with right P* to M? and left P* and M1.

ZW 102/3—Part of a skull with right C and P? and left I? to P?; associated
mandible with right dentition complete and left I, to P,.

Maxillary fragments with teeth as follows:
ZW 2953 —Right C and P? to M?.

ZW 104/5—Right P* to M?.

ZW 1856 —Right M1.

ZW 1858 —Left P4 and M!.

ZW 3243 —Right P* and M!.

ZW 3071 —Right P! to P®.

ZW 2952 —Left P* to M?.

ZW 3781 —Right M! and M?.

ZW 1314 —Left P4 and M! (SITE 2).
Right mandibular fragments with teeth as follows:
ZW 3778—I, to Mg.

ZW 2954 © to Mg.

ZW 1471—Part of C, and P, to Mg.

ZW 2998—Part of C, and P, to M,.

ZW 2000—P, to Mg.

ZW 2635—Part of P,, M, and Mg.

ZW 346 —P, to P,.

ZW 2636—P, and M,.

ZW 114 —P, and P3.

ZW 3779—P, and Ps.

ZW 2315—M.,.

Left mandibular fragments with teeth as follows:
ZW 393 —P, to Mg.

ZW 2316—P, to Mg.

ZW 1471—P, and M,.

ZW 1954—P, to Mg.

ZW 347 —M, and Mg.

ZW 107 —M,.

ZW 2318—Part of P,, M, and Mg.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 199

ZW 3254—Part of C and P,.
ZW 115 —M,.

Isolated carnassials as follows:
P4— four; M,—two.

(4) Lime Quarry, Saldanha
5308B—Left maxillary fragment with part of P* and M1.
5308D — Incomplete right M,.

Some of the Canis mesomelas specimens from this site have been lost.

(5) Sea Harvest, Saldanha

S 224 and 8 225—Skull fragments.

Right mandibular fragments with teeth as follows:
S97 —E,, P, and P, to M,.

S 135—P,.

S 728—M, and Mg.

Left mandibular fragments with teeth as follows:
S 14 —Cand P, to P, just erupting, M, and M,.
S 15 —Part of M, and M,.

S 198—M,.

S 199—P,.

S 726—Edentulous.

Isolated carnassials and molars as follows:

P4— four; M!—one; M*—one; M,—one.

(6) Coastal middens
(a) Slangkop, Cape Peninsula (34° 9'S; 18° 19’E)
Q 1803A — Parts of a skull, including:
Left premaxilla and maxilla with P* and M!; left mandible with I,
to Mg; right mandible with C, P, and P, to M,.
(b) Melkbosstrand (33° 44'S, 18° 27’E)
Q 1802A—Left mandibular fragment with P, to Mg.
(c) Ysterfonten (Approximately 33° 21'S, 18° 9’E)
Q 1801B—Skull lacking zygomata, nasals, left I’ to P? and M?, right I? to P3,
and the mandible.

Comment

Canis mesomelas is the best represented carnivore in the Quaternary fossil
assemblages from the south-western Cape and it is also the species for which the
largest comparative collection was available. Consequently, the examination of
this species was in the nature of a pilot study and some later conclusions were
based on observations made in this section of the report.

There are two extant species of jackal in southern Africa, namely, C.

200 ANNALS OF THE SOUTH AFRICAN MUSEUM

mesomelas and C. adustus, and their dental characteristics have been discussed by
Ewer (1956b). The classification of the south-western Cape material was based
on the diagnostic criteria determined by Ewer and by making use of statistical
data obtained in an independent comparative study. The two extant species
are most readily differentiated by their ‘carnassial:molar’ ratios (see Fig. 45).
Sixteen C. adustus skulls were examined in the course of the present study, while

24

=
=2,1
=
L > |
=
=

2,0

19

1,8

1,7

16

15 16 17 18 19 20 21
M, length
@ Canis mesomelas (n=63) x Canis adustus (n=16)

Fig. 45a. Lower carnassial: molar ratios of modern Canis mesomelas and C. adustus plotted against
the lower carnassial length.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 201

1,6

1,5

=

= 14
=
=
ce

1,2

13 14 15 16 17 18 19
p4 length

@ Canis mesemelas (n-72) * Canis adustus (n= 15)

Fig. 45b. Upper carnassial: molar ratios of modern Canis mesomelas and C. adustus plotted against
the upper carnassial length.

sixty C. mesomelas skulls from the Cape Province and thirteen from the northern
part of South West Africa were also available. No significant difference between
the two C. mesomelas series were observed and the two samples were treated as a
single unit. The comparative series included both males and females, with ages
ranging from young adult to aged individuals.

Description and Discussion

Owing to differences between the material from the various occurrences,
it is convenient to consider the fossil samples in descending order of age.

Coastal middens

Since the midden material almost certainly dates from the latter part of
the Holocene, it probably represents that form of C. mesomelas which inhabited
the area in historic times. None of the modern comparative specimens came from
the south-western Cape, although many were from immediately adjacent
regions. Although specimens of the locally occurring form might be preserved

202 ANNALS OF THE SOUTH AFRICAN MUSEUM

in institutions other than the South African Museum, it is most probable that
its osteological characteristics will have to be determined from fossil material
such as that from the middens.

In respect of the carnassial:molar ratios the midden specimens are well
within the ranges of variation observed in the C. mesomelas comparative series.
The most northerly of the specimens, the Ysterfontein skull, is virtually indis-
tinguishable from the skulls of the moderns in other respects as well. The
Melkbosstrand specimen differs from the moderns only in having a more promi-
nent subangular lobe. In this respect it resembles the mandible from Slangkop,
which is the most southerly of the midden records and which also has a stouter
mandibular corpus than most of the moderns.

One of the characters investigated in respect of the local Florisian C.
mesomelas was the relative breadth of the lower premolars, notably P, (vide infra).
This was found to be very variable in modern C’. mesomelas, but there was an
apparent tendency for the Florisian form to have the P, broader relative to that
of the moderns. Although it is not necessarily significant, it is worth noting that
the Slangkop P, is exceeded in relative breadth by only one of the 72 modern
specimens measured. In this respect, and also in its robust mandibular corpus
and prominent subangular lobe, the Slangkop specimen is reminiscent of the
Florisian C. mesomelas.

In itself this is hardly sufficient evidence to suggest the existence of a locally
endemic post-Pleistocene C. mesomelas population and one which was then
replaced late in the Holocene from the north or north-east by a population
which is still extant in those regions. There is, however, some evidence to indi-
cate that this might have been the case with the local Panthera leo and Hyaena
brunnea. The possibility is mentioned here in connection with C. mesomelas as
something which can be borne in mind until further evidence in this regard is
forthcoming.

Sea Harvest

The Sea Harvest specimens are essentially similar to modern C. mesomelas,
although some of the teeth are larger than those of the comparative series
(Tables 41, 42). It may be significant that the teeth of those specimens which
have a less aged appearance (e.g. S 14) are comparable in size to those of the
moderns, while the older looking specimens (e.g. S 197) are larger and more
comparable to the specimens of Florisian age. On the other hand, the assemblage
as a whole occupies a position intermediate between those of Florisian age and
the moderns in certain respects (Fig. 46) and this may be taken as an indication
that the total assemblage is of intermediate age. Since the Sea Harvest specimens
were all derived from the same shallow unconsolidated deposits, the latter
alternative is favoured.

In respect of the development of the subangular lobe of the mandible, one
Sea Harvest specimen (S 198) is comparable to the moderns and another (S 15)
is more like the Slangkop and Florisian specimens. The other mandibles belong

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 203

2,6 & Middens

e Sea Harvest

e Swartklip
oO Eft.-Bone Circle

x Elandsfontein - other

xx Transvaal

—+— Holocene
—H— Florisian

—t+— Pre-Florisian

17 18 19 20 21 22 23
M, length
Canis mesomelas — — — Canis adustus

Fig. 46. Lower carnassial:molar ratios of fossil Canis mesomelas compared with those of modern
C. mesomelas and C. adustus.

ANNALS OF THE SOUTH AFRICAN MUSEUM

204

Gin el 0z €l vl L u
E9p I-1€€ I O'7I-S‘O1 8°8-7'L SHOWS — ISei 6°6-8'L 66I-L‘91 vy-S‘€ 8°01-8°6 osuey | * (Uel[sUIOD) UroJUOJspuelg

1:8€T 911 18 €‘9l 671 6'8 6LI 0 vOl uvo

= ‘Ol SLE 891 CEI = = = = €E8SI
a OTT 9°L 8°91 S‘T1'9 == = = = ESS | ° (@[9sID ouog) urojUOyspurlq

L8r'T 80! GIL TOI 6TI 9°6 ‘ol = =a EBS
1:0S‘T L‘°6 OL OvI 601 SL EOI 8°€ S‘6 ESES | © (@UED0[O}{Z ) UloJUOJspurly

8 L 6 Ol : 9 u
189-1: Lp'T 9‘TI-€OI ¢*L-8°9 8°LI-Z'SI €€I-8'TI 8°6-1°8 TO7F'LI O's-9'€ ~=—s_—s HOI -76 SURYA Ss ieee dip11eMs

Eps‘t es O! Gi €‘O1 Ell 76 6°81 ev 0‘O! ues

= = = ae €Z1 a 76l =a = LEIS ‘907S
Ma Ha ess oS | eS : : * «4 YSOAIL FY BOS

= Zl ‘9 = STI 1°6 p81 = = LIS ‘61ZS ‘O1ZS
T6€'1 = = ist S*ZI 1°8 vLI a = WORD ai ieee oe

L:1¢'l 9°6 ‘9 Ov 6'1l L‘8 8°Ol 9°¢ 9°6 al0s1oO

TL OL tL IL TL 69 u
1:8S‘I-1:ZET 8°I I-76 0°8-L‘S L‘OI-O°Vl ~— eEI-L‘O1 ACL 8°8I-9'CI o's-+*E —_ 801-78 esuey | © spjawosaw siuvD UIspo!W

L:Sp'T 9‘O1 69 TSI SIT 1°8 VL 6€ 9°6 ura

tae q ” [ q oe I q i I q ; I

‘susutIoeds UIIPOUI JO satios & YIM posedut0d s0urIAOIg dey UsO}soM-YINOS dy} WIOIF svjamosaw sirup’ {ISSO} JO YI991 19ddn ay} Jo suOIsUsUIIC,

ib alavy,

205

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

91 oI vl Sl 61 o£ 61 rl u
7 Ol TI 617 88'T: I 6'L S0r | Ls poe. |- COC 7 Pe Jel |S Lor
-1°8 ESL -ES8'] -09'T: I =29 =O1Reo|laale =O LE | “BENZ. 0% M6) - 16'S =6% Pt Gt ee (uvI[PUIOD) UloJUOJspuRlq
0°6 6'81 1:07 SLT I Ie $6 6'L 06! 1:€7'2 Sr sol 6£ £6 uve
1 I (6 € € 6 Ol € 9 u
= = E72 96°1: I alt 66 | 86 Teo), tite" See ott.) or oO
— — SUC —68'1: 1 Cl Pa) =: rag TEO | TEBOe O'S =—8'OD | “8's: -B6 edury | (s}USWI9/9 ULISIIO]J JOYIO pur 9/d1ID oUuDg)
een eee ee UIs] UOJspurly
9‘6 L‘0@ 1:77 £61: T OL 56 5‘8 cic.) THe Oe seh I‘p 96 uvoW
_ _ a aa — ee L‘6 O17 = — — — _ OpsOW "US a sce
6 Ol Tl Ol ral 9] Ol Ol u
6'6 8°07 LPS 'z 10'2: I viL 86 | $6 Pec | Gere . Le .LIL | ee Lor
JENS = /L1 -1:07'7 a 3 | -9'9 aa ieee —'61 || “E00 “6 SOL | -1h -0 SORE oc, TS 8 ee ip neme
76 v6! [c¢% 681: 1 OL 06 L‘8 ‘IZ I:S1‘Z ose 2 rb 96 ure
= — € a ¢ € z — u
1: 0v'7 CL S‘6 0'6 1'2Z 1:07'7 9S §6 OI
a — al 4 A a -£°9 Sail Med het a! "Sat | -POoe . “Hou. 2 1 — — SOUR ly. ese ee, ISOAILH{ BIg
a = [7€Z 98'TI 9°9 L‘8 1‘8 p07 Lor‘? c's Ott —_ - urs
8°8 761 1:07'7 881 1‘9 88 8‘L r'6l 1:90°% OS: £01 Or 06 VEO8IO
[O_O FFF LT | : . : : * SUOPPIUW [vIsSvOD
78 0°81 ELU2 6L‘TI ¢‘9 76 ca 0‘07Z 1:8€°7 Le oi = — VZ0810
69 69 £9 L9 19 €9 89 IL IL 69 u
$6 L‘6l L:8€'7 SO'7 I S‘L O‘or | L'8 GOZ | [99 8 Pil) Se sor
=| SOlen =Inco] -OLT: 1 -8‘¢ “BR | =69 =—$'LT | -ES0' . Oe “06 re TL osuey [°° * * spjamosau sluDD UIOPO-!
1‘8 OLI L912 12 L‘9 88 Cz 06! 1:0€'7 S‘p = €OI ee 3's ure
TW Moreq | TW Mopeq | ATI | OLA q I q I q:I I I q I
sndio9 jo | snd1o5 jo
Yiprag WSIOH "WW WH "d ®d
*suoumtoads

UJOpOU JO soTJOs B YIM paredusos sduIAOIg 9dey Us9}s9M-YINOS Y} WIOIY spjawmosau siuD’) [ISSO} BY} JO sa|qrpueUT PUL Y}99} I9MOT ZY) JO sUOIsUDWICT

ob alavy,

206 ANNALS OF THE SOUTH AFRICAN MUSEUM

either to immature individuals or are incomplete, so the character of their
subangular lobes could not be determined.

Two specimens (S 197, S 135) have P, fully erupted and both these teeth
are broader than most of the modern specimens and compare with those of
Florisian age (Table 42). However, the incompletely erupted P, of S 14 has a
length :breadth ratio of 2,40:1 and is much narrower than the other specimens.
It is excluded from Table 42 since its dimensions may be determined by the
ontogenetic age of the individual and all the other specimens recorded are
fully grown adults.

Swartklip

The Swartklip C. mesomelas is taken to represent the typical local, Florisian
form and reference has already been made elsewhere to its relatively large size
(Hendey & Hendey 1968). Since 1968 a large number of additional specimens
have been recovered and the assemblage as a whole confirms that the Swartklip
jackal was a little larger than the moderns in most respects, although there are
overlaps in the observed size ranges of variation in the fossil and modern sam-
ples. The lower premolars of the fossil form are generally broader than those
of the moderns and the subangular lobes of the mandibles tend to be more
prominent. In other respects the Swartklip specimens are indistinguishable from
the moderns.

Saldanha Lime Quarry and Melkbos

The comparatively large size of the single Melkbos tooth has already been
mentioned (Hendey 1968). This specimen is taken to represent the local
Florisian form of C’. mesomelas partly because of its size, but largely because other
faunal evidence suggests that the Melkbos fauna is Florisian in age.

Cooke (1955) mentioned the presence of C. mesomelas in the Lime Quarry
assemblage. He gave no measurements of specimens and some of the material
seen by him has been lost. The few remaining teeth compare in size to the
larger Swartklip specimens and the Lime Quarry jackal is also taken to repre-
sent the local Florisian form of C. mesomelas.

Elandsfontein

In their original description of the Elandsfontein C. mesomelas, Ewer &
Singer (1956) concluded that it was intermediate in character between the
Makapanian C. mesomelas pappos (Ewer 1956b) and the modern form. Many
more specimens from this site are now available and the new material, together
with that from other local sites, necessitates a re-evaluation of the assemblage
as a whole.

The size ranges of variation exhibited by the Elandsfontein teeth are, in
general, greater than those recorded for the modern comparative series (Tables
41, 42), while the M,:M, length ratios overlap with the ranges observed in
both C. adustus and C. mesomelas (Fig. 46). This suggests the possibility that more
than one form is represented by the specimens here referred to C. mesomelas.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 207

HULL VL TLL LLL nm LUTTE WLLL LL

114 115 1i6

ZW 19528 *

1

mn

5 1

7 1

ae UCHLI na TUT UL

Ii

mm

mn
8 113

Fig. 47. Lateral view of the Canis mesomelas skull and mandible (ZW 1952) from Swartklip.

It has long been suspected that the Elandsfontein fossils are not all contem-
poraneous and that Cornelian, Florisian and perhaps even post-Florisian ele-
ments might be represented (Hendey 1969). It is also possible that the earlier
material is itself not homogeneous, but was accumulated over a relatively long
period of time. Consequently, the C. mesomelas assemblage was examined with a
view to establishing whether or not temporal variants within a single species
could be distinguished, since this could account for the apparent heterogeneity
of the C’. mesomelas sample and perhaps also assist in the analysis of other species
represented at the site.

No constant morphological differences between the teeth in the modern

208 ANNALS OF THE SOUTH AFRICAN MUSEUM

and fossil series were observed and it was evident that if the fossils were to be
separated into categories, this would have to be done on the basis of metrical
data.

Initially a detailed analysis was undertaken of the lower carnassials in the
various C’. mesomelas samples since this tooth is easily identifiable, it is one of the
most commonly represented of the fossil teeth and its relatively large size
facilitates an assessment of dimensional characters and their changes.

Variations in M, length in the two modern C. mesomelas samples were found
to be similar, 43,6°% of those from the Cape Province having a length of
between 19 and 20 mm, while the corresponding figure for the smaller South
West African sample was 50%. Frequency histograms of M, lengths in the
modern and Swartklip series show single peaks (Fig. 48a), although that of
the Swartklip series is between 20 and 21 mm, one unit higher than the moderns.
In all, about 82° of the moderns had M, lengths of between 18 and 20 mm,
while about 87% of those from Swartklip were between 20 and 23 mm in length.
There is, therefore, a very clear difference in the average lengths of the M, in
the modern and Florisian forms of C. mesomelas, although there is an overlap
in the observed ranges of variation and the pattern of the histograms is essen-
tially similar.

By contrast, the frequency histogram of the Elandsfontein C. mesomelas
M, lengths has two peaks and a more even distribution of measurements over
a greater number of units. The one peak between 20 and 21 mm suggests that

50

40

30

To

MODERNS SWARTKLIP ELANDSFONTEIN
M, length

Fig. 48a. Frequency histograms of M, lengths of modern, Swartklip and Elandsfontein Cents
mesomelas.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 209
n= 10

50
40
oe
20
10
16
Bone Circle, EC 26, 3363 & Remainder
5478
M, length

Fig. 48b. Frequency histograms of M, lengths of Elandsfontein Canis mesomelas.

a variety of C’. mesomelas corresponding to that from Swartklip is included in the
Elandsfontein assemblage, while the second peak between 18 and 19 mm
suggests that a second form, similar to the moderns in respect of M, length, is
also represented. The second peak is actually one unit lower than that of the
moderns, indicating that this form is not identical with the extant species, a
conclusion later confirmed by other evidence.

Included in the Elandsfontein C. mesomelas assemblage are a series of
specimens (15833) from what is known as the ‘Bone Circle’ occurrence (Inskeep

210 ANNALS OF THE SOUTH AFRICAN MUSEUM

& Hendey 1966). The preservation of the Bone Circle specimens differs from
that of most other fossils from Elandsfontein and, in addition, this assemblage
includes no extinct species. For these reasons it is thought to date from the
Florisian. In respect of M, size, the Bone Circle specimens are larger than most
others from Elandsfontein and are within the size range of variation observed
in the Swartklip form (Fig. 46), which is a further indication that the Bone
Circle occurrence is indeed Florisian in age.

A primary distinction was therefore made between those specimens from
Elandsfontein which date from the Florisian and those which do not. Since the
Elandsfontein fauna as a whole is clearly older than that from Swartklip, the
common ‘normal’-sized C’. mesomelas was regarded as part of the earlier element
of the fauna, although the possibility that it includes some post-Florisian ele-
ments was also recognized.

The separation of the C. mesomelas assemblage into categories on the basis
of M, dimensions is complicated by the fact that the Florisian variety, as
represented at Swartklip, has a size range of variation which overlaps with those
of the ‘normal’-sized moderns and the ‘normal’-sized Elandsfontein specimens.
Consequently, although three other unusually large specimens (EC 26, 3363,
5478) are tentatively assigned to the Florisian form, there might well be others
which belong to it, but which are not recognized as such because their dimen-
sions fall within the area of overlap of the various ranges of variation. ‘Corrected’
frequency histograms for the M, lengths of the Elandsfontein C. mesomelas
(Fig. 48b) still show inconsistencies. Furthermore, since the M, of the earlier
Elandsfontein form and the moderns are essentially similar in size, they cannot
be separated from one another on this basis.

The same applies in the case of all other teeth and the Elandsfontein
C. mesomelas can clearly not be categorized solely on the basis of individual
tooth size.

One other point which emerged from the examination of the lower car-
nassials is that there has apparently been a fluctuation in the average size of this
tooth through most, if not all of the Quaternary (Fig. 50). This was possibly
accompanied by changes in actual body size as well.

The Makapanian C. mesomelas pappos was found to differ ‘from living
C. mesomelas in having the lower premolars and second lower molar longer in
comparison with the length of the carnassial’ (Ewer 1956): 113). Ewer &
Singer (1956: 342; Fig. 1) indicated that the Elandsfontein form is intermediate
between the Makapanian and modern forms in this respect. They illustrated
their contention by direct comparisons of mean cheektooth lengths. Carrying
this reasoning further, it follows that the Swartklip form should be intermediate
between the early Elandsfontein form and the moderns in respect of mean
cheektooth lengths. However, it is found that this applies only in the case of M,
and the Swartklip premolars are either closer or actually equivalent to those of
C. m. pappos in length. Simple linear measurements are clearly misleading in
this instance and, although the earlier Elandsfontein form should theoretically

LATE CENOZOIC GARNIVORA OF SOUTH-WESTERN CAPE PROVINCE ZF

7

mm
8 1

NAIL UIE
3 210 211

Mm
5 |

TIAA AU
6 117 1

mm
3 1

UU

nn HU

HU

rm

mm

A LULL LL LULL LL qin TUT LLULLU LL LLL
7 8 9 110 Mit 112 113 114 115 1

Fig. 49. Buccal views of a Canis mesomelas mandible from Elandsfontein (17214) and a modern
specimen (SAM 35856).

occupy a position intermediate between C. m. pappos and the moderns, the
evidence put forward in support of this by Ewer & Singer is unacceptable in
the form in which it is presented.

A comparison between cheektooth lengths relative to the carnassial lengths
gives a different, and somewhat surprising result. The M,:M, length ratios of
the early Elandsfontein form are actually essentially similar to those of C. m.

ANNALS OF THE SOUTH AFRICAN MUSEUM

22

“SDjaUosa S1uD {ISSO} PUB UJOpour Jo sy}sug] "J JO suvsU puL sasuvs 94,1, 0G “S17

i? yyGua; ‘hw
q9G61 1394], £2 oo te Lz 02 GL aL Lt
i en en |e Tif += «a
U19}U0}x1IIS CS
jueluedeyew SUBIYIIEMS Let ==
leeIpwoly 9 =u x————————+
(Aj sea)
ueljauiog [ wlazuoyspue)q 0g =u Ga

dipyyaems = 9 = u = /————————-x

ueISI10]4 (a3e}) /
ulayuoyspue | ol =u #-———x
}SaAJeH eas ¢ =u x
9ua90|0H
SUJBpPOIN 99 =U Ym el

39v | a9v1aWassy

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 213

pappos (Fig. 46), indicating that the two forms are equally primitive in this
respect.

The Elandsfontein and Transvaal samples overlap with the M,:M, length
ratios of both the modern C. adustus and C. mesomelas series, but not with the
ratios of specimens from the Bone Circle, Swartklip and Sea Harvest. Conse-
quently, a distinction between pre-Florisian and Florisian forms of C. mesomelas
can be made on this basis, the earlier group having a ratio of 2,10 or less and the
later one 2,20 or more. This distinction is not necessarily entirely reliable, since
one of the Elandsfontein specimens (6865) has a ratio of 2,19 and could belong
in either category, or may actually represent the post-Florisian form of the
species.

The overlap with the C. adustus ratios does not necessarily indicate the
presence of this species at Elandsfontein. It may simply be accounted for by the
fact that C’. adustus and C’. mesomelas had a common ancestor and primitive forms
of either species might be expected to exhibit characters which are intermediate
between those of the modern forms. The specimen with the lowest M,:M,
length ratio (17021), which is therefore the one most like C. adustus in this
respect, was found in direct association with others (16867, 17000, 17001) in
which the ratio is well within the range observed in modern C. mesomelas. This
is taken as an indication of a single population in which the M,:M, length
ratios overlap the ranges of the two modern species, rather than that both these
species are represented.

Essentially the same pattern of similarities and differences was revealed
by a comparison of P4:M! length ratios. This ratio, which is less useful in dis-
tinguishing modern C’.. adustus from C. mesomelas (Fig. 45; Ewer 19566), actually
indicated the relationship between the fossil material and C. mesomelas more
clearly than the M,:M, length ratio. On the other hand, the distinction between
the local pre-Florisian and Florisian forms is less clearly indicated, while the
recorded C. m. pappos specimens overlap with the other forms of C. mesomelas
in this respect. On the basis of the available specimens, the local Florisian form
has a P4:M? length ratio of more than 1,45, the pre-Florisian form has a ratio
of less than 1,45, while the mean figure for C. m. pappos is 1,45.

The relatively broad lower premolars of the Florisian C. mesomelas have
already been mentioned. The length:breadth ratios of the P, of the local forms,
together with those of a series of Makapanian specimens from Kromdraai,
indicate an apparently consistent broadening trend, which is only reversed
in the modern form (Table 43). Although the P, length:breadth is very variable
in the modern comparative series, the broadening of this tooth in the fossil
populations is regarded as a real rather than apparent change, because a
graphical representation of the change (Fig. 51) matches that of the change in
M, lengths in the same populations (Fig. 50).

The relative breadths of P, also proved useful in demonstrating yet again
the heterogeneity of the Elandsfontein C. mesomelas assemblage. It was suspected
that the material included in the category ‘Elandsfontein (excluding Bone

214. ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 43

Length:breadth ratios of P, of modern and fossil Canis mesomelas from southern Africa.

P, lb
n Mean. Range

Kromdraai_. : : ; : : 4 2,360:1 , 2,32-2,43
Elandsfontein (excl. Bone Circle) : : 19 BI 2,02—2,46
Swartklip ‘ : ; : : 1&0) 2,15:1 2,07—2,17
Elandsfontein (Bone Circle) : : : 3 Orit 2,08-2,21
Sea Harvest . : : 3 : : 2 2,10: 2,00—2,20
Slangkop coastal midden . : : : I ; 2,06:1 ——

Melkbosstrand coastal midden . 2 : I 2,38:1 —

Moderns - : : ; , 71 2,30:1 2,05-2,56

Circle)’ in Table 39 might include specimens of Florisian age, so the ratios of
each individual P, were plotted on a frequency histogram and, as anticipated,
there were two distinct peaks. The first, between 2,15 and 2,20, corresponds
approximately to the mean figure of the Swartklip series, while the second,
between 2,25 and 2,35, compares with the mean of the modern series. This is a
parallel of the situation encountered when the M, lengths of the various series
were compared. Once again it proved impossible to separate the Elandsfontein
material into two distinct categories because of overlaps in the ranges of
variation. It is, however, clear that the mean figure for the early Elandsfontein
form, which is given as 2,23:1 in Table 43, is incorrect. The actual ratio for
this form is probably closer to 2,30:1 and this alternative is indicated in
Figure 51.

The possibility that some Elandsfontein specimens represent the modern
form of C. mesomelas has already been mentioned and this seemed particularly
likely in the case of the series of upper teeth, 5353. They not only occupy a
somewhat isolated position in relation to the rest of the assemblage in respect
of their dimensions (Table 41), but are also most unusual in the nature of their
preservation. They differ most markedly from other Elandsfontein specimens in
the relatively small size of the upper molars and this is reflected in the P*:M?
length ratio. Since pre-Florisian C. mesomelas is relatively primitive, the upper
molars are generally larger than those of the moderns, but the M! of 5353 is
actually near the lower size limit observed in the comparative series. It is
clearly not conspecific with the second jackal from Elandsfontein (vide infra),
although it might simply be an unusually small example of the pre-Florisian

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 215

ASSEMBLAGE AGE

Moderns
Holocene
Sea Harvest

(late) Florisian

Elandsfontein

Swartklip

Elandsfontein | Cornelian
(early)

Kromdraai | Makapanian

2,08 2,18 2,28 2,38 2,40

P, length: breadth

Fig. 51. The means of P, length:breadth ratios of modern and fossil Canis mesomelas.

C. mesomelas. On balance, the indications are that it probably does represent
the Holocene form of the species.

Non-dental characters provide little useful information on the nature of
the Elandsfontein C. mesomelas. The two braincases which are known apparently
both represent the earlier form and they do not differ significantly from those
of the moderns, although 17213 is larger and has a more prominent sagittal
crest than any of the comparative specimens. The mandible of this specimen,
and others belonging to the earlier form, all have more prominent subangular
lobes than the moderns and, in general, are larger. Judging from the length of
the cheektooth rows, the size of the mandibular corpora and the size of the
braincase of 17213, the pre-Florisian C. mesomelas was probably comparable in
overall size to the Florisian form and a little larger than the moderns. In this
particular instance the size of the M, in relation to that of the modern form
does not reflect the overall relative skull sizes of these two forms.

Summary

On the basis of metrical data the fossil C. mesomelas from the south-western
Cape may be grouped into three main categories and these are for the present
and for the sake of convenience correlated with the Cornelian, Florisian and

216 ANNALS OF THE SOUTH AFRICAN MUSEUM

Holocene. The age of the pre-Florisian material will be discussed again
later.

Depending upon the parts which are preserved, it is not always possible to
assign material to one or other of the groups and categorization of the hetero-
geneous Elandsfontein assemblage is particularly difficult. Most of the Elands-
fontein specimens are apparently Cornelian in age, but some, notably those
from the Bone Circle occurrence, date from the Florisian, while a few may
represent the Holocene C. mesomelas. It is also possible that the pre-Florisian
assemblage includes early and late Cornelian elements or even pre-Cornelian
specimens, while others intermediate in age between ‘typical’ Cornelian,
Florisian and Holocene forms might also be represented.

The two Pleistocene categories which are presently recognized are most
readily distinguished from one another on the basis of their carnassial:molar
ratios. The M,:M, length ratio of the Cornelian form is 2,10 or less and the
P*:M? length ratio is less than 1,45, while the corresponding hee for the
Florisian form are 2,20 or more and more than 1,45.

The Melkbos, Swartklip and Saldanha Lime Quarry material, like that
from the Bone Circle, represents the Florisian form of the species.

The Sea Harvest specimens are taken to be early Holocene in age and,
viewed as a whole, this assemblage is intermediate in character between the
typical Florisian form and the moderns. It may, however, actually be made up
of specimens which are Florisian in age and others which are Holocene, so that
the intermediate character results from a heterogeneous sample rather than
from phylogenetic reasons.

The distinction between the Florisian and Holocene forms is principally
a matter of a difference in the average size of individuals, the former being
larger than the moderns.

The C. mesomelas from local Late Stone Age coastal middens represent the
later Holocene form of C. mesomelas. The specimen from the most southerly of
the middens relevant in the present instance (i.e. Slangkop) may belong to an
endemic variety of the species.

The basic patterns in the nature and changes observed in the south-
western Cape C. mesomelas were found to be repeated in a number of other
locally occurring species.

Nomenclature

Although all the specimens listed earlier are referred to C. mesomelas, the
different categories can be distinguished from one another to at least some
extent. Consequently, the formal definition of distinct subspecies needs to be
considered.

In respect of the M,:M, length ratios the Elandsfontein Cornelian form is
essentially similar to the Makapanian C. m. pappos and on these grounds alone
the two forms could be regarded as belonging to the same subspecies. There is,
however, not the same correspondence between P*:M? length ratios. In addition,

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 257

the indications are that the Elandsfontein assemblage is younger than any of
those identified with the Makapanian. Finally, there is some doubt as to
whether or not C. m. pappos, as it is presently defined, should be regarded as a
valid subspecies.

Ewer (19565: 111, 112) recognized that there was ‘some heterogeneity of
the sample’ of C. m. pappos ‘since the fossils came from several sites which, on the
basis of other faunal evidence, are unlikely to be strictly contemporaneous’.
She goes on to say that, ‘The material may well include forms which are sub-
specifically distinct, or would be so regarded by a neontologist working with
living material’. To add yet another series of specimens to this taxon, and one
which is temporally and geographically removed from the C. m. pappos type
material, would serve only to complicate it further and there seems no point
and little justification for referring the early Elandsfontein material to this
subspecies. On the other hand, osteological differences between it and C. m.
pappos are such that a new subspecific name can hardly be supported by an
adequate definition.

Similar difficulties are encountered in attempting to justify a new sub-
specific name for the local Florisian form, although in this instance the problems
are mainly concerned with distinguishing it from the later Holocene form. In her
discussion on the various C. m. pappos samples, Ewer (1956a: 112) stated that
‘no useful purpose would be served by making a (taxonomic) separation which
could be based only on an arbitrary decision as to size limits’, and it is precisely
this kind of decision which would have to be made in order to distinguish the
Florisian C. mesomelas from the extant subspecies. However, since the extant
C. m. mesomelas and C. m. arenarum are more similar to one another than either is
to the Florisian form, this might be regarded as justification for affording the
latter separate taxonomic status.

Palaeontologists and neontologists may have a different basis for their
classifications, even though they may be studying essentially thesame taxonomic
units, largely because of the nature of the evidence which they have to evaluate.
In the present instance, subspecific distinctions would refer to a phylogenetic
succession which culminated in a number of geographical variants, each of
which is itself afforded subspecies status. A possible solution would be to name
subspecies according to the age with which they are correlated, thus giving a
separate name to the C. mesomelas of each of the three Pleistocene ages. The
inherent difficulties in this procedure are manifest and its advantages are
limited, since, to quote one example, ‘Makapanian C. mesomelas’ is as distinctive
and more explicit than ‘C. m. pappos’.

Since the question of the definition of new subspecies has already arisen
in connection with the Lime Quarry Arctocephalus pusillus, and since it will arise
again in connection with other species, a standard approach to the problem
was adopted during the present study. Quite simply, no new subspecific names
are proposed and the way is left open for others who may feel that trinomens
are justified and useful with some of the species described in this report.

218 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Canidae
Subfamily Caninae
Canis terblanchet Broom, 1946/1948

(Fig. 52)
Present Status

Extinct.

Materia!
Elandsfontein
15605—Left and right maxillary fragments, the latter with P! to M?.

Comment

The preservation of these specimens is atypical of the Elandsfontein
assemblage as a whole. They are part of a small association of fossils which
apparently occurred in a manner comparable to the Bone Circle association.
They are, however, encrusted with a partly consolidated calcareous sand and
have a more aged appearance than the Bone Circle fossils, although this does
not necessarily prove that the two assemblages are not contemporaneous. A
canid mandibular fragment from the same occurrence (15613) was earlier
referred to Canis mesomelas. The dental characteristics of this specimen are those
of C. mesomelas, but since the mandibular corpus shows signs of disease or severe
injury, it is possible that the development of the teeth was also affected and that
the specimen does in fact belong to a species other than C. mesomelas. Although
15613 clearly does not belong to the same individual as 15605, it may belong
to the same species, namely, C. terblanchet.

Description

The right maxilla of 15605 is almost complete and all the cheekteeth are
preserved intact. In respect of overall size (Table 44) and dental characteristics,
this specimen is essentially similar to the holotype of C. terblanche: from the
Makapanian of the Transvaal (Broom 1946, 1948; Ewer, 19565).

The similarities between this species and the extant C. adustus were noted
by Ewer (19565). Although most of the characteristics which were said to
distinguish C. terblanchei from C. adustus cannot be observed in 15605, the Elands-
fontein specimen does have a relatively broad P?, a character included in the
diagnosis of the Transvaal species. The P? and P? of 15605 are also broader
than those of C. adustus (Table 44) and, although the breadths of these teeth are
not recorded for the Transvaal C. terblanchei, they too were probably relatively
broad and in keeping with P? in this respect.

The premolars of 15605 are well spaced, as are those of the C. terblanchet
holotype.

The P*:M? length ratio of 15605 is a little less than those of the two pre-
viously recorded C. terblanchei specimens and, as with the holotype of this
species, it falls within the range of variation observed in modern C. adustus

219

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

"qgS61 IOMy x

GI V1 gI gl! GI GI G1 gI G1 u
1: 3651 oy gsi £6 Teo De ouned eo. - Tee L°'§ = G‘or bE 96 of 2G
—1:1‘1 —grii:1 | -S°6 -9SL | —Par —P‘11 bg -b't1 -3& -1'g | -gfe -&L -Siz -9'f o8ury snjsnpo siup’y
= 3 Se See ee ee ee eee ee ee ee ee ee UIIpOpy
1:60°1 09‘I: I 6‘o1 ‘9 vr Wer gL sor VE 96 of $9 An. “Pe uvoy
1:96‘! — ohio flay St Ons S24 of81 —_- — —_ — —- — 69°
——e—e——eeEeEeEEeEeEeEE————E——EE——EE——————EE—————————— ES a es ee ee 4 2YIUD] GLA} SUD!)
10 ie QS 11 Liv 92 — III — Shr ‘9 — 36 — gL oe 86S PD ad J,
UL6°% Cantus Er Ola 1 1e Orr s ort Gr, ofS: or 98 GE Oo SE 6 GogS1 UlD]UOJspuryry
sk I q I q | q I q I q I
WN rd vd‘ ed aN WN rd ed od td

‘SNISNPD *-) ULIPOU puL [eVASULIT, SY} WUOIF 1ay9IUY]GAa} *7) JO BSOY} YIM poreduro0o “1ayIUY]G4a} SUD) ULDJUOJspuL]y VY} JO YyII9} s9ddn oy} Jo suorsusWIC

bb alav yp

220 ANNALS OF THE SOUTH AFRICAN MUSEUM

(Table 44; Fig. 53). The referred specimen from the Transvaal has a P*:M}!
length ratio which is a little beyond that of the C. adustus range of the available
comparative series, but it is at the upper limit of the range for this species
recorded by Ewer (19566). It is this ratio which most clearly distinguishes 15605
from that material which was earlier referred to C’. mesomelas.

Discussion

On the basis of the material presently available, there are no grounds for
believing that 15605 and C. terblanche: are not conspecific. There are, however,
grounds for doubting whether C. terblanchet warrants recognition as a species
distinct from C. adustus. It may well be a primitive form of C. adustus, just as
C. mesomelas pappos is a primitive form of the extant black-backed jackal.

The most striking characteristics of C. terblanche: are the prominent sub-
angular lobe of the mandible and well-developed paramastoid process. Ewer
(1956b) recognized these to be related features since the occipito-mandibularis
muscle is inserted at the subangular lobe and originates at the paramastoid
process, so the development of these osteological features is related to the
development of this muscle.

HA il

||| Fig. 52. Occlusal view of the
Canis terblanchei maxilla (15605)
from Elandsfontein.

fy NHI THI

A prominent subangular lobe is a feature of extant canids such as the
crab-eating fox (Cerdocyon thous), the bat-eared fox (Otocyon megalotis) and the
raccoon dog (WNyctereutes procyonoides), that is, species which are somewhat
atypical of the Canidae in dietary and other respects. On the other hand,
it is also well developed in less remarkable species such as the New World grey
fox (Urocyon cineroargenteus). In addition, it is a feature of the Cornelian and
Florisian C. mesomelas and Vulpes chama from the south-western Cape. In none
of the South African fossils is the subangular lobe quite as prominent or as
posteriorly situated as it is in the modern species mentioned above. The develop-
ment of this feature, and those directly related to it, in the fossil forms is regarded
as a reflection of the differences between their dentitions and those of their
modern counterparts and perhaps relate to size and/or dietary differences.

221

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

‘SNISNPD *F) pUe SDJaWOSAU *-) UIOPOUL puUv [elIIo}eUI adA} 2949UD]GQ44} “7 24 JO 2804} YIIM poredur0s
SD]aUOSAUL *7) PUB 1949UD]GLa] SUD) UIDYUOJspuUL][Y IY} JO soNvi Iejour:yeisseures isddy °fS “S17

sejawosaw = siueyg — — — snjsnpe siue4)

yy Gua yd
02 61 8 l Lt 91 Sl bl El

ula}uUOjspue,y — Sejawosawl “y ¥

Zl
jeeasuels | - rT uo @
ula}uoyspue,Z - 1ayauejqia} “9 © a
a
Ben, Te e 6 | =
a
Sa * & =
He ¥ 5 x * 3
¥ ‘ >
¥ * A
-
x
x
cl

222 ANNALS OF THE SOUTH AFRICAN MUSEUM

There apparently is a connection between the nature of the cheekteeth
and the presence or absence of the subangular lobe in other groups of carnivores.
For example, the Felidae with their highly specialized shearing dentition lack
the subangular lobe, whereas the Hyaenidae with their broad and heavy
crushing teeth have this feature well developed. In the fossil Canidae the sub-
angular lobe may relate directly to broader lower premolars (e.g. Florisian
C. mesomelas), broader upper premolars (e.g. C. terblanchet), relatively prominent
posterior cheekteeth (e.g. Cornelian and Florisian V. chama—vide infra) and
other such dental characteristics.

The analysis of the mammalian masticatory apparatus is a complex
matter requiring detailed study (see Turnbull 1970) and the present specula-
tions made in connection with the development of the subangular lobe are not
necessarily desirable or useful. Nevertheless, the evidence of the local fossil
record suggests that its development, and those of directly related features,
could vary within a single lineage over relatively short periods in time and that
its presence does not necessarily indicate only a distant phyletic connection to
an otherwise similar form in which it is not developed.

Although C. terblanchei may be regarded simply as a less advanced form of
C’. adustus, its status as a separate species is retained. This is partly because
C.. terblanchei is relatively poorly known and its relationships are therefore more
likely to be misinterpreted, and partly because it was found convenient to
retain the names of two other Makapanian canid species (Vulpes pattisoni,
V. pulcher) and for the sake of consistency C. terblanchei was treated in the same
way. Ewer & Singer (1956: 345) state that the ‘exact point (at) which pro-
gressive change is taken to be of sufficient importance to warrant nomenclatural
recognition must be largely a matter of individual judgment.’ C. terblanchei
has been judged to merit separate species status and there is little point in
suggesting a change based on another subjective opinion.

Referral of 15605 to C. terblanchei rather than the more advanced C. adustus
is based largely on the relative breadths of P! to P%, in which respect it is appa-
rently similar to the C. terblanche: holotype and is definitely different to modern
C’. adustus.

The identification of a jackal in the Elandsfontein assemblage which has
teeth resembling those of C. adustus raises yet another problem with that material
referred to C. mesomelas. The teeth of the two modern species may be indis-
tinguishable from one another (Ewer 1956): 97, 98), and it is therefore
possible that some of the specimens classified as C. mesomelas actually belong to
C’. terblanchet.

Relative age

It has already been indicated that the Transvaal C. terblanche: is Maka-
panian in age, but this is actually not certain. The holotype was originally
reported to be from Kromdraai (Broom 1946), but apparently this is unlikely
and its source is unknown (Ewer 19560). The referred material was said to be

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 223

from Coopers (Ewer 1956d), although subsequently Ewer (1956c: Table 2) did
not list it as part of the carnivore fauna from this site. Whatever the actual source
of the Transvaal specimens, they are almost certainly from the breccia deposits
in the Krugersdorp area and they are most likely to be Makapanian in age.

Since there are no significant observable differences between 15605 and
the type material, it follows that they might be broadly contemporaneous.
The unusual preservation of 15605, and the specimens with which it was
associated, may be an indication that this material differs in age from most
other specimens from this site. The fauna associated with 15605 has not been
investigated, but there is nothing obvious which would suggest it is of Maka-
panian age. The mandible from this occurrence earlier identified as belonging
to C. mesomelas (15613) has dental characteristics which indicate that it could
belong to either the Makapanian or Cornelian form of this species. Conse-
quently, it is possible that some elements in the Elandsfontein assemblage pre-
date the main Cornelian fauna. They could date from the Makapanian or
perhaps from the earlier part of the Cornelian.

Taken in conjunction, the Elandsfontein jackals suggest that the fauna
from this site is made up of elements aged as follows:

(1) A few specimens may be Makapanian or early Cornelian.
(2) The bulk are Cornelian.

(3) Some are Florisian.

(4) A few may be Holocene.

The only real departure from previously held opinions is that the fauna
may include some specimens which are earlier than the main body of fossils.
The evidence for this is very slender and it is probably preferable and certainly
more convenient at this stage to regard all the early fossils as Cornelian in age.
Until there is really convincing evidence to the contrary, the only age categories
recognized in the Elandsfontein fauna are, in descending order of importance,
Cornelian, Florisian and Holocene.

Family Canidae
Subfamily Caninae
Vulpes chama A. Smith, 1833
(Fig. 54)
Present status

The Cape or silver fox is now very rare in the south-western Cape Province,
although it was formerly common.

Material

(1) Elandsfontein

21007— Parts of the skull and skeleton of a single individual, including:
Incomplete braincase; left maxillary fragment with P® to M?; right

mandible with C to M,; left mandible with I, and P, to M,.
Two vertebrae and parts of all four limbs.

224 ANNALS OF THE SOUTH AFRICAN MUSEUM

5474, 5477, 5472—Right maxillary fragment with P*; left maxillary with P®
and P*; isolated Ml.

5457, 5469—Right maxillary fragment with P?, part of P*4, M! and M?.

8073, 8114—Right and left M?.

Mandibular fragments with teeth as follows:

5093 —Right P,.

5453 —Right P, (incomplete), P;, P, (incomplete) and M,.
5455 —Right P, to M,.

14244—Right M,.

20022—Right M,.

20023—Left M,.

(2) Swartklip

ZW 1894—Left maxillary fragment with P*.
ZW 2317—Right mandibular fragment with P, and M,.

(3) Sea Harvest, Saldanha
S 39—Right P*.

Comment

As with Canis mesomelas, the material here referred to Vulpes chama is
heterogeneous and at least two forms are represented.

Description
Sea Harvest and Swartklip

The Vulpes specimens from these sites are too few and fragmentary to allow
for adequate description of the form which they represent but, viewed in
relation to the material previously referred to C. mesomelas and to other fossil
and modern Vulpes specimens, their affinities are reasonably clear.

Of the four teeth represented, only one, the P, of ZW 2317, falls within
the observed ranges of size of modern V. chama (Table 46). The M, of ZW 2317
is slightly broader than the broadest M, in the comparative series, while the
upper carnassials from both sites are longer than any of those of the available
V. chama specimens (Tables 45, 46).

Morphologically the fossil teeth are essentially similar to those of the
moderns, although the Swartklip M, differs in having a relatively larger
talonid. This tooth is broadest at about the midpoint of the talonid, rather than
near the posterior end of the trigonid as in modern JV. chama. The size of the M,
alveolus indicates that this tooth was relatively large. The large M, together
with the size of the M, talonid suggests that the upper molars of the Swartklip
Vulpes must have been correspondingly larger than those of the modern species.
This is indeed confirmed by the sizes of the M1 and M? alveoli of ZW 1894.

Consequently, it appears that while the premolars of the Swartklip Vulpes
were little or no different from those of modern V. chama, the carnassials and

225

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

"qQS61 IOMT »

1:96°1 — g‘ol Lea 1G O11 oe Oat — _— £g*y
OS a ee ee oe oe ee eee Sa ee Se (eee eee ee eee 4 1ayIUD] GLa} SUD‘)
Teeth gG‘1: 1 Hat gil — III — GHr9} — a6 adA qT,
et &% 1G BB bz £% u
1:96‘ GL‘ti1 £'6 £9 G11 bg zg Ss S‘o1 ht 2 ocZ puDya
—1: G11 —Ghit:1 | —bZ -6°F | -&*6 -1‘L | -6°F -€6 a —G‘¢ a2ury saqin4
a a |e ulopoyy
1: L5‘1 gS‘: bg 9‘S g‘Ol LL gS 96 C% 29 uv
eee moe ae = —_ = 1‘9 III as = 6§ S JSIAIVFT 89S
= — — — — _ og girl ~~ — b6g1 MZ dippjzemg
1:G6°1 QQ‘I:1 _ — GSI o‘O1 Lig =S*E1 ca 3 1° LL/aLvsS
a a ee ee ee ee ee ula}UOJspurly
1:S3‘1 gQ‘I: 1 S01 9 POD) PEI g‘O1 ob Pred GE 0g Loolz

“‘DUIDYI “A UIOPOUI JO SaIJas & PUL 21979UD]gG49] StUD’) JO BSOY} YIIM poieduiod fader) u19}saM-YINOS 9Y} WOT] DIUDYI Sagjny {ISSO} JO 4199} 1addn 9y} Jo suoisuswIg,

Ch alav

ANNALS OF THE SOUTH AFRICAN MUSEUM

226

“GQ9S6] OMA x

73°
odA]
u

osuPy

uvoy
LIEC MAZ
€60S
£7007
CCOO?

IevIPWOIy

SULIxIIEMS

= o/h = L‘9 68} 89 gS — — a = = =
16€ Eel‘ LVI: CHL, IG I We) 2 [ESI laSS v6 TP L‘Ol S‘€ 96 | €€
8 €7 €7 €Z €7Z €7Z €7 €7
9'L@ 1:86'1 CLT: I GG OH | OG Seal | Gee ON 18 | 67 GS | Sire
=6, 62 -1:79'T BPTI -0'p =O | ay =(OL | SUG He =(SO) | LEE JES || L0Ve6
LES L08‘T INT L‘v 9 8‘P SII L177 (645 el SE CO" CC
= ae LLY: = = 8‘s y‘ZI 1977 I‘€ OL = = os
cae — = = = a = L:€7'7 0'v 68. <— = =>
= es = = = Va) rrl == ca = = == ==
C80 2 az a = | 69 Ist =, ae oes jie See \ ar
0°67 “2 = (6 a a 79 E'S Me ee g‘€ p‘8 €‘€ 9‘L I‘€
PLZ “2 L:9L'T OST: I ¢‘9 88 | 69 CSI 1:97Z g‘¢ 9°8 I‘€ Gi | Oe
1°67 I: L6‘T 99'T: 1 a [an ORS | 1:€9°7 TE v8 S06 (EE I 17
9°97 1: 10°Z POT WA? 09 | 9b ral! LE 1E'7 Gs Ph | WG ROM |e exc
l’d-'d I I q I q I qi 1 q I q I q
TA TA TA 'd NI WA *d ®d

‘DULDYD *A UIOPOUL JO SoIi9s & pue [eeASULIT,

v8 v'7
(WG
b'9 61
C6 =5'|
aS [
OL =
Bi98 2) ||| SENG
ee 6'I
79 9‘T
- I q

4 10YOUD]GAa}
siupg

puipyd sadjn4
ulOpOo|

dipsems

UIoJUOJspur|y

44aY4ojnd sadjn{

2Y} WIOL 2ayIUD]gQ4a] SluD- pu saying ‘4 JO 9sOy) YUM posedurod nwuvys sadjng AipyeMG pue uUla}UOJspuLTy IY} JO YII9} JIMO] 9Y} JO suOIsUIWIC,

gh alav],

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 227

HL fi
110 i |

om

3 1

as

mm
5 1

mm

7 1

Il
8

Fig. 54. Buccal view of the Vulpes chama mandible (21007) from Elands-
fontein.

molars were appreciably larger. A possible exception is the Mg, the size of which
is not known.

That part of the mandibular corpus of ZW 2317 which remains is poorly
reserved, but it is clearly much more robust than the slender corpora of modern
V. chama. In addition, it has a well-developed subangular lobe, which is in
marked contrast to the mandible of the modern species.

The Swartklip and Sea Harvest Vulpes are evidently similar to the C.
mesomelas from these sites in that they are larger than their modern counterpart.
The Swartklip form also differs from the modern species in respect of the relative
sizes of certain teeth, a fact that will be enlarged upon later. It has already been
suggested that the Sea Harvest fauna is intermediate in age between that from
Swartklip and the present, and interestingly the Sea Harvest P* is intermediate
in size between that of ZW 1894 and those of the modern comparative series
(Fig. 55). In view of the small number of specimens involved this is not neces-
sarily significant, but at least the small size difference between the Swartklip
and Sea Harvest upper carnassials is in the right order.

Elandsfontein

The Elandsfontein material belongs to a canid with a skull comparable in
size to that of modern V. vulpes. Superficially it differs appreciably from both
V. vulpes and V. chama and initially it was thought to belong to a small jackal
rather than a large fox. The most important osteological character which
distinguishes the skulls of jackals from those of foxes, the inflated frontal sinuses
in the former (Huxley, 1880), cannot be observed in any of the specimens
presently available and their relationships had, therefore, to be determined
on other grounds.

228 ANNALS OF THE SOUTH AFRICAN MUSEUM

Breadth

) 10 HW 12 13 14
Length

@ Modern Vulpes chama —»> Decreasing age

0 Sea Harvest
O Swartklip

* Elandsfontein

Fig. 55a. Dimensions of the P* of modern and fossil Vulpes.

Most of the observations which follow are based on the specimen 21007.

This form is clearly not conspecific with either of the canid species
already described, although in respect of actual size of the posterior cheekteeth
and the carnassial: molar ratios it is similar to C. terblanchei. The pre-carnassial
teeth of 21007 are, however, appreciably smaller than any of the recorded
fossil and modern jackals of southern Africa, a character best illustrated by the
P,:M, length ratio and P, to P, length. The contrast in the sizes of the anterior
and posterior cheekteeth is reminiscent of the Swartklip Vulpes, although the
actual size of the individual teeth of this form is appreciably less. In addition,
the Elandsfontein Vulpes is smaller in overall size than any of the local jackals,
but in this respect it is similar to fossil material from the Transvaal which is
referred to a species of Vulpes, namely, V. pulcher from Kromdraai (Broom 1939).

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 229

Breadth

10 11 12 13 14 15 16
Length

e Modern Vulpes chama —> Decreasing age
oO Swartklip
* Elandsfontein

OO Kromdraai Vulpes pulcher
& Swartkrans

Fig. 55b. Dimensions of the M, of modern and fossil Vulpes.

The Vulpes which have been recorded from the South African Quaternary
are, in descending order of age, as follows:

(1) V. pattisoni from Taung (Broom 1948), a small species comparable in size to modern VP.
chama. It is poorly known and largely excluded from the discussions which follow.

(2) A specimen from Swartkrans belonging to a species intermediate in size between V. chama
and V. vulpes and which was referred by Ewer (19560) to V. pulcher.

(3) The Kromdraai V. pulcher, which is in most respects a little larger than the Swartkrans form.

(4) Specimens which are Florisian in age and which are referred to V. chama (see Cooke 1963:
Table 7), including those from Swartklip.

(5) Modern V. chama.

On the assumptions that these five units represent different stages of a
single lineage and that the Elandsfontein specimens are Cornelian in age, it
follows that 21007 should conform to the sequence in a position intermediate
between the V. pulcher holotype and the Florisian V. chama, which is represented
locally by the specimens from Swartklip. The characteristics of 21007 were
investigated with this in mind.

There was apparently an increase in size with time in the Makapanian
forms of Vulpes from the small V. pattisoni to the large V. pulcher holotype, with
the Swartkrans specimen in an intermediate position. This was followed by a

230 ANNALS OF THE SOUTH AFRICAN MUSEUM

decrease in size with time with the Florisian form being intermediate between
V. pulcher and modern V. chama. The length of the lower premolar row of
21007 is similar to that of the V. pulcher holotype, while the molars are longer.
This indicates that the early Pleistocene size increase was continued in the
Elandsfontein form at least in respect of the molars, and perhaps in overall size
as well. It is, therefore, not out of place in the hypothetical phyletic series in
respect of size.

The greater development of the molars relative to the premolars has
already been mentioned in connection with the Elandsfontein and Swartklip
specimens and, where possible, the P,:M, length ratios were compared in
order to determine the nature of changes in this respect. This ratio was found
to be quite variable in the modern form, but, assuming that the fossil specimens
represent fairly typical examples of their respective populations, it is apparent
that there was a trend in the relative development of these two teeth which was
consistent with the inferred overall size of individuals in the various populations
(Table 46). The largest M, in both a relative and absolute sense is that of the
Elandsfontein form.

The changes in the M,:M, ratios of the fossil series also appear to be
consistent, but in this instance the M, with the greatest relative length is
apparently that of the Swartklip form. The estimated M,:M, length ratio of
ZW 2317 is calculated to be below 1,60:1 and is, therefore, less than the mini-
mum observed in the modern comparative series. The trend towards the
lengthening of M, thus appears to have been reversed in the Holocene.

In at least one respect the V. pulcher holotype is unlike any of the other
specimens. The relatively narrow premolars of the Kromdraai specimen were
remarked upon by Ewer (19560) and its difference to the other forms in this
respect is clearly illustrated by the P, length:breadth ratios recorded in ‘Table
42. There is no detectable trend in the fossil series in the relative breadths of
the premolars.

Upper dentitions are either unrepresented or less well represented in the
fossil series, but they presumably underwent changes comparable to those in
the lower teeth. The relative sizes of the Elandsfontein, Swartklip and modern
upper carnassials are certainly similar to those of the lower carnassials and the
same apparently applies in the case of the upper molars. However, the P?:P#
length ratio of 21007 does not differ from the moderns in the same way as the
P,:M, length ratio. As a general rule, the upper teeth of the Canidae appear
to be less useful in distinguishing species than the lowers.

There are yet other indications of a relationship between the Elandsfontein
and Swartklip specimens. The relatively large M, talonid of ZW 2317 was
mentioned earlier and, although none of the lower carnassials from Elandsfon-
tein has the talonid quite as prominent, in four of the five known specimens it is
the broadest part of the tooth. This may indicate a development in the M, of
the Elandsfontein form which is consistent with the trend in the fossils towards
an increase in the size of grinding surfaces on the posterior cheekteeth.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 231

Another feature of ZW 2317 is the prominent subangular lobe of the
mandible. It is also well developed in the only three specimens from Elands-
fontein in which the posterior part of the mandibular corpus is preserved. The
three specimens belong to a young adult (20023), a mature adult (21007) and
a very aged individual (5453). The nature of the subangular region in V. pulcher
is not recorded.

The available evidence, limited as it is because of the small number of
specimens involved, suggests that there are grounds for regarding the South
African Quaternary Vulpes as belonging to a single lineage.

The fragmentary braincase of 21007 is comparable in size to corresponding
parts of the skull of V. vulpes and it resembles this species in having a fairly
prominent sagittal crest. It differs from the skull of modern V. chama in its
larger size, more prominent sagittal crest and less inflated braincase.

The body size of the Elandsfontein Vulpes can be gauged from a comparison
between limb bone dimensions of 21007, modern V. chama (1 male) and modern
C. mesomelas (1 male, 1 female, 1 sex unknown) (Table 47). It was clearly a
species which was much smaller than C. mesomelas. The individual limb bones
of 21007 were even a little shorter than those of the V. chama skeleton, although
the limb segment ratios are similar (Table 48). The fossil bones are, however,
more stoutly proportioned, indicating that although it was comparable in
stature to modern V. chama, it had a heavier body. The more robust body was
coupled with a head which was larger than that of the modern species.

Discussion

Having suggested that those fossils from South Africa which are referred
to Vulpes probably represent a single lineage, parallels can be drawn between
the nomenclatural problems involving this group and those which were dis-
cussed in connection with the jackals.

Little can be said of V. pattisoni because it is so poorly known, but V. pulcher
is like C. mesomelas pappos in that it includes material from more than one site
and is a taxon comprised of samples which are heterogeneous in a temporal
sense. The differences between V. pulcher and modern V. chama appear to be
more marked than those between C. mesomelas pappos and modern C. mesomelas.
This is largely due to the greater size of the V. pulcher specimens relative to the
moderns, since other observable differences are probably no greater in the foxes
than they are in the jackals. The fact that the fossil fox is afforded full species
status and the jackal only subspecific status thus reflects an inconsistent approach
to the problem of their taxonomy.

In respect of differences between the Elandsfontein Vulpes and modern
V. chama much the same applies. In some features (e.g. the actual size of
posterior cheekteeth; P,:M, length ratio) they are even more different from
modern V. chama specimens than is the material referred to V. pulcher, while in
others (e.g. the P, length:breadth and M,:M, length ratios) they are closer to
the modern specimens. Once again, excluding differences in actual skull size,

ANNALS OF THE SOUTH AFRICAN MUSEUM

232

0°91

8°77
eve
9°%
86!

ma

€‘LI

181
OLI

as

v'LI

9%
i 4

IZ

£‘81

10% O'OI 009SE N'VS DULDYI
sadjn,4
p87 S‘EZI | 5 P7BSEWVS
Cue €‘671 TSZIE INVS | SDJamosoum
SS en | neers ee Par siuDy
ZOE ose! PETBSE NWS
fice o‘Sol LOOIZ ul9}UOJ
-spurly
~
cq
o
x
(eo)
we
Q,
Ge
°
& | 4
Vv q
roy 2
4 =
re =
a >
= fe)

s‘7I | 98 p'6l | €‘L7I €‘LI €‘6l | VL £6 L‘s P901
gst | S‘Il | 67 | O'SHI 817 8‘Sz TLI 88 Oe! 98 TEI
List | TIL | ofsz | O'ISI (KE, L‘6z OSI 16 671 6'8 I‘Z€l
ESloleSeliet bee |) OST 617 £97 L‘9t £6 ¢‘7I ‘8 0‘OrI
ZEL | OCOL | 9'07 | O9TI 761 (GME O'rl S‘L €‘Ol 69 L‘t01
Me) ~ fC ~ ue)
8 ~ § oO ie] 5 ue) 5 =
Bol ce). 28 c z ig Z 6 S
2 nes : 2
paca OR) ae cee) ar ie eee
E Pee g E é
a a & a A
Bees ie | | 8 4 fs oe) ee |g
CBee eae ig 4 3 d é: d 3 §
())
Cee seer le ee PO awe |e
Fy ‘1 A 3 A i | A 6 . ow
5 5 5 8 4 5 a a a as 9
= = = e) => PS > 2 =, PD o)
RIqLL Inwo,{ snipey

"pupys ‘A Uapow suo puL
SDJAWOSAUL SIUDT UIZPOUI 9914} JO 9SOY} YM poredui0d UlojUOCJspuL]yY WoIy DUDYI Sagjng DY} JO UOJI[9¥s JeIUVID\sod oY) JO syUDWI[9 JO suOIsUDUNIC

Lb alavy,

snJouin fy

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 233

TABLE 48

Limb segment ratios of the Vulpes chama from Elandsfontein,
compared with those of modern V. chama and Canis mesomelas.

Elandsfontein 21007

Vulpes chama 83,6

89,0

Canis mesomelas

the contrasts between the Elandsfontein Vulpes and modern V. chama are proba-
bly no greater than those between contemporary populations of C. mesomelas.

In the case of Cornelian and Florisian C. mesomelas, distinctions were made
on the basis of relative tooth sizes (i.e. the P4:M! and M,:M, length ratios
differ), but not actual skull size. The Cornelian and Florisian Vulpes are dis-
tinguished mainly by actual skull size and not relative tooth sizes (e.g. P,:M,
length ratios are essentially similar). The differences between Florisian and
Holocene C. mesomelas were largely a matter of actual average size. Florisian
and Holocene Vulpes also differ in this respect, but in addition there
are differences in relative tooth sizes as well (e.g. P,:M, length ratios).

The fossil Vulpes therefore contrasts with C. mesomelas in that it apparently
underwent more marked changes in size, while the nature of the changes in the
dentition were not the same. For example, a relatively long M, was a charac-
teristic of early C. mesomelas, but developed to an extreme only in Florisian
Vulpes. In addition, changes in dental characteristics were not synchronous in
the jackals and foxes. A graded classification of fossil C. mesomelas could con-
veniently have distinguished between the Florisian and pre-Florisian forms,
each group perhaps being given species status (i.e. C. pappos for the pre-Florisian
group and C. mesomelas for the Florisian group). This would, however, have
served no useful purpose because of the impossibility of distinguishing Holocene
C. mesomelas from the two fossil categories.

By contrast, a graded classification based on the dental characteristics of
fossil Vulpes could conveniently distinguish between the Makapanian forms
(V. pattisont and V. pulcher), a Cornelian/Florisian form (Vulpes n. sp.) and the
Holocene V. chama. In this instance so few fossil specimens are involved that
definition of the four categories would present no real difficulties, although the
recovery of additional specimens could well render the definitions impracticable.
One difficulty in this categorization is that when the actual size of individuals
is taken into account, it is found that the Cornelian Vulpes is actually closer to
V. pulcher rather than to the Florisian form. In other words, the most readily
evident feature of the fossils (i.e. skull size) does not conform to a categorization
based on dental characteristics.

234 ANNALS OF THE SOUTH AFRICAN MUSEUM

If a new species name were to be given to the south-western Cape fossils,
the definition would of necessity have to take into account the heterogeneous
character of the Cornelian and Florisian specimens. Consequently, instead of
defining a heterogeneous new species, it might be preferable to refer the fossil
to an existing species and to extend its definition to include the characters of
the Elandsfontein and Swartklip material. In this event, the local fossils could
be referred to either V. pulcher or V chama. This material is in fact here referred
to V. chama because the Swartklip specimens are closer to the modern species
in morphology and time than the Elandsfontein specimens are to V. pulcher.
In the final analysis, this decision is probably as subjective as any other in
taxonomic problems of this kind.

The fossil populations of C. mesomelas were distinguished from one another
by prefacing them with the age name to which they belong (e.g. Cornelian
C’. mesomelas), and this is also done in the case of the fossil V. chama from the
south-western Cape. It serves to indicate that the Cornelian and Florisian forms
differ from one another, that both differ from V. chama, but that all are part of
the phyletic series which culminated in the extant silver fox.

While the naming of the local fossil Vulpes adopted here may not be an
ideal solution to the problem of nomenclature, it is regarded as simple and
convenient. Once again the way is left open for others who feel that new species
and/or subspecies names are more desirable.

As with C. mesomelas and C. terblanche1, the existing names of the Maka-
panian forms are retained. In this connection it is worth noting that there is a
precedent for distinguishing early Pleistocene foxes at the species level. In
Europe each of the extant species is preceded by only one Pleistocene fossil
species. V. alopecordes may be ancestral to both V. vulpes and Alopex lagopus, while
V. praecorsac is ancestral to V. corsac (Kurtén 1968).

Family Canidae
Subfamily Simocyoninae
Lycaon pictus Temminck, 1820

(Fig. 56; also Ewer & Singer 1956: pls 30, 31)

Present status

The hunting dog is now extinct in the south-western Cape, but it apparently
still occurred in the northern parts of the region in the eighteenth century.
Material
(1) Elandsfontein

(a) Previously described specimens (Ewer & Singer 1956):
EC 12—Right mandibular fragment with I,, I; and C. (Note: Number given
as EC 121 in the original description. )

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 235

EC 13—Left mandibular fragment with P, to M, (Note: The M, was added to
this specimen subsequent to its description. )
EC 30—Incomplete right M,.

(b) Additional specimens:
EC 37—Right mandibular fragment with roots of cheekteeth.
6701 —Right mandibular fragment with parts of P, and P,, Ps, part of Py,
and M,.
17058 —Right mandibular fragment with part of Mg.
20439 —Right mandibular fragment with parts of C, P,, P; and P,.
9194 —Part of right P*.

(2) Swartklip

ZW 137 —Right I?.

ZW 177 —Part of left I?.

ZW 2320—Left M?.

ZW 3070—Left I3.

ZW 2314—Left mandibular fragment with part of C.

Description
Elandsfontein

In their description of the Lycaon remains from Elandsfontein, Ewer &
Singer (1956: 341) noted the following differences between the modern and
fossil forms:

(1) ‘the fossil jaw is considerably longer than that of the extant species’.

(2) ‘the premolars are considerably longer, but not much broader’.

(3) ‘the canine and I, are significantly larger in both dimensions while the significance of the
slightly greater breadth of I, is less certain’.

On the basis of these differences the material was referred to a distinct sub-

species, Lycaon pictus magnus.

The discovery of additional specimens of this species allows for amplifica-
tion and modification of these observations.

Although it was stated that, ‘Apart from their greater length the teeth
differ in no way from those of the living Lycaon pictus’ (Ewer & Singer, 1956:
341), there are other differences in some of the teeth. The most striking of these
is the absence or small size of anterior accessory cusps on the P, to P, of the
fossil. In the available modern L. pictus specimens (n = 4) the anterior and
posterior accessory cusps are variably developed and are progressively less
prominent from P, to P,. They were, nevertheless, always present, although the
anterior accessory cusp of P, was very small. In EC 13 this cusp is barely dis-
cernible on the P; and P,, while it was not present at all on the P,. The anterior
portion of the P, of 6701 is lost, but in the P, the anterior accessory cusp is
absent and in the P, it is very small.

L. p. magnus also differs from the modern form in having a relatively
smaller M, metaconid. In addition, the three known lower carnassials of the

236 ANNALS OF THE SOUTH AFRICAN MUSEUM

rc

Fig. 56. Buccal view of the Lycaon pictus mandible (EC 13) from Elands-
fontein.

fossil form have a small ridge on the lingual surface of the talonid which is not
present in any of the available modern specimens. .

There is a curious anomaly in the length/breadth ratios of the lower
cheekteeth of the Elandsfontein Lycaon. The premolars of EC 13 are narrow
relative to those of the moderns, but the carnassial is comparable in width
(Table 49). The difference in the P, to P, of EC 13 is due largely to the lack of
inflation of the postero-internal cingula of these teeth. This inflation is evident
in the premolars of the available modern specimens as well as the fossil 6701.
In the latter specimen it is the M, which is unusual, since it is relatively broader
than the lower carnassials of EC 13 and the extant form. The modern species
thus has premolars with length/breadth ratios comparable to 6701, but the M,
length/breadth ratio is comparable to that of EC 13. Many more modern and
fossil specimens are required before the significance of these similarities and
differences can be assessed, but the earlier comment by Ewer & Singer on the
breadth of the premolars of L. p. magnus is invalid in view of the nature of these
teeth in 6701.

Standard measurements could not be taken on the incomplete P* (9194),
which is the only known upper tooth of the Elandsfontein Lycaon, but it is
appreciably larger than any in the comparative series.

Swartklip

The Swartklip mandibular fragment (ZW 2314) has a very slender corpus
and probably belonged to an immature individual. The preserved canine and
the alveoli of the cheekteeth compare in size to those of the available modern
specimens.

The M, (ZW 2320) also falls within the size range of variation of the
modern species. It measures 15,8 by 16,2 mm.

237

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

v
IIc FON 9S
alnGhaGan Grn) ane ence
res'z 9°6 x4
Weer Oral = apie
EIEG <9 01 L8¢
q: I q I
aa

‘suowid0ds uIapOU JO sorsas BJO 9SOY} YIM Ppoieduiod snja1d uovsd7] dipyIeMG pue uUTa|UOJspuR]y YI JO 499)

1:06'1
re ato

q:1

9's “9

v b b v b u
¢‘9 €‘TI 9°¢ 8‘Ol I‘ rE Ps SEI z‘9 L‘9
=¢'S =~ Ol | -3'7 =—3'2 | -G6'E —'s [=CL =6'8 ='S “8S adury
6's ae oe 86 €‘p oo | SL “EO! LS v9 urs
= i = << = = Lele OC} 80. E'9 6'L PIET MZ
OLOE MZ
OL L‘€l 9°99 == I‘g ‘9 _ — —_ _ a 10L9
v9 (ana 9°¢ ara | TS 1‘8 = aa = = €fl Og
— — = a = — 16 6TI ¢‘9 fight ra ey: |
q I q I q I q I q {
“d *d 'd 2)

6b alavy

Said UODIAT ULOPOW

* dipyuems

* ulojuojspuryq

JIMO] IY} JO suOoIsUsUIG]

238 ANNALS OF THE SOUTH AFRICAN MUSEUM

The only tooth in the Lycaon dentition which is common to the Elandsfon-
tein and Swartklip assemblages is the I,. The two fossil specimens (EC 12,
ZW 3070) are similar in size and larger than the I, in the comparative series.
The I? and I? from Swartklip (ZW 137, ZW 177) are also a little larger than
the modern specimens, in spite of the statement to the contrary made about
them earlier (Hendey & Hendey 1968: 64). ZW 137 measures 7,4 by 7,2 mm
and ZW 177 measures 6,7 by 8,4 mm.

Discussion

The Elandsfontein Lycaon pictus material is apparently a homogeneous unit
and represents a variety of this species which is appreciably larger than the
moderns in most observable respects. It almost certainly belongs with the earlier
(Cornelian) element of the Elandsfontein fauna.

The Swartklip L. pictus appears to be comparable to the moderns in size,
although the incisors at least are more like those of the Elandsfontein form.

The available comparative series is obviously far too small to allow for an
accurate assessment of size similarities and differences to be made, but it is not
unexpected to find that the Swartklip Lycaon differs from that from Elands-
fontein and is closer to but still different from the modern form. In keeping
with the earlier decision to avoid the use of subspecific names, the local forms
are categorized simply as ‘Cornelian’ and ‘Florisian Lycaon pictus’.

In this instance, however, the former variety already has a subspecies
name applied to it, and since its use may still be favoured by some, it is here
redefined on the basis of the additional specimens which are now available.

Lycaon pictus magnus Ewer & Singer, 1956

A variety of Lycaon pictus which dates from the Cornelian age of the South
African Pleistocene and which is presently recorded only from Elandsfontein
in the Cape Province. It differs from the extant variety in its larger size, in the
absence or near absence of anterior accessory cusps on the second to fourth
lower premolars, and in having a relatively smaller metaconid on the lower
carnassial.

Family Canidae
Subfamily Otocyoninae

Otocyon megalotis Desmarest, 1822

Although the bat-eared fox still occurs in the adjacent Cape west coast
and Karroo regions, and was much more common in these areas in the past,
there are no comparable records of this species in the south-western Cape.
However, the presence of a family group in the vicinity of Paarl near Cape
Town was recently reported by a local newspaper (The Argus, 29 December
1972) and the species is included here on the basis of this record.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 239

There is no obvious reason why Oftocyon should not have been common
locally earlier in the historic period and before, although since it is essentially
confined to the drier parts of sub-Saharan Africa it may well have found the
environment of the south-western Cape unsuitable, especially at those times
when it was cooler and wetter (e.g. the late Pleistocene). There are at least two
other carnivores, namely, Cynictis penicillata and Proteles cristatus, which occur
commonly in the adjacent more arid regions, that are known only from isolated
recent records in the south-western Cape. ‘Together with Otocyon they may have
run counter to the general trend by actually extending their ranges into the
south-western Cape comparatively recently. Their presence locally may thus
be, or have been, dependent on the more arid conditions prevailing today and
at other times earlier in the Quaternary.

Family Mustelidae
Subfamily Mustelinae

Ictonyx striatus Perry, 1810

(Fig. 57)
Present status

The striped polecat is one of the most commonly occurring fissiped carni-
vores in the south-western Cape today.

Material
Elandsfontein

9200—Braincase with associated right maxillary fragment which includes part
of P*.

Description

The fossil skull belongs to a small mustelid and is that of an aged individual,
probably a male because of the presence of a well-developed sagittal crest. It is
undoubtedly an IJctonyx, although it does differ in certain respects from the
available skulls of modern J. striatus (n = 6).

The frontal region of 9200 is damaged and incomplete, and the right
maxillary fragment is detached from the rest of the skull. The maxillary frag-
ment is indistinguishable from corresponding parts of the modern specimens
in all observable respects. The incomplete P? lacks most of its anterior half and
the wear on the posterior keel indicates that the individual concerned was
advanced in age. The posterior part of the C alveolus is preserved, as are the
alveoli of P?, P* and M1!. The teeth of the fossil were similar in arrangement, and
apparently also in size to those of the modern species.

The remaining part of the frontal region of the fossil is, in general, similar
to that of the comparative specimens, although the frontal sinuses are less
inflated. Only a part of the left post-orbital process is preserved, but sufficient

240 ANNALS OF THE SOUTH AFRICAN MUSEUM

remains to indicate that it was more prominent than those of the modern speci-
mens. Similarly the temporal ridges are well developed and clearly discernible.

The post-orbital constriction is appreciably narrower than that of the moderns
(Table 50).

TABLE 50

Dimensions of the skull of the Elandsfontein Ictonyx compared with those of a series of modern
I. striatus.

mu
Zz)
fo}
= :
—
: 3 : 2
te = n
ae : 3 a3
= fan) a 3) wa
gOS re) ie } Oo oO
cH 1S) = na : |
~~ 59} wn
2 wn = om fe) & a
o 5 § > 5 Ss +s
a2 6 x Sg os
= S ° g si (es
cmt cm ~ n Law] ad
ee g : 5 as
=e) Ay = = iS) Ha
Elandsfontein 9200 32,4 14,1 12,3
Mean 3539 1559 11,9
Modern | | | |]
Ictonyx striatus Range 32,8- 15,0- 10,5-
38,5 16,8 13,5
n 6 6 6 6 6

The braincase is correspondingly narrowed and its lesser development is
most evident when the specimen is viewed from the posterior, since the occipital
almost completely obscures those parts of the braincase anterior to it. By con-
trast, the squamosals and parietals of the modern species are visible as an arch
of bone over the occipital. The narrowing of the braincase is, however, con-
fined to the more dorsal parts and the mastoid width is within the range of
variation observed in the comparative series. The braincase is shorter even than
that of the smallest female in the comparative series and is appreciably shorter
than that of the largest male. The bone surface of the braincase is very rugose
and while older individuals in the comparative series also exhibit such rugosity,
in none was it developed to the same degree.

The width across the occipital condyles is less than in the modern specimens
and the foramen magnum is correspondingly smaller. In all the modern speci-
mens the occipital above the foramen magnum rises more or less vertically to
the nuchal crest, but in the fossil there is a very prominent shelf of bone project-
ing over the dorsal margin of the foramen. The older of the modern specimens
exhibit a similar feature, but in none is it as prominently developed.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 241

The basicranium of the fossil is essentially similar to those of the moderns,
except that the tympanic bullae are a little more inflated and there is a more
pronounced ridging of bone along the midline of the basi-occipital.

Discussion

At least some of the characteristics of the fossil braincase are apparently
due to the advanced age of the individual. Anderson (1970: 15, 16) mentions
that with advancing age in Martes, the post-orbital constriction is narrowed, the
gross size of the braincase decreases, the post-orbital processes are enlarged and
a prominent sagittal crest is developed in males, while there is also ‘an increase
in the mass of the bone of the skull’. Presumably the same applies in the case
of other mustelid genera.

Judging from the wear on the P? of 9200, this individual was older than
any in the comparative series and since the ageing characteristics mentioned by
Anderson are also the principal characters which distinguish the fossil from the
moderns, the differences may be accounted for by this fact. An increase in the
size of the brain cavity of 9200 would result in an inflation of the parietal/
squamosal walls, widening of the post-orbital constriction, straightening of the
occiput dorsal to the foramen magnum, increasing size of the foramen magnum
itself and perhaps also an overall lengthening of the braincase. A reduction in
the mass of the bone of the skull would reduce or eliminate the rugosity of the
bone surface. ‘These changes, coupled with a reduction of the post-orbital
processes and sagittal crest would result in 9200 bearing a far closer resemblance
to the available modern J. striatus specimens.

It is, however, likely that the fossil braincase would still be shorter than
those of the males of the comparative series and that the bullae would still be
more inflated. In addition, there might well be other differences between the
modern and fossil specimens which cannot be determined on the basis of the
available material. The differences do not, however, preclude the possibility

rs
110 | 12 1 Z 1

mM

NAINA
4 AS

IAIN

1t5

i
0 1 |

mm
at

Fig. 57. Dorsal and ventral views of the Ictonyx striatus braincase (9200) from Elandsfontein.

242 ANNALS OF THE SOUTH AFRICAN MUSEUM

of a close relationship between the fossil and J. striatus. Judging from its preserva-
tion, 9200 belongs in the Cornelian element of the Elandsfontein fauna and it is
here regarded as representing a relatively primitive variety of the modern species.

I. striatus is the most widely distributed small mustelid in Africa today, but
it has not previously been recorded as a fossil and nothing definite has been
known of its origins and ancestry. The Elandsfontein record indicates that the
species has undergone comparatively little change during the latter part of the
Quaternary. Ictonyx has probably been a well-established member of the small
carnivore fauna of Africa for much longer than that and it perhaps had its
origins during the Pliocene when the Eurasian element in the African fauna
was still well represented. It is remarkable that it should have remained so
successful in spite of the radiation of the small viverrids in Africa.

Family Mustelidae
Subfamily Mellivorinae

Mellivora cf. capensis Schreber, 1776
Matertal

Baard’s Quarry, Langebaanweg
L 179/12—Right mandibular fragment with parts of P, and P,.

Description

This rather poorly preserved specimen has only the P, reasonably intact.
The principal and posterior accessory cusps are damaged, but sufficient remains
to show that in size (Table 51) and morphology it is indistinguishable from the
corresponding tooth in the available modern Mellivora capensis series (n = 5).
The less complete P, is apparently also similar to that of M. capensis. The roots
of P, and the posterior part of the C alveolus are preserved and, at least in
respect of size and relative positions, these teeth resembled their counterparts in
the modern species. The M, is lost and the alveolar region of this part of the
mandible is damaged, but it appears that this tooth was relatively longer than
that of M. capensis.

The mandibular corpus resembles that of the modern species. There are
two mental foramina below P3, but the fossil differs from the available compara-
tive specimens in having a shorter symphysis. This terminates below Pg, rather
than below the anterior part of P,.

Discussion

With the exception of the possibly longer M, and slightly shorter sym-
physis, there are no grounds for believing that this specimen should be referred
to a species other than M. capensis. The reservation in the identification is made
simply because the specimen is so incomplete. Its significance lies chiefly in the
fact that it is clearly not conspecific with the ‘E’ Quarry Mellivora, and it is one
of the species which indicates that the Baard’s Quarry fauna includes a post-
Langebaanian element.

243

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

5 ¢ G G u
00S ele bg S‘*99 ow 1°i4
-9‘GP —V‘9 6°61 -2°6S —0'9 —L‘6 aBuryYy sisuaqvo vLOANIaTAY UIIPOYW
a oc L‘v1 g°zg bg Z‘O1 Uva
= —_ ofgI °9 6‘19 o‘9 L‘6 w1/6Z1 "J | Aaien() s.preeg
—_. __ |_— >>??? He] samuveqosuery
o°EP 19 oP g‘bS GG L‘6 C9f9 J Aaren() (qf,
q I 1 I
1/001 x qt 'W [/oor xq’ ‘d
“stsuadva “Ay

urspoul pue sisuadvs “Jo vioayjapy Arren() S,paeeg 2Y} Jo asoy} YM poreduios ‘vsoapyapy AssenK) .q¥, SamueeqoZur'] sy) Jo yI991 Vy) Jo suotsuswIG

1S aldv ye

244 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Mustelidae
Subfamily Mellivorinae
Mellivora capensis Schreber, 1776
(Fig. 58; also Ewer & Singer 1956: pl. 32)
Present status

The honey-badger or ratel may still occur in isolated areas in the south-
western Cape, but it is certainly rare.

Maiterial
(1) Elandsfontein

(a) Previously described specimen (Ewer & Singer 1956):

EC 14 —An almost complete skull lacking only the zygomata, right I! and

C, left It to I? and mandible.
(b) Additional specimens:

EC 44 —Braincase.

8640 —Parts of the skull of a single individual, including: Incomplete brain-
case; right maxilla with C and P? to M!; left I’; left maxilla with G
and P? to M!; right mandible with C and P, to M;; left mandible with
Cy 2; and Mi.

15616/7— Maxillary and mandible fragments of a single individual with teeth
as follows:

Right 1°, C and P? to M?; left GC, P?, P* and part of Mea nen Ge =.
and Pc deft Cs P,. P._and, partotm Ni.

15833 —Parts of the skulls of at least two individuals, including:

Incomplete braincase; right I°, P?, part of P*, and M?; Left maxillary
fragment with damaged P?; left maxillary fragment with P* and M';
right and left mandibles lacking incisors.

20021 —Braincase.

20887 /8— Parts of a skull including:

Incomplete braincase; left maxillary fragment with P® and P*.

20916 —Maxillary fragments with right P?, P? and part of P*, and left P®
and: Re

20981 —Parts ofa skull including:

Braincase; left maxillary fragment with P? to M?.

(2) Swartklip

Previously described specimens:
ZW 1 —An almost complete skull lacking only zygomata, left I? and I°, right
CG and mandible (Singer & Fuller 1962).
ZW 142—Left maxillary fragment with P? (Hendey & Hendey 1968).

(3) Lygerfontein
Q 88—An almost complete skull lacking only zygomata, left and right I’ and
I?, and mandible.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 245

(4) Sea Harvest, Saldanha
S 787—Braincase.

Description

All the fossils are morphologically indistinguishable from modern Mellivora
capensis, but the specimens were categorized according to relative size.

The Sea Harvest braincase (S 787) is that of an immature individual with
all the skull sutures unfused. It compares in size to the braincase of modern
M. capensis males (Table 52), and it is tentatively regarded as belonging to a
young male of a population which was little or no different to that which is still
extant.

The Tygerfontein specimen (Q 88) is virtually indistinguishable from the
skulls of modern M. capensis males (Tables 52, 53), and it is also regarded as
belonging to the Holocene variety of this species.

Both the Swartklip specimens have already been described (Singer &
Fuller 1962; Hendey & Hendey 1968), and neither can be distinguished from
modern M. capensis. However, Singer & Fuller (1962: 208) concluded that this
Mellivora ‘is phylogenetically intermediate between the (Elandsfontein) form
and the extant honey-badger’ because the skull (ZW 1) is rather small.
Judging from the nature of the sagittal crest, this skull probably belonged to a
young male and if this is indeed the case then it is smaller than the skulls of
males in the available comparative series.

Singer & Fuller’s statement does, however, require qualification, since
the Elandsfontein assemblage includes two varieties of M. capensis. The speci-
mens numbered 15833 are from the Florisian Bone Circle occurrence and
although the braincase is as large as that of the modern males, it is identified as
belonging to a female because of the presence of temporal ridges rather than a
sagittal crest. It is concluded that the Bone Circle Mellivora is a Florisian variety
of M. capensis which was larger than its modern counterpart.

The other Elandsfontein specimens, which apparently represent both males
and females, probably belong with the Cornelian element of the fauna and they
are smaller than the available modern M. capensis specimens. This was men-
tioned by Ewer & Singer (1956) in their description of the specimen EC 14,
which is also distinguished from the modern variety by its shorter palate.
Unfortunately, the palate is not complete in any of the additional specimens.

Discussion

All the material is referred to Mellivora capensis and four temporal variants
are recognized.

The earliest (Cornelian) form was smaller than modern M. capensis and is
represented by most of the specimens from Elandsfontein. There are apparently
two varieties represented in the local Florisian assemblages. The first is that
from Swartklip, which was a little larger than the Cornelian variety, but still
smaller than the moderns. The second is that from the Elandsfontein Bone

ANNALS OF THE SOUTH AFRICAN MUSEUM

246

‘O}CWIXOIddY

83S -I93A J, ee |

SISUAADD DAOAITJaAW UIOPO||

w4oJ AJIVO—uIOJUOJspuLlA

L‘v€ O'SE pve O9E 197 L‘Ov 6CE = LZ 6‘LZ 0°67 0°67 787 = ae
OEP 6'IP 6 Iv L‘Sv 6‘9r TSP 0‘6r 797 OEY OLE — pLe 1°6€ OIr (aa
7'8L STL €S8 b°96 €‘L8 I€L €‘76 crs = 66S — 6°99 PvL S‘TL‘°9 = Shh
5‘09 ¢‘€9 ¢‘79 (GG 1‘€9 €‘99 0‘OL oe L‘t9 ¢‘Ss 89S a‘LS 96S 709 1‘09
0*L9 ¢‘99 0°69 SILL S‘OL 0‘OL CEL SOL 0°L9'°92 | 06S ¢‘09 0'8sS ¢‘P9 0°79 0*r9
be } } ‘ue 2 “anf Pe rowA IMZ

TEIQE 9ZEE STEE Or66l wLSSe | L&LS 88 O ELSI diy dé dé bd b% Pe Pi

WvVS WVS' WYS WVS WVS_ | }S0AreH} | UlojUOJ | O 9UOG | -11eMG | [8607 L8807 bhOH tI OA O98 1Z00Z

UOIJOIIJSUOD [%}IGIO-JSOg

* yysroy [eIWdI99O
* YIPIM proyseyAl
* YIPIM osvouleig

,UISUS] OSvOUIvIG

‘susut1oads UIOPOUI JO Salios B JO ssOYy} YIM posredwioo ‘soutAOIg oder) 94} WIOIy Sisuadvs vs0A2r7;aP [ISSO} JO [[NYS IY) JO suoisuowIdg

oG aTaVvy,

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 247

WM A AL GT AUT A

Fig. 58. Dorsal views of the Mellivora capensis skulls from Elandsfontein (EC 14) and Tyger-
fontein (Q 88).

ANNALS OF THE SOUTH AFRICAN MUSEUM

248

“yIsuZ] [ensul]—|] ‘yysus] yeoong —]{q x

g G G G ¢ u
O61 L‘o1 OG oS1 QhI G‘9 £6 cy 9‘9 VL v6
Vo! =a'L -1'P —1o1 —6‘11 6G -z‘9 VE =—8°G | -G*G -—1‘L IBULY sisuagvs vs0n1yjayy UIIPOW
Oo £‘9 ov OU Bel 0‘9 9‘8 6°E oFG V9 a‘Q ued
OSI 1°Q QV OUGiIy manor 9‘9 1°6 ov 9‘9 CSL 9 OO % 988 0 UIO]UOJIOBA J,
ane =e —_ a = — — VV £9 — — oVI MZ
oI Tha Ly Sor gQ*aI £9 L‘g ge EG v9 Z‘Q I MZ dipyarems
O%SI are) gv 9} og'o1 Sa — — — — — — SEQSI QDI VUOg UTD} UOJspuLly
— — — SG) (OI £9 £°Q o°v 6G — — g160a
Gor G°G QV OD (ii 9‘S Sle g‘E 0G v9 QL g19S1 2
ee @) le!
— os ~ O@ OOr% | GH ECM, BD — — — — 8980z ae
> o
66 ofS QE GO HOLD] OG 1° — — — — 19602 3 5
Q,
G6 rg 6 G6 gor oS of Ge e | of by PLOW ;
Tier 0‘9 ov OD S71 6G EU, VS 1G 6S o‘L ovgg
q «II *14 q I q I cl I le ee al
IN od ed ed ©)

‘suoutioods usopoul Jo dsoy} YM poreduios ‘adeg usd}sam-YINOS 9Y} WIT sisuadvs vsoayjapy Isso} Jo y}90} 19ddn 9y) Jo suotsusuIIq

6S alav J,

249

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

¢ G G ¢ 1 u
ME Pg gL nein 6'G bia |= Gk 6‘G c‘9 1‘O!

-V‘9 —6°E1 | -o‘9 -L‘6 | -9‘P -ofL | -S°E -—o'G | -o'g -9L aBury sisuaqvs vLsonyjayy ULIPO|

o‘L L‘vi bg Z‘OI a‘G Cae, Ox BG a 6‘9 Uva
Tig BGI 89 v6 SG 99 | SP gS | og Z‘OI GEQS1 IID 90g UTs}UOJspuLyy

— —— GEG G6 gb 69 — — 3°9 9°98 gig9S1
| sd -- | w110} Aj1ed UIOJUOJspurTy

Z‘9 oS VG v6 VD Alo may AS 8 oS S‘9 fobs) ogg

q I q I 4 I q I q I
"W "d “d ‘d 2)

‘susuttoods ULapoOUr JO satsas v JO asoy} YUM poreduiod ‘adey us9}saM-YINos ay} Woy sesuadns vsoAIYay [ISSOJ JO 4399} J9MOT IY} JO suOIsUDUITG

SG alavye

250 ANNALS OF THE SOUTH AFRICAN MUSEUM

Circle occurrence and this one was larger than the modern variety. The last
of the temporal variants is the modern form itself and it is represented by the
fossils from Sea Harvest and Tygerfontein, as well as being recorded in the
historic period.

It was concluded earlier on the basis of the Canis mesomelas from the Bone
Circle and Swartklip that these two occurrences were broadly contemporaneous
and, while this probably is the case, the M. capensis remains suggest that the
Swartklip fauna is closer to the Cornelian element of the Elandsfontein fauna
than is that from the Bone Circle.

The south-western Cape Mellivora is, therefore, categorized as follows:

Cornelian M. capensis— Elandsfontein
‘Early’ Florisian M. capensis—Swartklip
‘Late’ Florisian MM. capensis— Elandsfontein Bone Circle
Holocene M. capensis—Sea Harvest, Tygerfontein and modern fauna

The difficulties in categorizing specimens solely on the basis of size were
discussed earlier and it is certain that if larger samples of the four groups of
M. capensis were available, each would overlap in size characters with preceding
and/or succeeding groups. Consequently, it might well prove impossible to
categorize certain specimens of unknown age if they differ in size from the
‘typical’ examples of the categories which are recognized here. This taxonomi-
cally informal and flexible categorization has, therefore, a very restricted
application and the arrangement is necessarily provisional.

Perhaps the only point of real significance to emerge from the present
study of M. capensis was the suggestion that the Swartklip fauna predates that
from the Elandsfontein Bone Circle.

Family Mustelidae
Subfamily Lutrinae
Aonyx capensis Schinz, 1821
(Fig. 59; also Ewer 1962: pl. 1)
Present status

The Cape clawless otter is now very rare in the south-western Cape, but
it was formerly common.

Material
Swartklip

(a) Previously described specimen (Ewer 1962):
ZW 6a—Left M,.
(b) Additional specimens:
ZW 2672A—A crushed and incomplete skull comprising the right half of the
braincase and the most posterior part of the right maxilla with M?.
ZW 2672B—A crushed and incomplete skull made up largely of the left half of
the braincase.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 251

ZW 2672C—Part of the right tympanic region of a skull, possibly belonging
with ZW 2672B.

The above three specimens were recovered from a single block of matrix and
they represent at least two individuals.

ZW 2930—Left maxillary fragment with P* and M?.

ZW 3486—Right M,.

Description

The specimen ZW 6a was described in detail by Ewer (1962), and she
concluded that it belongs to a species other than Aonyx capensis.

The additional specimens also differ from the modern species in certain
respects, although in this instance there is no doubt that the material should
be referred to A. capensis.

Since the skull fragments are crushed and incomplete, they could only be
measured in a few places (Table 55), but it is in any case evident that they are
larger than corresponding parts of the few available skulls of the modern species
(n = 4). The nuchal crest of ZW 2672A, and what little remains of the anterior
part of the sagittal crest, are more strongly developed than in the modern
A. capensis specimens, while the post-orbital process is more prominent. The
tympanic bulla is less inflated than those of the moderns, but it has an appre-
ciably greater transverse diameter. The difference between the antero-posterior
diameters is less marked. Similarly, the glenoid fossa has a greater transverse
diameter, but otherwise it and adjacent parts are similar to modern A. capensis.

Since the Swartklip canids are also larger than their modern counterparts,
the size difference between the Swartklip otter and modern A. capensis is not
unexpected. Unlike the canids, however, the otter teeth are less consistently
larger than those of the modern form (Tables 55, 56). The upper teeth are either
within the range of variation observed in the moderns, or are only slightly
larger. They are morphologically indistinguishable from corresponding teeth
of the comparative series.

The situation in respect of the known lower teeth, two carnassials, is more
complex. One of the specimens, ZW 3486, is a little larger than any in the
comparative series and its size is consistent with the upper teeth of the fossil
form. It is essentially similar in size to the A. capensis M, from Florisbad (Ewer
1962), and it is accommodated quite well by the M, alveolus of an Aonyx
mandible from a previously unrecorded locality in the southern Cape (Lake
Pleasant). The Lake Pleasant occurrence is very similar to those at Swartklip
and is also regarded as Florisian in age. The otter mandible (Q 1745) is appre-
ciably larger than those of the moderns and, as far as can be judged, its size
is consistent with that of the Swartklip skulls.

The previously described lower carnassial (ZW 6a) is very narrow com-
pared with the other fossil specimens and also the M, of the moderns. It was
this characteristic which prompted Ewer (1962) to suggest that it should not
be referred to A. capensis. This specimen does not occlude well with the upper

ANNALS OF THE SOUTH AFRICAN MUSEUM

252

6 6 v 14 v 17 v u
Q‘L& Gz 1‘Qz 1:Z0‘1 1:08‘0 6g gb gb gb sisuadva xduop
—-o0‘L€ -L‘Va —S‘Gz —1:96‘o -1:SL‘o —E‘o1 -9's1 | -O'€1 —-S*ar osury ulopoyy
PLE 1‘Gz 6°92 1:00‘ 1:gL‘o aL PEI GSI 6°E1 uray
ce = — 1:S0‘1 1:39'O L‘g1 L‘é1 Lor vb 066% MZ
On a4 36% — — — — -— — — azloz MZ dipx31eMsg
ooP °9 £°6% 0's& — 1:6g‘0 L°G1 oF ~ — Volgz M
ey[nq o1uedurA} | ejjnq d1uvdurA} | vssoy prousys I q:] q I q I
po Ue oe Jo WN rd. IN WN rd
*ysod-"juy ‘WIvIp ‘suey, | “WIvIp ‘suviy,

‘susuitdods UJOpOUL JO solios & JO OSsOYy} YUM poredusod sisuadvs xduop dipyxIIeMG IY} Jo {[NyYs pue yj}00) 19ddn sy} Jo suotsuswiId

GS alavy,

253

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

‘SQG1 IMT »

-6°9g —o‘gI —b‘6g —1:SS°1 —O'1r 8=—ESL1 -3°9 -L‘g asur yy sisuadva xtuop ULIPOYy
S‘6 161 G16 A | Ltt 6°91 oc 6‘g uvd
— — a 1:9S‘1 ee | o°O% — — x06VI D peqsaoyq
g‘O1 L‘1% 6‘g6 1:QQ‘I *9 Ss1's ofle 3 | 6'9 g‘ol GPLI © yuesea|g dye]
— a= — 1:9L‘1 QI 90% — — ggh& MZ
——__—————_— |“) Mm ii, Y*Y'—_-_-Acm i cr dipyaemg
mae a= ee 1:06‘1 6'6 ‘gl = — ®9 MZ
"WW Moog A Mopq S[qtpueu q: | q I q I
sndioo sndioo jo yysu9] WN W @)
ORC eo ae Jo WYBIOH teas) <

‘BOY YING wo sisuadvs xCuop |Isso} puke UWPOUI JO YI99} JOMOT PuLK soyqrpuvUl dy) JO suOISUIUICT

gS alavy

254 ANNALS OF THE SOUTH AFRICAN MUSEUM

teeth from Swartklip and it is too small to fit the M, alveolus of the Lake Plea-
sant mandible. On the other hand, Ewer (1962: 275) stated that, ‘The cusp
pattern of the fossil tooth is the same as that of the extant species and its length
falls within the range of a sample of 11 A. capensis’ and concluded that it clearly
belonged to the genus Aonyx.

OY
114 LS 116 3 114 Lis 116

Fig. 59. Occlusal views of the Aonyx capensis lower carnassials (ZW 3486, ZW 6a) and maxilla
(ZW 2930) from Swartklip.

Coetzee (1967: 14) gives one of the diagnostic characters of modern
A. capensis the fact that the M, ‘measures over 16X10 mm’. The breadth of
ZW 6a falls just short of Coetzee’s minimum measurement, while its length is
not inconsistent with that of the moderns. It is the length/breadth ratio of this
tooth which really sets it apart from other specimens, both modern and fossil.
In terms of its actual length and breadth, ZW 6a actually differs more markedly
from the fossil specimens than the moderns.

Discussion

The Florisbad, Lake Pleasant and Swartklip material, exclusive of ZW 6a,
evidently represents a Florisian form of Aonyx capensis which was larger than the
extant variety. Ewer (1962: 276) suggested that ZW 6a belonged to ‘a conserva-
tive, little modified descendant of the same stock as gave rise to the modern
Aonyx capensis’.

These interpretations of the fossil evidence mean that not only were there
two varieties of Acnyx present in the South African Florisian, but that both are
represented in the Swartklip assemblage. While this possibility cannot be
discounted, it is perhaps more likely that only a single variety is represented
and that ZW 6a belonged to an aberrant individual. This specimen is now even
more problematical than when it was first described and the situation will

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 255

probably not be resolved until more Florisian Aonyx specimens are described.

Dreyer & Lyle (1931) referred the Florisbad otter specimens to a new
species, A. robustus, but Ewer (1962: 276) concluded that they represent what
is simply ‘a rather large specimen of A. capensis’. She also stated that if ‘the fossil
teeth belong to an average-sized individual from a population whose mean
tooth dimensions are significantly greater than those of the extant otter’ then
‘sub-specific separation would be warranted’. ‘The Lake Pleasant and additional
Swartklip specimens do indeed indicate that the Florisbad otter was not just an
unusually large individual. Although the material is here categorized as
‘Florisian Aonyx capensis’, a name is available to those who prefer to recognize
it as a distinct subspecies, that is Aonyx capensis robustus Dreyer & Lyle, 1931.

Family Viverridae
Subfamily Viverrinae
Viverra civetta Schreber, 1776
Present status

The African civet is largely confined to tropical Africa and has not been
recorded in the Cape Province in historic times.

Material
Elandsfontein
15601 /2—Right P, and part of right P*.

Description

These two teeth evidently belong to a single individual. The P* is damaged
and the protocone and parastyle are lost. That part which remains is indis-
tinguishable from the P* of the only available modern Viverra civetta skull. The
P, is intact and is similar in size (11,3%5,7 mm) and morphology to the
corresponding tooth of the modern specimen, although the posterior cingular
region is narrower and less crenellated.

Discussion

There can be no doubt that these specimens belong to a civet which, in
respect of at least some teeth, was comparable in size to modern V. civetta.
The less complex structure of the posterior cingulum of P, is what would be
expected of a civet which was more primitive, but still closely related to V.
cwetta. The only other Pleistocene civet known from South Africa is that from
the Kromdraai australopithecine site (Hendey 1973a), but since this is repre-
sented only by an incomplete humerus, it cannot be compared with the Elands-
fontein form. It was, however, a large species.

On the basis of the material presently available, the Elandsfontein civet
can only be referred to Viverra (Civettictis) civetia and, judging from the preserva-
tion of the teeth, it probably belongs with the Cornelian element of the Elands-
fontein fauna. The indications are that there was a steady decrease in overall

256 ANNALS OF THE SOUTH AFRICAN MUSEUM

size in South African fossil civets from the Langebaanian V. leakeyi, through the
Makapanian (Kromdraai) Viverra to the Cornelian JV. civetta, and that they were
representatives of a single lineage which culminated in the modern J. civetta
(see Fig. 10).

The presence of this species in the Pleistocene of the south-western Cape is
notable, because in recent times it has been recorded no closer than the northern
coastal plain of Zululand (Shortridge 1934), which is about 1 500 km to the
north-east. The failure of a number of herbivores to extend their ranges into,
or to maintain their presence at the southern continental extremity was earlier
attributed to environmental factors, but this control is far less likely to apply
in the case of an essentially carnivorous omnivore such as V. civetta. The distri-
bution and relative numbers of civets will be discussed in more detail later, but
the indications are that they have become a progressively less significant element
of the southern African fauna during the Pleistocene and Holocene.

Family Viverridae
Subfamily Viverrinae

Genetta genetta Linnaeus, 1758
&
Genetta tigrina Schreber, 1776
Present status

The common and large-spotted genets still occur in the south-western
Cape, but neither species is common.

Comment

Genets have a poor fossil record and only a single specimen belonging to
this group has thus far been recorded from a Quaternary fossil occurrence in
the south-western Cape. The taxonomy of the modern representatives of this
group is complicated and about eleven species are currently recognized, with
distinctions being based principally on external characters (see Coetzee 1967).
Consequently the identification of fossil forms may be extremely difficult. In
the present instance the fossil specimen was compared only with small series
of the two species which still occur in the area and it is tentatively identified with
one of them, i.e. G. tigrina.

Genetta cf. tigrina
(Fig. 60)
Material
Sea Harvest, Saldanha

S 1106—Right mandible lacking incisors and P,.

Description

The mandibular corpus and ascending ramus of S 1106 are largely intact,
although the angular process and immediately adjacent parts, together with a

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 257

portion of the symphyseal region are missing. The mandible is appreciably
larger than those of the available series of modern Genetta genetta (n = 4) and
G. tigrina (n = 6) (Table 57), but it is otherwise similar to the comparative
material. The modern specimens have two mental foramina in each half of the
mandible and in G. tigrina the more posterior of these is situated below the
anterior root of P;, whereas in G. genetta it is more anteriorly situated, being
either below the posterior root of P, or between the posterior root of P,
and the anterior root of P,. The fossil specimen has the posterior mental foramen
situated as in G. tzgrina.

Of those teeth remaining, only one, the M,, compares in size with the
corresponding tooth in one of the modern species (G. genetta). Otherwise the
teeth are all larger than those of the comparative series and are thus in keeping
with the overall larger size of the mandible.

HU
5 1

ee

gg

TUTTLE LLU LL
3 114 1 6 1

Fig. 60. Buccal views of the Genetta cf. tigrina mandible
(S 1106) from Sea Harvest and a modern G. tigrina specimen
(SAM 36105).

The lower dentitions of G. genetta and G. tigrina are not easily distinguished,
but the most obvious distinguishing characteristics observed in the available
comparative series are that G. genetta has a larger P, and higher crowned pre-
molars than G. tigrina, while the M, was also usually larger. The P, of S 1106
is lost, but judging from the size of its alveolus it was proportionately as large
as that of G. tagrina and smaller than that of G. genetta. Similarly the remaining
premolars are less high crowned than those of G. genetta and more like those of

=
+)
isa
iva)
5
=
Z
<
S
%
fy
<
a
isa
re)
e)
i?)
ca
an
fH
fx
)
ive)
4
<
Z
Za
<

258

9 9 9 9 9 9 9 9 9 9 9 ¢ u
8°79 Sit 701 8°8€ OL'7: 1 ie Ge |W G2) rie I6 6D) SE ED) ee SS] OS Ly
co 29 =€'€ =GaL —€ ‘pe Sala call a7 6 ALG || Se =k, |) =H a oe eae BOP | Clea Gia | Oeil sO Gan || SONG Oe suey DUIAS1] D1JQUIH
[fa md uIOpO|
6°8S I'v 8°8 L‘9€ aaa | Sic is (Sh Je SC LO | YE O© | Oe SS 6. S17 urs
v v v v Y v v v v + v v u
p'e9 I'v L'8 7:0F plz its Oy |) Oty TOE | tay iay He US| Se GO) Ce Wi Oe ab
-6°9S One =6il -0°9€ -€6 1:1 G16 ONS. || SES a7 Sedov a! NG SQ) | GG ACD | SG SS. || Sarg SOs osury D1jaUuas DjJaUayH
Se aa a oe a | a al ee ulopoyy
6'6S g‘€ €‘8 SLE 10°71 OE BC ge OL OT sz 99 SG 179 | Or 16S Cie Sis urs
6°L9 GAS 601 Lt co) rae | TE BC rr 78 PIII Oe SY, i Oe OG CS HS aes 7S 9011S
(e[Apuos
01D) NW Mojeq | A Mojeq I I q I q I I I q I q I q I
g[qipueur | ajqipueur i \or®) ATW “1A W ai*d ‘d d *d 5)

Fo igen JO YIpeoig | JO 14SI0H =

‘DUIS “) PUL DyIUaT “4 UIBPOUI JO Sotias & JO 9SOY} YIM poseduio0d “\saxrIePT VIG WOLF YJJaUay [ISSOJ JO YII9} IOMOT PUL I[qQIpUeU dy) JO sUOIsUsUTIG]

LG atavy,

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 259

G. tigrina. The relative lengths of the fossil P,; and P, are also most similar to
those of G. tigrina. In actual size the fossil M, corresponds with those of G.
genetta, but in its length relative to that of M,, it is intermediate between those
of the two modern series.

Discussion

While the Sea Harvest genet was evidently a larger variety than either
modern G. genetta and G. tigrina, its dentition most closely resembles that of the
latter species and it is identified accordingly. The reservation in the identifica-
tion is made because of the complexities in the taxonomy of the modern species,
the fact that the present comparative study was inadequate and because the
fossil variety is so poorly represented.

Family Viverridae
Subfamily Herpestinae
Herpestes ichneumon Linnaeus, 1758
(Fig. 61; also Hendey & Hendey 1968: pl. 7D)
Present status

The Egyptian mongoose is now extinct in the south-western Cape, but it
still occurred in the region during the ninteenth century.

Material

(1) Elandsfontein

15833—Right P*.

6867 —Right mandibular fragment with P, and P3.
6880 —Left mandibular fragment with P, to P,.
6885 —Left mandibular fragment with P,.

8101 —Left M,, possibly belonging to 6880.

(2) Swartklip
(a) Previously described specimen (Hendey & Hendey 1968):
ZW 111—A crushed skull lacking the braincase and mandible and with the
following teeth missing:
Eeft 7, PY, P*, M2 and M?*; right [, part of P4, and M?’.
(b) Additional specimens:
ZW 2647—Part of an occiput.
ZW 1627—Right mandibular fragments with C and M,.
ZW 1845—Right mandibular fragment with P, and P,.
ZW 2001 —Left mandibular fragment with P,.

(3) Sea Harvest, Saldanha

S 138—Left maxillary fragment with P? to M?.
S 209—Right mandibular fragment.

260 ANNALS OF THE SOUTH AFRICAN MUSEUM

Comment

The study of this material was hampered by the fact that only one skull
of the modern species was available for comparative purposes. However, since
the nature of the changes undergone by locally occurring fossil populations of
this species is apparently essentially similar to those of other species (e.g. Canis
mesomelas), the identification of this material and the observations made about
it are probably secure.

Description

The maxillary fragment from Sea Harvest (S 138) is virtually indistinguish-
able from corresponding parts of the skull of a modern Herpestes ichneumon from
Pondoland (SAM 1883) (Table 58). The same applies to the edentulous mandi-
bular fragment (S 209), except that the corpus is a little broader and more
heavily built (Table 59).

The incomplete skull from Swartklip (ZW 111) has already been described
elsewhere (Hendey & Hendey 1968: 66). It is generally similar to the modern
specimen, although the P* protocone is a little larger and the parastyle a little
smaller. The Swartklip mandibular fragments do, however, differ quite appre-
ciably from the modern and Sea Harvest specimens. The corpora are much more
robust, the C is larger than that of the modern specimen and the cheekteeth are
broader. The P, has a less distinct anterior accessory cusp and the protoconid
of M, is only a little higher than the paraconid and metaconid, so that the
trigonid is composed of three more or less equally sized cusps.

The only upper tooth recorded from Elandsfontein is an isolated P#
(15833) from the Florisian Bone Circle occurrence. It resembles the Swartklip
P* in having a smaller parastyle and larger protocone than the Sea Harvest
and modern specimens.

The remainder of the Elandsfontein specimens probably belong with the
Cornelian element of the fauna. The lower teeth are very similar to those of the
modern specimen, but the mandibular corpora are more robust. There is a well-
developed subangular lobe in 6885, which is the only specimen in which this
region of the mandible is preserved. The Swartklip, Sea Harvest and modern
specimens do not have subangular lobes. Judging from the size of the M,
alveolus in 6885, this tooth was larger than in the later specimens.

Discussion

It has already been indicated that the south-western Cape fossil Herpestes
ichneumon apparently differs from the modern variety in much the same way
as the fossil Canis mesomelas differs from its extant counterpart. The characteris-
tics may be summed up as follows:

(1) Sea Harvest— Specimens are essentially the same as the modern form.

(2) Swartklip—The fossil form is larger than the modern in most respects and there are also
some minor differences in dental morphology.

(3) Elandsfontein—The Florisian form (i.e. that from the Bone Circle occurrence) resembles
that from Swartklip.
The Cornelian form is similar in size to that from Swartklip and differs from the moderns
in certain respects (e.g. presence of a subangular lobe; larger M,).

261

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

EEQGS1 ulsyUOJspuryy
Ill MZ dipawemsg
gbI g ISdAIeFT VIG

£881 WVS

‘uswtoads usiapour & Jo 9sOY} YIM poreduros ‘sourAOIg odeD Us19}S9M-YINOS DY} WOT UOwWNaUYII sajsadiafy [ISsOy JO yI99} 19ddn ay) Jo suotsusUTIC]

gS aldvy,

ANNALS OF THE SOUTH AFRICAN MUSEUM

262

— — 0G 06 | — — | — — | — — j — — 101g
9G o.f1 — — | oF 99:9 | — — | — — | — — [899
| J | | | _] ___] ur quyspureyy
rake) GSI — — | ov PIL, OD | ite Vg | &E Gq} — — 0889
a? nds zat Se es eG 99 | 6% gS | — = L989
rele) teh 2) — — | vv 1‘ — — | — — | — — 1003 MZ
&°9 L‘61 — — | S‘Y g‘L robkel L°g) |; — — | — — Ch81 MZ dipyjreMg
—- — crc 96 | — — --~ — | — — | 0G Give LoOl MZ
69 6‘o1 = — | — oo — — | — — | — — 60% S JSOAIVFT FIG
VG VOI 1‘G 96 | L‘é 9‘8 ohne 99 | L‘z qc | 9S 6‘9 | £881 WYS
"d Mojeq "d Mojeq q I Sl I q I | I q I
sndioo snd.io9 OWN val Ua oq ®)
Jo yypeoig Jo 1410 z
‘usultoods

UJOpOU & JO 2804} YUM poaieduiod ‘soutAoIg ade U19}S9M-YINOS 9Y} WOI UOwWNaUYI sSaysaduazy [ISSO} JO YI99} IOMOT PU IIqIPUeW dy} JO sUOIsSUIUTIG

6G alav],

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 263

10
[eae 2 5]

Fig. 61. The mandible of the Herpestes ichneumon from Elandsfontein reconstructed from the
specimens 6880, 6885 and 81or.

H. ichneumon is recorded as a fossil from a number of localities in Africa
(see Cooke 1963; Hopwood & Hollyfield 1954). It is still one of the most widely
distributed of the African viverrids and is also recorded from Spain and Israel
(Dorst & Dandelot 1970).

Family Viverridae
Subfamily Herpestinae
Herpestes pulverulentus Wagner, 1839
(Fig. 62)
Present status

The Cape grey mongoose is still common in the south-western Cape.

Material
Sea Harvest, Saldanha

S 786—Incomplete skull lacking most of the braincase and with the following
teeth:
Right I? and I? and P? to M!; left C to P*.

S2 —Right mandibular fragment with C to Mj.

Description

The mandible (S 2) is without P, and in this respect it is similar to the
available Herpestes pulverulentus comparative series (n = 14). This tooth is often
absent in Herpestes (Galerella), whereas in Herpestes (Herpestes) it is apparently

264 ANNALS OF THE SOUTH AFRICAN MUSEUM

always present. The other lower teeth of the fossil are morphologically indis-
tinguishable from those of H. pulverulentus and, although they are relatively large,
they fall within the size range of variation observed in this species.

The mandibular corpus is most clearly distinguishable from those of the
comparative series in that it has a greater dorso-ventral diameter in the symphy-
seal region, but the post-symphyseal parts are also more robust than even the
largest of the mandibles in the comparative series (Table 61).

The incomplete skull (S 786) evidently belongs to the same species as the
mandible, but it is from a second individual. It differs from the skull of modern
H. pulverulentus in its larger size (Table 60), and in having a more prominent
post-orbital process.

ANCUUTIIAIA LULU
3 114 IS 116 117

Fig. 62. Buccal view of the Herpestes pulverulentus mandible (S 2) from
Sea Harvest.

Several of the teeth of this specimen are lost, but only I! and M? are not
represented at all. The M? was a double-rooted tooth and, judging from the
size of the alveoli, it may have been a little larger than any of those in the
comparative series. The other upper teeth are generally similar to those of
modern H. pulverulentus, although, like the lowers, they are relatively large.
The Cand P? are actually longer than any in the comparative series (‘Table 60).
Both left and right P! of S 786 are absent and although the postcanine diastema
is long enough to have accommodated this tooth, it seems that it was never
present in this individual. Only one specimen in the comparative series had
P! absent from both sides, while in two others it was present on one side
only.

The right I? and C of S 786 were broken off during life, with the result
that the right cheekteeth are more heavily worn than those on the left side.

265

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

G1 G1 VI v1 VI F1 G1 FI oI u
o'Gt 6°G1 6‘6& g‘bz i al oP ors og | gf oS G‘% gt snjquaynasayng
P11 La | —G‘EE -9‘1% ~G -L'E|-0G -gig | -0% -FF -9‘1 6's a3ueyy saysaqapy
|] —H—_. UIIpoyy
E°E1 V1 1‘9& BES 89 iv | 3G BL |) Mee L‘vV 1% PE uvayy
L‘S1 o‘QI v‘ov 09% One bb | ofS gil | gf a‘G Gz uv 9g g | Is2AreP vag
rhe) Rove teeter ea ajyejed rd JO | I q I 4 I q I
[eyq41o-sog | [¥iUqso-s9}UT | = jo yIsus] spue ysod IW rd ed ad D

uvIPIYy SSO1Oe YIPIM

*susutdeds UIOPOUI Jo satias & YIM poreduroo snyuapnsaayng sajsaguazy ISIAIFY LIC 2} JO Yy}I9} 19ddn pur [[nys 94} Jo suoisusuICT

og alavy],

266 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 61

Dimensions of the mandible and lower teeth of the Sea Harvest Herpestes pulverulentus compared
with those of a series of modern specimens.

as Height of Breadth of
C mandible mandible
] b post. to M, post. to M,
Sea Harvest S2 4,1 57] 453
Mean 4,0 — 3,6
Modern a
Herpestes Range 3,5- — 3,0-
pulverulentus 453 = 4,2
n 12 14
Discussion

The small viverrid from Sea Harvest belongs to a species of Herpestes
(Galerella) resembling H. pulverulentus, but one in which individuals were larger
than those making up modern local populations of this species. The absence of
P! and relatively large size of M* may prove to be more significant differentiating
characteristics, but on the basis of the material presently available it is con-
cluded that the two forms are conspecific.

This is apparently the first fossil record of the species. It is remarkable
that it should be so rare as a fossil in a region where it still occurs quite com-
monly, whereas the larger H. ichneumon is more common as a fossil, but no
longer occurs locally.

Family Viverridae
Subfamily Herpestinae
Atilax paludinosus G. Cuvier, 1829

The water mongoose still occurs in the mountains adjacent to the south-
western Cape and it has been recorded from within the region in the past
(Sclater 1900). It is also known locally from a Late Stone Age coastal midden
(Van Noten 1967).

Family Viverridae
Subfamily Herpestinae
Cynictis pencillata G. Cuvier, 1829

In none of the standard textbooks on South African mammals (e.g.
Sclater 1900; Roberts 1951; Ellerman ef al. 1953) is the yellow mongoose
recorded as occurring in the south-western Cape. Its presence locally was,
however, recently recognized and as this species is a rabies vector it has been the
subject of a detailed study (I. F. Zumpt, pers. comm.). This species is essentially

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 267

confined to the drier western and central parts of the interior, where it is often
found in association with Suricata suricatta and Xerus inaurus, two species which
have not been recorded locally in recent times.

In view of the long standing and widespread human settlements in the
south-western Cape, it is surprising that C. penicillata had not previously been
recorded from this region and the possibility that it is a relatively recent immi-
grant cannot be dismissed (vide supra).

Family Viverridae
Subfamily Herpestinae

Suricata major n. sp.

(Figs 63, 64)
Holotype

8669—Parts of a braincase, a right maxillary fragment with P* and M? and left
mandibular fragment with P, and P,.

Referred material

EC 7, 6331, 9492—Incomplete braincases.

6873 —Right maxillary fragment with P? and P*.

6893 —Left M?.

16558—Right M!.

17126—Left premaxillary and maxillary fragment with I*, C and P?.

9499 — Mandible lacking ascending rami and with right C to P,; parts of
leit-P, and P,.

6884 —Right mandibular fragment with P, to Px.

9197 —Right mandibular fragment with P, to P, and Mg.

6883 —Left mandibular fragment with P, to M,.

8405 —Left mandibular fragment with C and P, to P,.

8655 —Left mandibular fragment with P, to M,.

Locality

Elandsfontein.

Age

Cornelian.
Diagnosis

A Suricata of large size; skull relatively long; sagittal and nuchal crests
prominent; post-orbital process prominent but no post-orbital bar; transverse
slot of ectotympanic continuously open and situated in a groove with well
defined posterior and medial margins; external auditory meatus T-shaped
in lateral view; Pj present; P, without anterior accessory cusp; M, protoconid
and metaconid approximately equal in size; M, paraconid and protoconid
approximately equal in size and talonid equal in width to trigonid.

268 ANNALS OF THE SOUTH AFRICAN MUSEUM

Etymology

The specific name serves to indicate the large size of this species relative
to the extant Suricata suricatta.

Description

The fragmentary braincase EC 7 was referred to by Ewer & Singer (1956:
335) as follows:

‘In size and general conformation the specimen closely resembles Herpestes
ichneumon (Linn.), but in view of its incompleteness no detailed comparisons
are possible.’

The reticence of these authors in giving further comment on this specimen has
proved to be justified, since the more complete braincase 9492, which is almost
identical with EC 7 in all observable respects, differs significantly from that of
FH. ichneumon. In dorsal and lateral view this braincase actually resembles that
of Cynictis and differs from H. ichneumon in being more expanded and relatively
shorter. In addition, the tympanic region is quite unlike that of H. ichneumon
and of the extant viverrid genera it most resembles Mungos and Suricata.

The bulla of the fossil species is known from three specimens (the holotype,
6331, 9492), and it is one of the more important diagnostic features of this
species. In general, it is more like that of Mungos mungo than any other viverrid
species which occurs in southern Africa today. However, the bulla of M. mungo
has a more inflated entotympanic, while the ectotympanic medial to the trans-
verse slot is a little less inflated. The transverse slot of the fossil resembles that
of Suricata in that it is relatively wide and continuously open, although it is
open for a greater distance than is usually the case in S. suricatta. On the other
hand, it is like M. mungo in that it is situated in a groove, which in the fossil
has more clearly defined posterior and medial margins. This groove, or depres-
sion, has the effect in both the fossil and M. mungo of compressing the external
auditory meatus dorso-ventrally. In the former the open transverse slot results
in the meatus being T-shaped in lateral view. In M. mungo the slot is usually
closed at the lateral margin so that the meatus is simply elongated antero-
posteriorly. Crossarchus, which also has a discontinuous ectotympanic slot, has
the meatus more or less circular in lateral view, as does Suricata.

Although the braincase of the fossil species resembles that of Cynictis, it
differs from this genus, and Swricata, in that the post-orbital process is relatively
shorter and does not form a post-orbital bar by linking up with the zygomatic
arch. The fossil is similar to Mungos in this respect. The development of a post-
orbital bar is an advanced characteristic in viverrids and is an ontogenetic
feature. For example, it is completely developed in Cynictis only after the animal
has reached the age of about one year (Zumpt 1969). Consequently, the fossil
species is less advanced in respect of the development of its post-orbital process
than Cynictis and Suricata. However, since this process is relatively more prominent
in the fossil than it is in Mungos, the latter is the less advanced in this respect.

269

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

ZI 9
lui

if S
nut

I b

t

£

l ¢é

1 I

|

“UloyUOjspuL[y Wor (6P6) {[nyxs solpw vzv9UNG
24} JO SMOIA [eIjUZA pu [esIOp “[eIDIeT “fg -S1y

I

L\I — ol S|t v £
Hu

if rani Wt 0
i nn

270 ANNALS OF THE SOUTH AFRICAN MUSEUM

The fossil braincase is relatively shorter and more bulbous than that of
M. mungo taenianotus, the southern African subspecies. It is, however, similar in
shape to that of the M. mungo gothneh specimen (AMNH 51112) from Central
Africa illustrated by Allen (1925: Fig. 49), except for being slightly less inflated
anteriorly. It is also about 50% larger. The fossil braincase is not as inflated
as that of S. suricatta (Table 62).

TABLE 62

The dimensions of the braincase of Suricata major from Elandsfontein, compared with those
of some extant Viverridae.

Braincase Braincase
length breadth I:b
Mungos mungo SAM 33329 42,8 28,1 1,52:1
taentanotus SAM 3484 41,5 28,2 1,47:1
Mungos mungo gothneh AMNH 51112 (52,5)* (40,0) * 1,31 Bp
Suricata major 9492 50,0 c. 38,0 9228
Cynictis penicillata’ SAM 17079 37,0 28,6 1,29:1
SAM 4105 3735 29,2 1,28:1
Suricata suricatta’ SAM 19124 32,0 20,5 1,08:1
SAM 36009 30,3 29,2 1,04:1

* Measured on fig. 49 of Allen (1925); scale x 3/2.

The nuchal crest is developed to about the same degree as in M. mungo,
but the sagittal crest is more prominent.

The upper dentition of the fossil species is less well represented than the
lower dentition. The specimen 17126, which includes I, @ and P!, agrees in its
state of preservation and degree of wear on the teeth with the holotype and it
may belong to the same individual. It is significant only because it shows that
P!, a small, single-rooted tooth, was still present in this species. This tooth has
been lost in modern Mungos and Suricata.

The other upper cheekteeth differ from those of the two modern species
in their larger size (Table 63) and in the relative sizes of certain cusps. The
P? has an internal cusp which is proportionately similar in size to that of
Suricata, but smaller than that of M. mungo. The P? differs from that of Suricata
in having the parastyle and metastyle approximately equal in size and it
resembles M. mungo in this respect. The fossil M? is represented by three very
worn specimens and none exhibit any observable differences from the M?*
of the modern species.

Apart from their larger size (Table 64; Fig. 65), the most significant
difference between the lower teeth of the fossil and modern species is the fact

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 27%

TABLE 63

Dimensions of the upper teeth of Suricata major from Elandsfontein, compared with those of
modern S. suricatta and Mungos mungo.

P3 Ppt Mi
] b ] b l b

8669 — — 6:°7;0 8,0 ¢. 455 c. 8,5

6873 5,9 C. 455 Fe C. 755 = ==
Suricata a ee ee ee eee
major 6893 — — — — 4,2 8,2

16558 aes a a. ant 4,7 7,6

Mean 3,8 3,1 454 4,8 355 6,0
Suricata ee
suricatta Range | 3,5-4,0 2,6-3,5 | 4,2-5,0 4,0-5,7 3,1-4,1 5,6-7,0

n 8 8 8

Mean 455 4,0 5,2 535 3,7 6,2
Mungos |] |] |]
mungo Range | 4,5-4,6 3,8-4,1 | 5,0-5,3 5,2-5,8 | 3,4-3,9 5,8-6,5

n 3 3 - 3 2

that P, is present in the former. In only one of the seven hemi-mandibles repre-
sented is there no evidence of the presence of P,, although this specimen (8655)
may have lost the tooth during life. This was certainly the case with 6884, in
which traces of the P, alveolus are still visible. The fossil P, is a simple, more or
less conical tooth, which is very much smaller than P,. Morphologically P,
and P, resemble those of the modern species and are similar to S. suricatta and
M. mungo gothneh (Allen 1925: Fig. 50) in that they are relatively high crowned.
The P, differs from that of both modern species in lacking an anterior accessory
cusp, although this cusp was also virtually absent in two of the thirteen modern
S. suricatta specimens examined in the course of the present study.

In both modern species, the M, protoconid is a little larger than the meta-
conid, whereas in the fossil these cusps are approximately equal in size. They
are also less divergent than in M. mungo. The fossil M, differs from the moderns
in having the talonid as broad as the trigonid, while the paraconid and proto-
conid are more or less equal in size.

The lower cheekteeth are less crowded than those of S. suricatta and are
spaced as in M. mungo gothneh.

The fossil mandibular corpora are much larger than those of the modern
species, but they are similar in shape and have two mental foramina situated
anteriorly and posteriorly of P, as in the moderns.

€ € € € € € € € u
ONE LiL S°C OE hy ve 6b | TE Go| Be SY | kG Ene
—p‘€ -1‘9 = (0K MG SIS Gis AGH | SKC =n || 7G in se =O osuey osunu Sosun|y
s‘€ 89 SUC 87 Or €‘€ ig | jee 6r | 97 ey || Scr ES uvoy
s ees! rail
=)
a eee, eee eee ee eee ee en OO Oe
= am || ee ed ef Pe ee
= I'v OL VIZ GCG be Oe 6b | 67 OS | &e OY) OC Le
Z = L5G -0'8I So! i Geta Sy 9 67 017 || <8 UNS | =6,06 —OGelee Ole =O 6 osuey DIJDIIANS DIDIIANS
o g‘€ 89 061 LG 9‘€ I‘€ vv || eG Wie | ©C 9°¢ 61 €‘€ ueroW
% a ee | |__| fee
jaa
< a fe) (i a LE || Eva enn (ce mE ( SI PR NE SOR] Este OE
a0 = == a = = — LY L‘9 << = €€ mS = = 1889
2 p's 0'6 [Nlie = = = = | O72 79 I‘€ 6b | 67 yO | = = SOPs
e 6S 00! 0°67 2 LAS 0s ‘9 a = vy ¢‘9 = — = a MND CEH L616
a ¢‘S OO! C87 9 = — — — — — — — = an = = 6606 4ofou DIDI1ANS
E g‘s 66 = a == 9‘ €9 |] tr 9 | Ee? SSE | OF 0S NED) re €889
a 8p €‘Ol 0°87 2 = a cv CO | Sy co} PE GE | OS SP = = ¢s98
3 09 ETI oa = — — — = — — — = — == == 6998
g "WW MoOjeq | TA MOlaq I q I q I q I q I q I q I
snd109 Jo | snd1o5 jo
yIpeoig 1YysIOH "Wd “A WH Yd ®d *d ‘d

‘OSUNUL SOSUNTY PUL DIZVIUNS DIVILING UIBPOUL JO IsoYy} YIM poreduroo ‘uIayUOJspuLlY WoL solpw DJvILUNG JO 1991 JOMOT PU I[QIPUeUT dy) JO sUOISUsUTIC

to aldvy,

272

=I

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

"Yy1991 94} UO
TvoM pso0UvApe PUL oIvJopoUr “VYsI[S SuImoys ulajUOJspuLly wos (SSgg
‘6889 “Sobg) sajqipueur sofpm vyvoung ayy Jo smatA jeoong ‘bg “S14

I e | 1 An I

mn HULL Hi

-

S|

eI c| I I

HM

I

91

Sy

U

nnn

ANNALS OF THE SOUTH AFRICAN MUSEUM

274

‘osuNnU SOduNPY PU DYDIIINS °¢
UJOPOUI JO ssoy}) YIM poleduiod ‘soln vivnsng Jo *gq pue *q ay} Jo suotsusuIC “SQ “81,7

Jofew ejyeaiing ¥ B}}eIIINS eeIIINS oO ofunw

y Gus]
¥ E c

Yipeaig

sofunw e

yipeaig

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 275

Discussion

In their study of the mongooses, Gregory & Hellman (1939) recognized
three groups, one of which included the genera Mungos and Suricata. It is with
these genera that the fossil species evidently has its affinities. It also bears some
resemblance to Cynictis (e.g. braincase shape, presence of Pj), a genus which
Gregory & Hellman regarded as belonging in another group, but which had
paralleled the evolution of Suricata. The dentition of the fossil is much more
specialized than that of Cynictis and it is unlikely that there is any direct phyletic
connection between the two.

Before considering the possible relationships of the fossil species to Mungos
and Suricata, it is necessary to examine the differences between M. mungo
taenianotus and M. mungo gothneh. In several respects the latter is the more
specialized of the two subspecies and they were at one time recognized as
separate species (e.g. Allen 1925). M. m. gothneh has a shorter and more bulbous
braincase, a broader tympanic bulla, shorter tooth rows, while the P, and P,
are higher crowned than in M. m. taenianotus. In these respects the Central
African subspecies approaches S. suricatta, although in other osteological
characters and in its external features it is clearly closer to M. m. taenianotus.
The implication is that extant M. mungo includes a regional variant (i.e.
M. m. gothneh) which has paralleled the development of some of the osteological
and dental characters of §. suricatta.

The more important similarities and differences of the fossil to M. m.
taenianotus and S. suricatta may be summed up as follows:

(1) The size of the fossil is appreciably greater than that of the modern species.

(2) The configuration of the braincase is intermediate between that of M. m. taenianotus and
S. suricatta.

(3) The post-orbital processes are relatively longer than those of M. m. taenianotus, but are not
as well developed as those of S. suricatta.

(4) The general morphology of the tympanic region resembles that of M. m. taenianotus, but
the relative development of inflated parts of the bulla, and the nature of the transverse slot

of the ectotympanic are more like S. suricatta.
(5) The dentition is a combination of characters evident in both modern species.

The fossil species is clearly not readily referable to one or other of the
modern species and in several respects is actually intermediate between them. In
respect of skull morphology and dentition, M. m. taenianotus is less advanced
than S. suricatta while M. m. gothneh is advanced in the direction of S. suricatta
to about the same degree as the fossil species. In view of the temporal and
geographical separation between the fossil and M. m. gothneh, and the fact
that they do differ in some observable respects, they cannot be regarded
as conspecific.

It is concluded that the fossil species, although having some of the charac-
teristics of Mungos, is actually an early member of the Suricata lineage and it is
accordingly referred to this genus. M. m. gothneh is regarded as a product of
parallel evolution, its characteristics having developed either at a slower rate,
or after a later start.

276 ANNALS OF THE SOUTH AFRICAN MUSEUM

In deciding upon the specific identity of the fossil form, account was first
taken of Mungos dietricht Petter, 1963 from Garussi (= Ngarusi) and Olduvai
Bed I in Tanzania. The only specimen of this species which has been described
is a mandibular fragment with a complete series of cheekteeth (Dietrich 1942;
Petter 1963), although part of the upper dentition has also been figured (Petter
1969). The Elandsfontein species is a little larger than M. dietrichi, it has a
higher crowned P, and P, and less widely spaced lower cheekteeth. It also has a
smaller internal cusp on P%, while the P* protocone is more anteriorly situated.
In her illustrations of the M. dietrichi mandible, Petter (1963, 1969) does not
show an anterior accessory cusp on P,, but the original photograph of the
specimen subsequently designated the holotype of the species (Dietrich 1942:
pl. IV, 36) shows this cusp very distinctly. The two fossil species thus differ
in this respect as well. They are, however, similar in that both retain P,, and
probably also P!, and both have the M, talonid as broad as the trigonid.

Leakey (1967) gives the age of M. dietrichi as ‘lower Pleistocene’ and it is
therefore probably appreciably older than the Elandsfontein species. The indica-
tions are that the two fossil forms are not conspecific. M. dietricht might well be
ancestral to the Elandsfontein species, or both might have an as yet unrecorded
common ancestor further back in time. There is nothing in the M. dietricht
dentition which suggests that it belongs in the Suricata rather than Mungos
lineage, and in most respects it is not dissimilar to M. mungo taenianotus. There
can, therefore, be no doubt about its generic identity.

Fossil Suricata has previously been recorded from the Cave of Hearths in
the Transvaal and this material was referred to the extant species (Cooke,
in Mason 1962). In addition, Cooke (1955: 166) listed ‘Suricata sp. (or possibly
Cynictis)’ as part of the fauna from the Saldanha Lime Quarry occurrence.
This material has been lost and its affinities remain unknown.

There is no other recorded species to which the Elandsfontein form can be
referred and it is, therefore, given a new species name. The holotype is the only
group of specimens presently known in which cranial and dental elements are
associated, although since they are so fragmentary the description of the species
was based principally on the various referred specimens.

The phyletic relationships of Suricata major which are suggested here (Fig.
66), conflict with the conclusion of Petter (1969) that Suricata was derived from
Herpestes (Galerella). In the light of the preceding discussion and the fact that
there is a far greater similarity between Suricata and Mungos than between
Suricata and Herpestes (Gregory & Hellman, 1939), Petter’s opinion seems
largely insupportable.

There are further points which must be considered in respect of the
taxonomy of S. major. The differences between it and modern S. suricatia are
perhaps only a little more marked than those between the Cornelian Vulpes
chama and modern V. chama. Consequently, for the sake of consistency, S. major
should perhaps have been referred to as ‘Cornelian S. suricatta’ instead of having
been given a new species name. The distinction between the taxonomy of the

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 290

Lu
Zz
O
O Mungos mungo Suricata suricatta Herpestes
rs) Tropical Africa South Africa
As
Lu
a Suricata major
U South Africa
O
~
nn
i
al
a
?

Mungos dietrichi

Tropical Africa
Lu
4 ?
Lu :
UO
e)
a
a

Herpestes

Fig. 66. Tentative phylogeny of some Viverridae.

Elandsfontein Suricata and Vulpes was made simply because a Florisian form of
V. chama is known, whereas a Florisian Suricata has yet to be described. As a
result, the phyletic relationship between Cornelian V. chama and its modern
counterpart is more securely established than the suggested relationship
between §. major and S. suricatta. In addition, the characteristics of the various
fossil V. chama are such that it is difficult to make unequivocal definitions for
the different forms. This is not the case with §. major. The time may well come
when forms intermediate in age between S. major and S. suricatta are known and
a situation comparable to that involving the post-Makapanian V. chama is
encountered. For the present, however, a specific distinction seems warranted
and supportable.

It is worth noting that although there is no certain record of Suricata in the
south-western Cape in post-Cornelian times, S. major is the most commonly
represented viverrid at Elandsfontein. This is attributed to the fact that Suricata,

278 ANNALS OF THE SOUTH AFRICAN MUSEUM

at least at the present, is a colonial animal which occupies subterranean lairs.
The death of individuals in their lairs, coupled with a higher population
density, creates a more favourable opportunity for the preservation of their
remains than is the case with more solitary animals which spend most of their
lives on the surface of the ground. It follows that other carnivore species which
make use of subterranean lairs should also be relatively abundant as fossils
at this site and this is indeed the case with Mellivora capensis, to quote but one
example. The question of carnivore lairs at Elandsfontein will be discussed
again later as this is regarded as an important factor relating to the origin of
bone assemblages at the site.

The fact that neither Cynictis nor the ground squirrel, Xerus, both of which
commonly occupy the same burrows as Suricata today, are not recorded from
Elandsfontein, suggests that the association of these animals was not charac-
teristic during the Cornelian in the south-western Cape. The significance of
the disappearance of Suricata from this region is not known, but it may well
relate to environmental factors. )

Suricata is a genus endemic to the drier parts of southern Africa, while
Mungos is an essentially tropical form, which is confined to the more humid
subtropical east coast of South Africa in the most southerly part of its range.
There is apparently no overlap in the distribution of S. suricatta and M. mungo
taenianotus. The two modern species are not dissimilar in external appearances
(see Dorst & Dandelot 1970: 124, 128; pl 18), and bearing in mind the more
arid environment in which S. suricatta lives, its lighter coloration is not unex-
pected. Mungos and Suricata also share a number of habits. For example, both
are very sociable and live in warrens, they are diurnal, very noisy and have a
wide variety of calls, their diets are essentially similar and their mild disposition
make them easy to tame (Dorst & Dandelot 1970). These habits, which could
be regarded as heritage characteristics, add some substance to the theory that
M. mungo and S. suricatta had a common ancestor in the relatively recent past.
Possibly their separate identity was the result of evolution in different geo-
graphical environments.

The identification of S$. major in the Elandsfontein assemblage lends
support to Petter’s (1969) conclusion that the diversification of the African
viverrids took place comparatively recently.

The taxonomy of the Viverridae has long been problematical and con-
troversial. It was earlier concluded that Vishnuictis, with its curiously modified
skull, should rank only as a subgenus of Viverra, and there is a parallel with the
inferred relationships of Suricata and Mungos. In this case, however, the question
of whether or not Suricata should be regarded as a subgenus of Mungos could
be additionally investigated by further studies on the anatomy of the modern
species, as well as by further studies on non-anatomical aspects of the species.
Such investigations are clearly beyond the scope of the present study and so no
alteration to the presently accepted taxonomic status of Suricata is proposed,
although the possibility for a future change is recognized.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 279

Family Hyaenidae
Subfamily Hyaeninae
Hyaena cf. bellax Ewer, 1954

Material

Baard’s Quarry, Langebaanweg
L 179/11 A-C—Right P4 and part of C; left I°.

Discussion

These teeth were briefly described previously (Hendey 1970a) and tenta-
tively referred to Hyaena brunnea. While they are superficially similar to the
corresponding teeth of H. brunnea, the size of the P* (L 179/11C) and the pro-
portions of its buccal cusps (Table 65), suggest that the material has affinities
with H. bellax from Kromdraai (Ewer 1954). This is a Makapanian species
which is apparently closely related to H. brunnea (Fig. 17). The Baard’s Quarry
P4 lacks the protocone, but this cusp was perhaps more posteriorly situated and
a little larger than that of the H. bellax holotype. Although this tooth is similar
in length to the P4 of the ‘E’ Quarry Percrocuta, it has a much larger parastyle
and the two species are clearly not conspecific.

TABLE 65

Dimensions of the teeth of the Baard’s Quarry Hyaena cf. bellax, compared with those of the
H. bellax holotype and a series of modern H. brunnea.

Kromdraai Modern
Langebaanweg Hyaena bellax* Hyaena brunnea

L 179/11 l r Mean Range n
- l 12.0 11,2 11,5 | 11,4 11,0—-12,1 4

b 13,1 12,6 12,5 | 11,2 11,1-11,4
C ] 16,5 19,2 19,8 | 15,2 13,9-15,9 9

b 12,5 — 14,5 | 11,5 10,7-11,9
c. 40,0 39,2 39.2 | 34,6 33,2-36,2 13
| of parastyle 11,8 11,3 11,3 — — —
i 1 of paracone 13,6 13,3 13,1 — — ==
1 of metastyle c. 14,6 14,6 14,8 | — = -
P* metastyle | x 100/P#] 36,5 37,8 35,2* — 15

P4 parastyle | x 100/P4

paracone + metastyle | 41,8 40,6 47.9" — 15

* Ewer 1954.

The I? (L 179/11A), which is intact, is similar to those of both H. brunnea
and H. bellax, although it has a very stout root which is perhaps more in keeping
with the larger size of the latter species.

280 ANNALS OF THE SOUTH AFRICAN MUSEUM

The G (L 179/11B), which lacks most of the root and part of the crown, is
a little larger than those of the available modern H. brunnea series (n = 9),
but smaller than that of the H. bellax holotype.

The Baard’s Quarry hyaenid is evidently an Hyaena and, although the
available material is too fragmentary for confident specific identification, it is
tentatively referred to H. bellax.

Family Hyaenidae
Subfamily Hyaeninae
Hyaena brunnea 'Thunberg, 1820
(Fig. 67; also Hendey & Hendey 1968: pl. 6; Hendey 1968: pl. 9)
Present status

The brown hyaena is now extinct in the south-western Cape, but it apparently
still occurred in the region during the nineteenth century.

Material
(1) Elandsfontein

(a) Previously described specimens (Ewer & Singer 1956):
EC 2, 3, 4, 6—Maxillary and mandibular fragments with teeth as follows:
Right P, left P? to M!; right I, to M,; left C to M,.
EC 11—Right maxillary fragment with dp? and dp‘.
(b) Additional specimens:
15833 — Parts of the skull of a single individual, including: Incomplete brain-
case; facial region with right I° and P? to P‘, and left I? to G and P?;
right mandible with P, to M,; left mandible with C to M,.
Maxillary fragments with teeth as follows:
EC 33/34—Right P! to M!; left P® to M!.
16669 —Right P?.
Mandibular fragments with teeth as follows:
5307 —Left P, to P,.
6870 —Left P; and P,.
9562 —Right P, to P,.
16584—Right P, to M,.
16686—Incomplete C, P, and parts of P; to M,.
17125—Right P, to M,.
20041 —Right P, and P3.
20081—Right C to M, (apparently associated with left P! and C numbered
20074/5)-
Isolated teeth as follows:
A ce ps bProbably one individual
17128—Right P,.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 281

The following are the remains of juveniles:

3950 —Left mandibular fragment.

14153— Parts of the skulls of at least two individuals, including:
Left maxillary fragment with dp? and dp*, and M, just erupting; two
right mandibular fragments, one with dp, to dp, and the other with
unerupted M,.

16780, 20566 and others— Unerupted permanent teeth.

(2) Elandsfontein Wes
16317—Right mandibular fragment with C to P,; left mandibular fragment
with P, to M,.

(3) Melkbos

Mb 116/7—Right mandibular and maxillary fragments with part of P,, P,
and P,, and parts of P? to P#.

(4) Swartklip

ZW 2659—Left maxillary fragment with P? to M1?.

ZW 394 —Right premaxillary fragment with I’.

ZW 1836—Left mandibular fragment with incomplete C and P, to M,.

ZW 3790—Right P®.

The following are the remains of juveniles:

ZW 1311/3—An incomplete skull and mandible with teeth as follows:
Right dp’, dp* and M! just erupting; left P!, dp? to dp* and M!?
just erupting; right dt, dp, to dp, and M, just erupting; left dp,.
(SITE 2)

ZW 141 —Right maxillary fragment with dp* and dp?.

ZW 2662 —Right maxillary fragment with parts of dp? and dp’.

ZW 2677 —Part of dp,.

(5) Sea Harvest, Saldanha

S13 —Right maxillary fragment with P® just erupting.
S 200 —Incomplete left C.

S 133 —Right P4.

S 722 —Right mandibular fragment with P, and P3.

S 721 —Left mandibular fragment with P, and M,.

S 724/5—Left C.

S134 —Left Py.

S 36, S 207—Incomplete right M,; left M,.

S 37—Premolar fragment.

Description

Ewer & Singer (1956: 336) recorded some slight differences between the
lower teeth of the Elandsfontein material available to them and those of modern
Hyaena brunnea, but they considered it unnecessary ‘to place the fossil specimens
in a distinct subspecies’. Their observations were based on the dentition of a

282 ANNALS OF THE SOUTH AFRICAN MUSEUM

single adult individual (EC 2/6) and although some of the additional material
apparently represents the same form, other specimens evidently belong to a
second form which is itself slightly different to modern H. brunnea. The possi-
bility of the existence of a second fossil variety of H. brunnea in the late Pleistocene
of the south-western Cape has already been suggested (Hendey 1968).

The first form, which is regarded as the earlier of the two, and which
includes the specimens EC 2/6, EC 33, 6096, 17125 and 20081, differs from
modern H. brunnea in the following respects:

The M? is larger (Table 66).
The lower cheekteeth are generally smaller and relatively narrower (Table 67).

1)
)
) The P, anterior accessory cusp is relatively prominent.
)
)

2

The M, metaconid is relatively prominent.
The mandible and probably also other skull parts are slightly larger.

3
4
5

The second form, which includes the material from the Bone Circle
occurrence (15833), is larger than modern H. brunnea in most respects. For
example, although some of the teeth of 15833 are within the size range of varia-
tion observed in the comparative series, the mandible and braincase are larger.
The only convenient measurement which could be taken on the fragmented
braincase to illustrate this point was the maximum occipital width (99,7 mm).
This figure is appreciably greater than the mean of 13 modern H. brunnea skulls
(90,6 mm) and is outside the observed range of variation (86,8—94,0 mm).

Other specimens which are assigned to the later (Florisian) variety are
6870, 9562, 16669 and 16686. They resemble the earlier (Cornelian) form in the
size of the mandible, and probably also other skull parts, in having a fairly
prominent P, anterior accessory cusp and in having a relatively narrow M,
with a prominent metaconid. The lower premolars are, however, as broad as
those of modern H. brunnea.

The criteria used in separating the two Elandsfontein forms are insufficient
to allow for categorization of all specimens. The juvenile and poorly preserved
or incomplete adult specimens were not categorized, although they clearly
belong to H. brunnea. In addition, one of the mandibular fragments (20041),
which is somewhat unusual in its preservation, was not referred to either of the
two forms because of its exceptionally broad P;. This specimen is more similar to
corresponding parts of the mandible of modern H. brunnea than it is to other
fossil specimens from the south-western Cape, although its P, is broader even
than the broadest specimen in the modern comparative series (Table 67).

The specimens from Elandsfontein Wes, Melkbos, Swartklip and Sea
Harvest also represent a form, or forms, of H. brunnea larger than the moderns
in most respects. They are similar to the Elandsfontein Florisian variety in size,
the relative breadths of the lower premolars and in the nature of P, and M,.

Not all of these specimens can be distinguished from modern H. brunnea.
For example, the Sea Harvest lower carnassial S 207 falls within the size range
of variation observed in the moderns and has a reduced metaconid comparable
to that of the modern form. It is, however, a little narrower than any M,

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 283

FEU AULA LUT AL LL LC ra TURAL LLU LUA LLU TL LLL rn ii}

Fig. 67. Buccal and lingual views of the Hyaena brunnea mandible (20081) from Elands-
fontein.

ANNALS OF THE SOUTH AFRICAN MUSEUM

284

G‘o1 °9

G°C1 ‘2

‘s][npy sunox y

I] H |} rit Avery

=e = 9°91
660% 6°sE o‘gI
OC1% 9°SE 6°S1

| i _} ——— rr spur

—] ft} |] |] Yi] rit 272 FT

(| || | |

— — CC
rag OD Os —
G1% 6°VE 6°L1

G1
Q°ss o‘Q& 6‘QI
Vou = EE -9‘V1
V1z QVE 1‘QI
ot I Sl
rd.

ae 9‘
o'LI G‘g
0°61 —
Z°QI °9 —
o‘QI —
Sot a) —
L‘Lt z‘Q

—1'91 -9°9
691 aL
[ |

+ 5/VLo00z
g609
6690
9/% OF
69991
6EQS1
gIIl dW BOS P INA
06LE MZ
dippzemg
659% MZ
6E1¢
‘~51 G ‘00ZS JSIAIVFT BIG
u
osury pauunsg Duavd Fy
uLOpOW
Uva

“DaUUNLG *FT UJOPOUL JO SalJos & JO YIM poseduios ‘oder Us19}saM-YINOS IY} WIOIy VaUUNLG DUADATT {ISSO} JO Y}90} 19ddn 9y} Jo suoIsUsUICG]

99 ATdV]

285

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

“s]]Npe dunoxX »

ass —s 70S (angi €°7 ZES CII 917 €19 rere 661 6°99 56 Url rang S‘SI «18002
SPP = = —— IHG = = ¢‘% ‘9 aaa = — _— — — a — STIL] | wuo04
Pema | Aa eS a fe LL lL) Ls, SL a oe alta ek ao Sl el eer
O'S 0'9€ iy NS ee ee AS) ae 799 Det. .-.8'61 919 = O'OLC.s«B HI S21 ee 9/7OF 2
a i ese a a a a Nee ee eee ee il ee ee ee a
— at — a a2 9°6S orl ts 74 6°69 €‘Sl 6°17 — == a: ao ae OL89 a
ec PE | NS OF a a ee | ee a eee a
= O'lr 2 = co = 16S 6'€1 SEZ 169°9 27S 2 O77 ‘2 —— — L‘9| ‘2 ae — Z9S6 | W404 =
a a eS TST | GSE | | I ea SS a a | Ge ae Cea
0'8r 2 O'Ir O'7S ORL 2 10'se | Cies-2 “Rp? oOGe 2 | g'69°9 Csi 2° Tez p69 9 S‘Z1'9 O81 9 = aa 9899] e
¢‘tr 0‘0r oar a 9°St™ 92 | ¥'09 61 0'€Z Tel? “Got Teo. COL"? Le: plo | Ste sz €E8SI
— S‘I€ ae ao = = nel — bbl S‘rl S‘6l ZL 601 e's! — ns Ip007 | é
== s‘8e 9 | gos Lol 0'SZ 9°09 9 EPI 9972 | O'L9 7‘SI L‘% L‘69 CTI S‘9I e712 O's LIE9I SOM UI9}
-uojspurlq
— 0‘Or = =o == OLS 9. Cele? BL oe 8] PL, Sl. 20 te — — — 9‘TZI 9°61 LIL WW SOq[9W
= = v'0S €‘TI bbz L‘6S I'l 9°€Z 8°69 O's SIZ a a = = CLI | «9€8IMZ dijxeMg
‘ ‘ ‘ ‘ ‘ ‘ ‘ LOS
= == 9‘0S TTI 74 ae = = L‘OL 7S SIZ ae = ao — ae #bEIS
= = = Sel = = = = = = = = = = <= = #9£S | ISOAIBH POS
== — Sics Sel b'97 819 09! 6ST = = = = a = = = *«17LS
ar S‘6£ = a = = — = +69 os! 9°17 I‘€L cc L‘9I a = 7ZLS
€l S ra Cl €l II Ol u
9'tP 9'se 8°9S Gul 0'SZ 6°19 byl 8°€Z 6'IL Lvl b'IZ ILL ‘ZI 6‘SI 6‘€1 Gael pouunsq
-L°9€ -0'7E SEG SEG Syeheere “LT oG =pcle ele =0°99:. -6<I —S 6! =0L ° -so0l* SE pI Cel ee OIo) edury puavd H
T_T | UIBpPO[
TOP pve les €‘TI CAS PSs bel O'€7 9°89 Oa p07 I‘pl pit PSI OE! 8°9l urs
“A MOTEq | ®q MOleq I I ] I
g[qipuew | sqrpuew —— oe =s=5 —
JO 1Y4SZIOH | Jo IYsIOF OOIxq q l OOIXQd. 4 I OOIXd. q I OOIxq | q I
WH "d ®d *d 2)

“pauunsg * FJ UJBPOUL JO Satsas & JO asOY) YIM poredurod ‘ade+D ussd}sam-YINOs 3Y} WOT VaUUNLG DUADAT] [ISSOJ JO YII9} I9MO] Pue sa]qrpueUL dy} JO sUOISUDUTICT

Lo alav,

286 ANNALS OF THE SOUTH AFRICAN MUSEUM

in the comparative series and in this respect resembles most of the other fossils.
By contrast, the specimen S 721 is as broad as the moderns, although it is
larger in overall size and has a more prominent metaconid.

The upper dentitions of the fossil forms are less well represented and there
are fewer observable differences between them and the moderns. While the
larger size of the M! of the early Elandsfontein form is regarded as a primitive
and possibly useful distinguishing characteristic, the slightly larger size of the
M? of the Swartklip specimen, ZW 2659, is probably simply in keeping with the
overall larger size of the individual to which it belongs. As with the Canis
mesomelas from the site, overlaps in the tooth size ranges of variation of the
modern and fossil forms limits the usefulness of size as a distinguishing criterion.
Discussion

Although local Cornelian and Florisian forms of Hyaena brunnea can be
distinguished, and although both can be distinguished from that which is still
extant, it is not possible to define the categories in such a way that all the
available specimens can be classified. The differences between early and late
specimens can be striking, but it is evident from the total assemblage that there
is a gradation in characteristics from one extreme to the other. This point is
clearly illustrated by the following examples.

The Sea Harvest specimen, S 721, and a mandible from Plandsiadieam
20081, belong to young adults of approximately the same ontogenetic age and
can therefore be conveniently compared. The size differences between the P,
and M, of these two specimens exceed the ranges observed in the modern
comparative series, while the relative breadths of P, are almost the same
as the two extremes in the comparative series. The Swartklip fauna, which is
intermediate in age between the Elandsfontein Cornelian and the Sea Harvest
faunas, includes the H. brunnea mandible, ZW 1836, which also belongs to a
young adult. Theoretically, the P, and M, of this specimen should be inter-
mediate in character between those of 20081 and § 721. It is in fact found that
these teeth are intermediate in all dimensional respects (Table 68), although the
teeth are not dissimilar morphologically.

Exactly the same applies in the case of the P, and P, of mature adults from
Sea Harvest (S 722), Elandsfontein Wes (16317) and Elandsfontein (EC 2),
although in this instance there are no other grounds for believing the Elands-
fontein Wes specimen to be intermediate in chronometric age.

Owing to the variation encountered in all species, there obviously will be
exceptions to this pattern and the problematical Elandsfontein specimen,
20041, is a case in point, but the general trend in the evolution of the later
Quaternary H. brunnea of the south-western Cape is evident. There was appa-
rently an increase in size, at least in some respects, from the early (Cornelian)
form to a later (Florisian) form, which was eventually culminated in the Sea
Harvest (? early Holocene) H. brunnea. The Cornelian variety is characterized
by relatively narrow lower premolars and larger M!, in which respects the
later material is indistinguishable from the extant H. brunnea. .

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 287

TABLE 68

Dimensions of the lower cheekteeth of typical examples of Cornelian, Florisian and ?early
Holocene Hyaena brunnea from the south-western Cape, and mean dimensions of a series of
modern specimens from elsewhere in southern Africa.

Py P;
bX 100 b xX 100
] b ] ] b |
M.A. Modern Hyaena brunnea WEA: © Eb ISA 74,1 20,4 14,0 68,6
M.A. ?Early Holocene (S 722) 16,7 ~ 12,2 73,1 21,6 15,0 69,4
M.A. Florisian (16317) 16,5 11,5 69,7 22,7 15,2 67,0
M.A. Cornelian (EC 2) 14,8 10,0 67,6 19,5 1931 66,2
rs M,
bX 100 b X 100
] b ] l b l
M.A. Modern Hyaena brunnea 28.0. FS4. 58,4 23,2 12,3 53,1
Y.A. ?Early Holocene (S 721) 25,9 16,0 61,8 26,4 13,8 52,3
Y.A. Florisian (ZW 1836) 23,6 14,1! 5937 24,4 12,3 50,4
Y.A. Cornelian (20081) 21,6 11,5 5352 22,3 11,2 50,2
M.A.— Mature adult. Y.A.— Young adult.

All the local fossils tend to differ from the modern form in the following
respects:
1) The M, is relatively narrow.

(
(2) The M, metaconid is relatively prominent.
(3) The P, anterior accessory cusp is relatively prominent.

Since no skulls of the H. brunnea which inhabited the south-western Cape
in historic times were available, it is not known whether or not it also exhibited
these characteristics in its Py and M,. The local modern form was apparently
no different in size to the more northerly variety which is still extant, so the
large ? early Holocene Sea Harvest H. brunnea was apparently succeeded by
one of ‘normal’ size comparatively recently.

The presence of a prominent M, metaconid and P, anterior accessory cusp
are primitive characteristics evident also in H. brunnea dispar from Swartkrans
(Ewer 1955a) and H. bellax from Kromdraai (Ewer 1954). The relative breadth
of the M, in the former subspecies cannot be determined on the basis of the
described material, but the M, of H. bellax is even narrower than those of the
south-western Cape fossil H. brunnea.

288 ANNALS OF THE SOUTH AFRICAN MUSEUM

The published records of other fossil H. brunnea from elsewhere in South
Africa, namely, Florisbad (Dreyer & Lyle 1931) and Bolt’s Workings (Toerien
1952), do not make it clear if these specimens also had the primitive charac-
teristics of P, and M, evident in the south-western Cape fossils. However,
judging from the illustration of the ‘semi-fossilized’ Bolt’s Workings mandible
(Toerien 1952: Fig. 4), this specimen is more like modern H. brunnea than, for
example, the Sea Harvest specimen, S 721. Consequently, it is possible that
modern H. brunnea derives from more northerly fossil populations, and that the
south-western Cape late Pleistocene/early Holocene H. brunnea represented a
locally endemic and recently extinct lineage.

Family Hyaenidae
Subfamily Hyaeninae
Crocuta crocuta Erxleben, 1777

(Fig. 68; also Ewer & Singer 1956: pl. 27-31)

Present status.

The spotted hyaena is now extinct in the south-western Cape, but it was
apparently still common in the region during the eighteenth century.

Material
(1) Elandsfontein
(a) Previously described specimens (Ewer & Singer 1956):
EC 1—Braincase.
EC 5—Left maxillary fragment with P? and part of P+.
EC 8—Left mandibular fragment with part of M,.
EC 9—Right mandibular fragment with part of P,, P, and P,.
(b) Additional specimens:
15833 — Parts of a skull, including:
Maxillary fragments with right I? to P* and left P?, P? and part of P*;
mandibular fragments with right I, to M, and left P, and Ps.

(2) Bloembos
669—Braincase.

(3) Sea Harvest, Saldanha
S 203—Left P?.

Description

In their original account of the Elandsfontein Crocuta, Ewer & Singer
(1956) referred the material to C. spelaea, but in a subsequent reappraisal of
South African fossil hyaenids, Ewer (1967: 113) concluded that ‘there is no
justification for regarding it as differing from the extant species’.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 289

ll TU LULL nin UAL AL

mi

HU

WW

nm
0 it 1

WU
a oN

nl

OO nm ITU LAVA LULU LLU LLL ww WDUUYLUTULVAUUUY AAU UTE DU A

4 11S 116 117 118 119 210 2i1 3 2

Fig. 68. Buccal and occlusal views of the Crocuta crocuta mandible (15833) from Elandsfontein.

Little can be added to the published description of the Crocuta from this
site, although some of the incisors, the canines and the cheekteeth, with the
exception of M!, are now represented by complete specimens. None of the
teeth differ from those of modern C. crocuta and, as in the modern form, the M!
is small and only sometimes present. In 15833 the M! alveolus is preserved in
the right maxillary fragment, but it is absent on the left side.
| The preserved teeth of 15833 and the isolated P® from Sea Harvest (S 203)
are similar in size to the moderns (Tables 69, 70), although S 203 and the P,
of 15833 are slightly longer than those of the available comparative specimens.
They resemble the premolars of EC 5 and EC g in this respect. Most of these
teeth are, however, within the size range of variation observed in a large series
of East African C. crocuta (Kurtén 1956). Since the southern African variety of
C. crocuta is apparently a little larger than that from East Africa, it is probable

ANNALS OF THE SOUTH AFRICAN MUSEUM

290

‘9S61 u91INy x

OLGG9) OU IN elon ove — — — oa —_ — C9F
—_————-]-- | Ss] || uIo]UOJspuLryy
0.0z 9 Fok Gey VES Cr OPS ull, g‘L Vol oLi 9 SEQS1
— —- ELI 9°bs — = — — — — 60% S JSIAIVFT ba
ae L6 = L6 — L6 — _ u
— Z‘OV — 1°VS — ILI
— . -L‘6z — ~-2‘9gI ee LPI — --- — —- osury x (BOLLS Sey) 0jNI0L9 DjNI0L7 UIIPO|W
— ofGé — 1°1G — VV — — — — ueoyl
OI OI OI Ol 9 U
1‘°SS 6°96 6‘L1 L‘&% 6°61 G‘L1 OL, 06 gal oes
—L‘g1 —S‘SE —€‘91 -L‘1% —O11 8 -6°V1 v9 —1°L —O‘II -O‘QI osuey (volIpy °S) 07NI0L9 DINIOL) UIOPO|W
£°0% o‘Lé Tia 6‘zz 1°SI z‘Ql ol 1°8 Q‘II 9‘9I Uva
sl I I I I I il I Ges al
rd ed zd 1d O

"susUUIIads ULIPOUI JO SoIJas eB JO 9s0Y} YIM poseduiod ‘souTAOIg odeD UI9}s9M-YINOS IY} WIOIJ 07NI049 VjNIOI‘) {ISSO} BY} JO 4190) 19ddn oy} Jo suotsuswIG

69 aTav]

291

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

‘961 ug1INy y

-— —- g‘b1 Cbs ‘9 rC1 Q‘%z — ogi ‘9 — — 697
gal G‘o& 2 — = = = a = = = 8 OW SEO eee Ele
PSI 1‘0& 6S G‘1z PGI L‘0% 6‘o1 691 Lata aoe EEQS1
— G6 — 16 — g6 — 96 — u
L‘o& G‘bz 9‘%z ce
— + -9'%s = —S‘gI — —1‘Z1 — -g‘Il — — asury | y(Bouyy yseq) 7NI019 vINI0I UIIPOJY[
=o L‘9z — L's — 6°61 = GF oa — Uva
he ae oe an ee eee eee 6 | Fee
L‘S1 6°sE ov gba g‘G 1°SS 6‘11 zg! g‘b1 €‘g1
-I1‘II —1‘9Qz —E%I —_ -Q‘1Z —EF1 0% -1‘O1 —-OFI 6511 = -3°G asuey (wor *S) 07NI049 0jNI04-) UIOPOYY
ISI £°6% ca 6°ss €°G1 BS Oc11 PGI g‘Z1 g‘S1 uvoyy
q I q I sil I | I q I
'"N ‘d *d *d 2

*susWIIOdS UIOPOUI JO SITIOS & JO BsOY} WWM Poredwiod ‘vjnI049 01NI0L/) UII}JUOJSpUL|Y IY} JO YI99} JOMOT 9Y} JO SUOISUDUIIC]

of a1aV

292 ANNALS OF THE SOUTH AFRICAN MUSEUM

that all the south-western Cape fossil teeth are actually within the size range
of variation of the modern southern variety.

The Elandsfontein braincase has already been described in detail (Ewer &
Singer 1956), and the specimen from Bloembos is essentially similar. Both
specimens are comparable in size to the skulls in the available comparative
series (Table 71).

TABLE 71

Dimensions of the skull of fossil Crocuta crocuta from the south-western Cape, compared with
; those of a series of modern specimens.

Modern Crocuta crocuta
(S. Africa) Bloembos | Elandsfon-
a 669 tein EC 1

Occipital height (base of occip.

cond. to top of sagittal crest) 100,6 | 93,7—-106,1 5 c.99,5 | 104,8
Maximum width of occiput 101,6 =| 96,3—106,5 5 97.0 101,3
Maximum width across occipi-

tal condyles 52,4 | 49,9-53.9 5 52,0 47,0
Post-orbital constriction 48,4 46,9-50,4 5 49,8 —

Discussion

The fossil Crocuta crocuta from the south-western Cape is apparently indis-
tinguishable from the extant southern African variety of this species, although
it is perhaps slightly larger than the average size of the moderns. The original
material from Elandsfontein may be Cornelian in age, 15833 is from the
Florisian Bone Circle occurrence, the Bloembos skull is probably also Florisian
in age and the Sea Harvest tooth is regarded as early Holocene. The age cate-
gories cannot be distinguished from one another on the basis of the specimens
presently available.

Family Hyaenidae
Subfamily Protelinae

Proteles cristatus Sparrman, 1783

The aardwolf is not known as a fossil in the south-western Cape and until
recently the only known local record of this species was that of a specimen from
Stellenbosch referred to by Sclater (1900). During July, 1972 an adult female
was killed on the main road 31 km south of Hopefield (SAM 36685). Since the
aardwolf is a shy, nocturnal animal it may be more common locally than this
meagre record suggests, but its numbers must in any case be very limited. The
possibility that it is a recent immigrant to the region was mentioned earlier.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 2903

Family Felidae
Subfamily Machairodontinae
Megantereon sp.
Present status

Extinct

Material

Elandsfontein

3060 —Ascending ramus of a left mandible.

3058/9—Parts of the shafts of left and right humeri, probably belonging to the
same individual as 3060.

6762 —Left humerus lacking proximal end.

5410, 277—Left femur and tibia belonging to one individual.

9486 C, D—Right tibia and left femur, probably belonging to one individual.

321 —Right femur lacking distal end.

3461 —Proximal end of right femur.

Comment

Casts of two parts of a right mandible, an unnumbered specimen no longer
in the South African Museum’s collections, were also examined. The teeth

preserved are I, to C, P, and M,. These fragments may belong to the same
individual as 3058/60.

Description

The ascending ramus (3060) has the reduced coronoid process charac-
teristic of the machairodonts and is similar in size to that of the Megantereon
eurynodon from Kromdraai (Ewer 1955c) (Table 72). The unnumbered mandible
will presumably be described elsewhere, but judging from the available casts it
was on the basis of this material that the Elandsfontein machairodont was
identified as Megantereon gracile (see Boné & Singer 1965: Table 1), a species
which resembles M. eurynodon (Ewer 1955c).

TABLE 72

Dimensions of the ascending ramus of the Elandsfontein Megantereon, compared with that of
M. eurynodon from Kromdraai.

Elandsfontein Megantereon
Megantereon eurynodon
3060 T™ Ka 64*
(1) Height of ramus from inferior margin to top of coronoid
process : ' : : 2 45,0 655,550
(2) Height of coronoid process above condyle . : : 21,5 C. 24,0
(3) (1) :(2) 4 : : , : : : 1:0,48 & 120,47
(4) Transverse diameter of condyle : : q 28,8 =

* From Ewer (1955c¢: pl. 2; fig. 2).

2904 ANNALS OF THE SOUTH AFRICAN MUSEUM

The fossil humeri are essentially similar to those of modern felines in
morphology, but their proportions are different. The estimated overall length
of the most complete specimen (6762) is only a little more than that of humeri
of modern Panthera pardus, judging from two specimens of the small south-western
Cape variety, but it is considerably more stoutly built (Table 73).

The femora and tibiae are referred to Megantereon since they too belong to a
medium-sized felid with relatively short and heavy limbs. The femora are
larger than those of the P. pardus comparative specimens, while the tibiae are
approximately the same length as those of P. pardus although they are far stouter.
In three of the four femora, the lesser trochanter is as large as those of available
modern P. leo specimens (n = 7), although in overall size the fossil femora are
much smaller.

Discussion

The Elandsfontein machairodont is a species which is comparable in size
to Megantereon gracile from Sterkfontein (Broom 1948) and M. eurynodon from
Kromdraai. Ewer (1955¢c: 608) thought it unlikely that these two forms could
be conspecific, although the observable differences between them are slight.
The suggestion has already been made that the Elandsfontein form is conspecific
with M. gracile and on the basis of the material presently available, including
the casts, there are no grounds for doubting that this is the case.

If it is assumed that only one Megantereon lineage was represented in South
Africa, and once again there are no grounds for believing otherwise, then it
would be expected that the Elandsfontein Megantereon would have greater
affinities with the Kromdraai M. eurynodon, since this form would be the next
oldest in the series. The relative age of M. white: from Schurveberg (Broom
1937a) is not known.

The situation may be summed up by the statement that although the
Elandsfontein Megantereon is phylogenetically closer to M. eurynodon, it is
morphologically more like M. gracile. Since the fossil forms are so poorly repre-
sented, this statement cannot be substantiated. There is, however, a simple
and convenient explanation of the available record and that is the three forms,
and perhaps M. white: as well, are conspecific and that their supposed dis-
tinguishing characters are no more than individual intra-specific variations.
Just as temporal variants of a single species have already been recorded for
other South African Carnivora, so the Schurveberg, Sterkfontein, Kromdraai
and Elandsfontein Megantereon could be regarded as temporal variants of one
species.

A re-examination of all the South African Megantereon specimens now
available would no doubt resolve the question of their specific status and the
identity of the Elandsfontein material is, for the present, left in abeyance.

According to Kurtén (1968: 75) the body of Megantereon is characterized by
‘short but massive front legs and relatively feeble hindquarters’. The forelimb
of the Elandsfontein Megantereon is known only from the humerus, which is

295

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

yIsuZ] VIqry, :anwo7
Yyisus] INUIIy :sn19uIN;T

D “ag9gr6:zg9L9

26'0: 1 v6‘0: I
tate ai wiels t
vbr o'si 661
S'1z g‘Lt bbs
g‘1 L‘Lz o‘gt
b‘GP gb g‘ES
gt 1°66 G'S. +3
Of01S 0‘Z0% o*£0z
LlLo
L‘g1 gh -— G‘bz 6‘bz
HENS, ave — — aPS
‘ov Wee — — b1S
CIs L‘61 a‘oS o's —
ght o‘ov 1‘oL g‘hl _-
0‘Q%z o'G1z — — —
19h& 136 o1bS
GG QF 961
L‘gz L‘ba —
gov 9‘or -
0°S0% o‘s61 —
2 bd
1S09& JWVS boEQE IWYS gSo&
snp10g -q snp.0g "q

L‘g1 ; oo yeys Jo ‘weIp ‘suvi) ‘UIPy
L‘Gs * pus “}sIp }@ ‘urerp “ysod-"jue *xepy
9°gt ; : : * pUd "SIP je ‘WIeIp ‘sues) “xepy
g‘19 * pus *xoid je ‘uieip *ysod-"jue *xepy
o'1G : : : pus *xoid je ‘wierp ‘suvsy *xeyy
o'h61 : : : : YIBUg] []V1IVAG
ry99h6 VIGIL
ove ‘9 . ; ‘ *“-4yeys Jo ‘WIeIp ‘suvsy “UIT)T
G*LS -9 ; ‘ pus “jsIp 1e ‘urerp *jsod-"jue “xepy
6‘0S j : * pUd ‘jsIpP }¥ “UILIP ‘suvI} “XeTT
b°6z ; ae: * pvoy jo ‘urerp ysod-"jue “xeypy
g‘oL ‘ pues ‘xoid jv ‘wieIp ‘suei} *xeyy
o°ESz : ; ysug] [[e19AO
q9gr6 aoWaa
G‘oz , . yeyYs JO “WIRIpP ‘sueI) “UIT)T
bgf ‘ ‘ pud “jsIp je ‘wieIp “ysod-"jue ‘xepy
zg j , * puUd “]SIp je ‘UILIP ‘sueI} “xe/T
00&Z ‘9 : ; ; : pe aes ‘ yyBuzq] [[e12AGQ
zglg snuawnH

eee
“sping vsayjung UI2POUr JO asoy} YUM poieduioo ‘uossajuvsapy uloUOJspuLyy IY} Jo souoq [etues93sod ay} Jo suorsusuq

€L a1av yp,

296 ANNALS OF THE SOUTH AFRICAN MUSEUM

indeed relatively short and robustly proportioned. The hindlimb is repre-
sented by both femora and tibiae and, although they are stout, there evidently
was some reduction in elements of the hindlimb relative to those of the forelimb.
The humerus is slightly longer relative to the femur compared to those of
P. pardus, while the tibia is appreciably shorter than the femur.

The femur and tibia, 5410 and 277, which evidently belong to the same
individual, are of interest since they exhibit pathological conditions similar to
those observed in the Langebaanweg Machairodus. There is osteitis of the distal
and proximal ends of the femur and tibia respectively, and facets of eburnation
which correspond to one another are worn on adjacent medial articular sur-
faces. Although the total number of machairodont limb bones from Langebaan-
weg and Elandsfontein is small, an appreciable proportion exhibit some sort of
bone pathology.

Family Felidae
Subfamily Felinae
Felis libyca Forster, 1780
(Fig. 69; also Hendey & Hendey 1968: pl. 7C)
Present status

The African wildcat still occurs in the south-western Cape, but it is now
rare.

Material
Swartklip

ZW 110 —Part of a skull comprising the braincase and posterior portion of the
left maxilla with damaged P*.

ZW 1841—Part of the facial region of a skull, the dentition of which lacks the
right P?, part of P*, and M!.

ZW 3487 —Part of the facial region of a skull with incisors, canines, left P? and
meht Pe:

ZW 377 —Part of the facial region of a skull with right I? to G, and left I’ and
re.

ZW 1843—Left maxillary fragment with P? and P*.

ZW 183 —Left P*.

ZW 178 —Left mandibular fragment with Ij, C and part of P3.

ZW 3489—Right mandibular fragment with P, and part of M,.

Comment

The difficulties which may be encountered in the identification of fossil
felines were referred to earlier and two of the above specimens, ZW 110 and
ZW 183, were previously tentatively identified as belonging to Felis serval
(Hendey & Hendey 1968: 63). It is, however, evident that they belong to a large
variety of the wildcat rather than to a small serval.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 297

Description

The smallest felid from Swartklip differs from the extant southern African
form of the wildcat, Felis libyca cafra, principally in its larger size and in the shape
of the braincase. The fossil braincase ZW 110 is comparable to the ‘narrow-
waisted’ type which characterizes the northern races of F. libyca (see Pocock
1951: Fig. 4). It is appreciably longer than the braincases of the available
F. libyca cafra specimens (n = 6) (Table 74). In addition, the post-orbital
processes are long and narrow, which is also a feature of the northern varieties
such as F. libyca ugandae. The sagittal crest is well developed and extends
anteriorly almost as far as the post-orbital constriction. It was much less promi-
nent in the F. libyca skulls examined in the course of the present study (n = 7).

1

1

8

ill

4

5

mm
113 7 1

|
1

|
1

mm
6

Fig. 69. Ventral view of the Felis libyca skull
(ZW 1841) from Swartklip.

The facial region of the Swartklip wildcat is best represented by the speci-
men ZW 1841, which includes the complete dentition on the left side. The face
of the fossil form is larger than that of modern F. libyca and this is indicated by
the maximum width of the snout anterior to the infra-orbital foramina (‘rostral
width’ of Table 74). Most of the upper teeth are also larger than those of the
moderns (Table 75), although P? and M!? were actually a little smaller than the
means of the comparative series. Consequently, the small size of these teeth
relative to P? and P? is more pronounced in the fossil form. In modern F. libyca
P? is sometimes absent, but in those specimens in which the relevant part of the
maxilla was preserved (ZW 377, ZW 1841, ZW 3487) the P?, or its alveolus,
was present. There are no differences in the morphology of the fossil and modern
teeth and a similar variability in features was evident. For example, a small
anterior accessory cusp is present on the left and right P? of ZW 1841 and in the
P3 of ZW 1843 it is represented by a trace, while in the available skulls of

ANNALS OF THE SOUTH AFRICAN MUSEUM

298

G G G ¢ 9 G U
9°Vo—0'ZS G“Ch_-9'1P o‘Qb_-V‘GP 9‘aS—6‘Qh 9‘Lb-2'b& L‘Vs—-GS‘ao asury DIGy SYay ULNPOW
L‘So 9&P G‘ov 01S 0‘9& G‘Ss uvay
— — — — — o1E IVQI MZ
——_—__|—.-$]$}$?-—| :&Y- 1 - |] | HH dipyems
L‘Lo G‘60 °9 02S 00g G‘o§ — Oll MZ
sjesourenbs (uOT}911}SU09
so[Apuoo *di1990 yIpIM ye YIPIM [e11q.10-jsod UOI}011]SUOD yIpIm
SSOIO® YIPIM ployse yy oseoUIeIg UIOIJ) YISUIT [e11qG.10-1sog [e11s0y
UINULNIXe I] gseoUIeIg

‘sudULIDedSs UJIPOU JO Salios & JO 9sOY) YIM posvduro0s ‘nIdgQy sijay AIpyIWeMG IY} JO [[Nys sy} Jo suotsusuIcy

bL alavy,

299

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

5 9 9 9 9 s u
1°3S bY bg arch | rl 62 as os ob BG
—6'gz —2‘¢ -6°F -L‘o1 —6°E bg | -S‘1 -6% | -HE —S‘P asury vIdgu Stjaf UIIPOW]
bof gE 9‘G G‘rI L‘°t ofL 6°! Lz L‘g QP rns
= — 19 oF! — — — 2 — — SBI MZ
— —~ 9‘S o*G1 ab gil — —— — — EbgI MZ
— = — — — — — — . oS 09 LLE MZ dipywemMsg
aces 1°E gS ESI 6°E gl g‘I gz oF ofS IvgI MZ
td-D WN td ed ed Oo

‘suswtoads UI9POUI JO satios & JO soy} YIIM Poseduros ‘v9¢g17 sya dipyeMg IY) JO YyII03 saddn 9y} Jo suotsusuIq

GL alavy,

300 ANNALS OF THE SOUTH AFRICAN MUSEUM

modern F. /ibyca it was absent in two specimens, represented by a trace in three

and present in two.
The lower teeth of the fossil form are within the observed size range of

variation of modern F. libyca (Table 76), but the mandibular corpus of ZW 3489
is larger than any of the moderns.
TABLE 76

Dimensions of the lower teeth of the Swartklip Felis libyca, compared
with those of a series of modern specimens.

Swartklip

ZW 3489 — — 79 3,6
Mean 4,4 3,4 It 9359
Modern i EG Lia ae SA
Felis Range LO= ye 6,9- 3,3-
libyca 533 4,0 8,4 3,9
n 5 6

Discussion

The Swartklip wildcat is comparable to some of the other carnivores from
this site in that it is larger than its modern counterpart. Otherwise differences
between it and modern F. libyca are not marked, although the relatively small
size of P? and M! may prove a significant differentiating characteristic.

The relatively elongated and ‘narrow-waisted’ braincase which it has in
common with the more northerly varieties of F. libyca is perhaps a more primi-
tive condition than the shorter and ‘broad-waisted’ braincase of the extant
southern African subspecies (F. l. cafra, F. 1. griselda, F. |. melland:).

Although the: Swartklip wildcat does differ in certain respects from the
extant local variety, it is undoubtedly closely related and it is distinguished from

the moderns by being termed ‘Florisian F. libyca’.

Family Felidae
Subfamily Felinae
Felis serval Schreber, 1776
(Fig. 70)
Present status

The serval is now extinct in the south-western Cape, but it still occurred

in the region during the nineteenth century.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 301

Material

Swartklip

ZW 1480/1 — Parts of the skull of a single individual, including:
Right premaxilla and maxilla lacking only I’ and P?, found in
articulation with a right mandible with I, to M,; left maxillary
fragment with P? and part of P*, found in articulation with a
left mandibular fragment with P, to M,.

ZW 1448 —Left maxilla with P? and P?.

ZW 1649 —Left C.

ZW 2950 —Right mandibular fragment.

ZW 3438 —Right mandibular fragment with Cite’ M,.

ZW 3488 —Left mandibular fragment with P, and M,.

Tentatively referred material
Sea Harvest, Saldanha

S 202—Right mandibular fragment with P, to M,.
S1 +Left mandibular fragment with M,.
S 216—Left M,.

Description

The material from Swartklip belongs to a felid which is comparable in
size to the extant Felis caracal. This species is usually larger than F. serval in
respect of skull and tooth size, although overlaps in the size ranges of variation
were observed in the available series of modern comparative specimens (F.
caracal n = 27; F. serval n = 13). The fossil teeth were all larger than those of
the F. serval series (Tables 77, 78), and in most respects were within the ranges
of variation observed in F. caracal. In other respects, however, the dentition of
the fossil form is most similar to that of F. serval.

The most important F. serval characteristic exhibited by the Swartklip
fossils is the presence of P?. In ZW 1480 the root of this tooth is preserved, while
in ZW 1481 and ZW1448 the P? alveoli are present. Judging from the available
comparative material, P? is invariably present in F. serval, whereas in only 3
of the 27 skulls of F. caracal was it observed. In one of these specimens (BMNH
2.9.1.20a) the P? is comparable in size (3,2 x 1,9 mm) to that of F. serval, while
im) tae others (SAM 35101, 36268) it is very small (1,5 Xc1,2; 1,3 X1,0).
Judging from the P? alveoli of the fossil specimens, these teeth were comparable
in size to those of modern F. serval and they occupied much of the space inter-
vening between C and P®. The P? alveolus of ZW 1480 measures 3,8 x 2,2 mm,
while that of ZW 1448 measures c4,5 X 2,7 mm.

There are few other constant differences between the dentitions of F. serval
and F. caracal. There is apparently a tendency for the posterior keel of the
F. serval P, principal cusp to be more steeply angled than that of F. caracal,
while the P, principal cusp of F. serval tends to be higher crowned. The anterior
accessory cusps of these two teeth are a little more prominent in F. serval,

ANNALS OF THE SOUTH AFRICAN MUSEUM

302

or

0‘66-6'z&

9‘SE

GSS Oe ee ee Oe Oe sss

G1

G<G-oԤ

GD

0‘9

J9}9UIeIp
OSIOASUCL J,

WN

"J00daS * 7 UIOPOU JO 9SOY} YIM poreduso0d ‘7vaLas stay AipyIIeMG IY} Jo W190} 19ddn oy} Jo suoisusuIIq

rd

*((Ar0jsIFY [eINeNY) WANasNyY YsIIG UT sUsUIIDOdS) voIIpY Iseq pu [eI]UID

‘[evAsULI], ‘UIOOI}SJOYIIOg z

GI

ed.

0‘6

G‘9

£6

€1
Vs CP
—6'1 —9‘%
OF TWAS
CS 1‘V
L's rch 4
q I
zd.

LL a1av J,

0G

G1

Ol

J0UIAOIg 9der) UI9}SIM-YINOS “JS9AA JOSAIWIOG _

osueyy

ues

1°92'9'66
HN

6SE9& WVS
6791 MZ
ghhi MZ

OgVI MZ

e]DALS SUA

2/DaLas Siaq

q]Dasas sya

dippzemg

393

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

*((ArojstPy [eanyeNy) Winasnyy ystig ur susumdeds) vorypy jseq pue [eUayD ,
‘Q0UTAOIg odeD U19}SaM-YINOS “jS9AA JOSIBWIOS ,

*jevAsue1y, ‘UI00I}SJOYII0g z

g]DQLS S177

g/Dasas SIaq

Joasas SYaT

1 GI oI GI GI 1
3°9 ae EVI 1‘SP gv P11 Gv £01
-3'v = Ah —S‘o1 —G*ce —3'v —06 | -L‘E -3‘8
9S G‘9 6°11 1°9f GP zor | oF v6
3°9 3°9 1S €‘sP ofS OCI vv O11
G°Z L‘9 of G‘sP gv 6‘o1 | HP L‘ol
— — — — 9S Q‘rI = —
— £'g 9 Gr - LG PSI — —
aL LL 6‘F1 — EG C1 | 6F g‘or
— vg ofS — — — — —
— oy QS — GG Pir | .G° Q‘I1
38 == —= — E'S G11 | 9g? OcII
ocL 6° oS o‘Gh 6°G Grr 1°G eG
(tjooaye (oRjans I q I q I
usamyoq) | Wl Mojeq | yesonq)
eulayseip | sndioo jo | "Ww Mojoq 4N-O WwW |
r) 180g yjpesig sndioo 7
a Jo 1SI0Hy

1)

JSOAIVFT BIG

dippzemg

"1004S “+7 UIIPOUL JO ISOY} YIM poieduiod ‘pvasas sia y2aIeP] VIG pue dipyIEMG 9} Jo W199} IaMO] PUL So[qIPUeUT 94} Jo SUOISUSTIG,

gl a1aV I,

304 ANNALS OF THE SOUTH AFRICAN MUSEUM

Il

1

|

Fig. 70. Buccal and occlusal views of the Felis serval
maxilla (ZW 1480) from Swartklip.

HATA

while the upper canine of this species is more deeply grooved on its buccal
surface than that of F. caracal. In all these respects the Swartklip form resembles
F’. serval rather than F. caracal.

In the modern varieties of these species the proportions of the individual
teeth are essentially similar, although there appears to be a fairly constant
difference in the lengths of P, and M, relative to one another. In F. serval the
tendency is for these teeth to be approximately equal in length, but in F. caracal
M, is apparently always longer. The former condition is evident in the Swart-
klip fossils (Tabie 79).

The only other observable skull character which may be of significance in
the present instance is in the nature of the external opening of the infra-orbital
foramen. In modern F. serval and in the fossil that part of the maxilla which
forms the lateral margin of the infra-orbital foramen projects further forward
than in F. caracal.

The specimens from Sea Harvest belong to a felid which is comparable to
that from Swartklip in almost all observable respects. A problem in identifica-
tion does arise in connection with S 202, because the relative lengths of the

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 305

P, and M, are comparable to those of F. caracal rather than F. serval. On the
other hand, the P, and P, of S 202 are similar to those of F. serval in respect of
their morphology. The isolated lower carnassial, S 216, is an unusual tooth in
that it has a small cusp projecting from the lingual cingulum at about the mid-
point of the tooth opposite the carnassial notch. No such accessory cusp was
observed in any of the other felid lower carnassials examined in the course of
the present study, and it is probably an individual peculiarity which is of no
real significance.

Discussion

Relatively few of the characteristics which distinguish the modern varieties
of F. serval from F. caracal are observable in the Swartklip material. It is, however,
really only in respect of size that the fossils resemble F. caracal rather than F.
serval and the material is accordingly referred to the latter species. This is yet
another species from Swartklip which is larger than its modern counterpart.

The Sea Harvest specimens are less certainly referred to F. serval. This
reservation is based largely upon the relative lengths of the P, and M, of S 202.
However, although it is usual for these teeth to be approximately equal in
length in F. serval, there are individual exceptions. For example, in the specimen
BMNH 99.10.23.3 the P,:M, length ratio is 1:1,24, so that in this instance the
M, is relatively longer even than that of S 202 (Table 79). Only when other
cranial and dental characters of the Sea Harvest species become known will its
identity be firmly established.

TABLE 79

Comparisons of the lengths of P, and M, of the Swartklip and Sea
Harvest Felis serval, modern F. serval and modern F. caracal.

IP, IM, IP,:1M,
Felis caracal (n = 19) it 1o7. 12.2 I:1,13
Sea Harvest F. serval (n = 1) 10,8 12,3 eee
Felis serval (n = 13) 9,4 10,2 1:1,08
Swartklip F. serval (n = 3) Mia, RIG 1:1,01

The record of F. serval from Swartklip and Sea Harvest indicates that this
species was not uncommon locally during the Florisian and early Holocene.
Since it was still present in the region until fairly recently, it is probably one
of the many species which became extinct as a result of human activity. This
is in Curious contrast to the similar sized F. caracal, which has survived locally
in spite of determined efforts aimed at its eradication. The serval apparently
still occurs, but is rare, in the eastern Cape about 750 km east of the area

306 ANNALS OF THE SOUTH AFRICAN MUSEUM

under consideration. Its disappearance from the large tracts of un- and
under-developed country in the southern Cape is also remarkable, since the
caracal still occurs there in greater numbers than it does in the south-western
Cape.

It is possible that the species was already in decline in the most southerly
parts of South Africa and that human activity in the historic period merely
hastened its extinction. There is, however, no real evidence to support this
and as late as 1775, more than 100 years after the first European settlement at
the Cape was established, Forster (1781:4) recorded the conspecific F. capensis
near Cape Town, reporting that it still occurred in ‘mountainous and woody
tracts’ near this city. As far as could be determined, the last local record of this
species was a specimen from Somerset West near Cape Town which was
acquired by the South African Museum in 1898 (SAM 36359).

A second possibility to account for the local disappearance of F. serval
and its apparent absence from the southern Cape is that Forster’s F. ‘capensis’
represented a relict population of F. serval, which survived for some time away
from the main area of F. serval distribution in tropical Africa. There are no
obvious differences between the Somerset West specimen and others from
elsewhere in Africa, but detailed comparisons have yet to be undertaken.

At least one other extant carnivore species, namely, Viverra civetta did
disappear from the south-western Cape and adjacent regions during the latter
part of the Quaternary and there are certainly no indications of human involve-
ment in this instance. There might well have been natural limiting factors
operating in the case of F. serval as well.

Family Felidae
Subfamily Felinae
Felis caracal Schreber, 1776

Present status

The caracal still occurs in the south-western Cape, but it is classed as vermin
and threatened with extinction.

Material

Elandsfontein

15833—Left mandibular fragment with C to M,,; isolated right P, and Mj.
20072 — Right mandibular fragment with P, to M,.

Comment

The material 15833 is from the Florisian Bone Circle occurrence, while
the other mandibular fragment, 20072, is in an unusual state of preservation
and may also be post-Cornelian in age.

6

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

P v v v v b v u
rb v6 b‘E1 SG OSI 6b gol | oF 19 | EL (worypy “q)
-0'9 -9'9 GI -0'S —b11 | -S*P -g'6 | -9'f -9‘9 | -9‘P -g‘S aBur yy 1090409 * J UIIPOYW
L‘9 vg 1°S1 EG QiIl Lv z‘Ol 6°E C‘Z 1G L‘g urs
C1 GI Gi GI GI G1 oI u
G‘9 o‘O1 G‘gi 19 oF GG PSI hd £6 1‘9 £9 (eorayy *S)
—EP —o‘L —2Sl -6‘P -of11 | -PP -ofo1 | -—9‘& —b‘L | -S‘P -~3'9 aBury 1090409 “JT ULIPOYW
v9 S‘°9 oF GG are | oS 6‘o1 ob zg 1°¢ 6‘9 uvody
SL *9 gil €E1 09 Gor | 3G Sor | oF a — — zLooz ulayUOJspuLly
89 G‘6 ofS gS oF GG aD | GP £6 1‘9 £9 zLogt WVS 090409 "J
UIIpoyy
g‘S C6") C°Gr *9 L°¢ G°E1 PG PSI gb Oi) SoS L°L €E9S1 ula} UOJspur|y
(oRjans q I 4 I q I 4 I
BUIO}SeIpP NAL Mojoq eoonq)
ouruvo sndioo NA Mopeq ’
-1S0g jo W)peoig sndioo W *d “d e)
JO 1Y4S1IOH 2

"]0904D9I “J UIBPOU JO 9SOY} YUM poiedurod uloyUOJspuL][y WOIJ 709IV4D9I $1797 JO YIII] I9MOT PUL So[qIPULU IY} JO SUOISUDUIIP 3],

0g ATaV I],

308 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

The Bone Circle specimens probably belong to a single individual and they
are virtually indistinguishable from corresponding parts of the mandible of a
large Felis caracal (SAM 36072) included in the available modern comparative
series. Although they are also similar in size to the Swartklip F. serval specimens,
they exhibit none of the features which characterize this form.

The specimen 20072 belongs to a smaller individual, which is comparable
to average-sized specimens in the F. caracal comparative series (Table 80).
The dentition is indistinguishable from that of F. caracal.

Discussion

Although the mandibles of F. caracal and F. serval are less easily distinguished
from one another than the skulls of these species, the fossil specimens are so
similar to F. caracal that their identification with this species is unavoidable.

Since the Mellivora capensis and Canis mesomelas from the Bone Circle are
larger than their modern counterparts, it is possible that the same applies in the
case of the F. caracal. ‘Thus the recorded material may represent an average-
sized individual of the fossil population, rather than an unusually large indivi-
dual as was the case with SAM 36072, the modern specimen to which it com-
pared most closely in size.

The smaller specimen, 20072, could be even younger than the Bone Circle
material and since it belongs to an individual similar in all observable respects
to an average-sized modern caracal, it might well be Holocene in age. The possi-
bility that it predates the Bone Circle material cannot, however, be dismissed.

The absence of F. caracal from the Swartklip and Sea Harvest assemblages
is surprising in view of the Elandsfontein and recent records of this species. As
the Swartklip and Sea Harvest sites are both hillside animal lairs, the absence
of F. caracal may simply indicate that this species did not make use of such lairs.

Family Felidae
Subfamily Felinae
Felis sp(p).
Material
Elandsfontein
Mandibular fragments with teeth as follows:
EC 38—None.
8402 —Right P, and P,.
9198 —Incomplete right P,.
11127—Incomplete left C.
Isolated teeth as follows:
7086— Right M1.
7527—Left C.
7787—Left P,.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 309

Discussion

These specimens represent only a part of a larger assemblage of remains
belonging to a caracal-sized felid, or felids. The remainder of the assemblage,
which includes the specimens tentatively identified as Felis serval by Ewer &
Singer (1956), is no longer in the South African Museum’s collections and will
presumably be described elsewhere.

At least one of the specimens listed above, 9198, differs in certain respects
from those previously assigned to F. serval and F. caracal, while another, 8402,
may belong to a large F. caracal.

J udging from their preservation, these specimens belong with the Cornelian
element of the Elandsfontein fauna and since an assessment of this material
could be better made in conjunction with the rest of the assemblage, it is not
described in detail. The indications are, however, that there is at least one
additional caracal-sized felid in the Elandsfontein assemblage. On the basis of
the observations made on other Carnivora from this site, it is most likely that
the present material represents Cornelian varieties of the serval and caracal.

Family Felidae
Subfamily Felinae
Panthera pardus Linnaeus, 1758
(Fig. 71)
Present status

The leopard became extinct in the south-western Cape during the nine-
teenth century, but small numbers still occur in the mountains adjacent to
this region.

Material

(1) Sea Harvest, Saldanha

S 131—Right P*.

S 132—Right M,.

S 213—Premolar fragment.

S 32 —Femur lacking distal end.

(2) Fish Hoek
Q 1800—Right mandible with C to M,.

Description

The Fish Hoek mandible is essentially similar to those of a series of leopards
from the Cape Folded Mountains (n = 6), although the cheekteeth are rather
narrow. (Table 81).

The Sea Harvest specimens are all appreciably larger than corresponding
elements in the comparative series, but they are similar in all morphological
details.

ANNALS OF THE SOUTH AFRICAN MUSEUM

310

9 9 9 9 9 s v 9 u
= | | | J ——_____—_——_. snpsvg
Svan gba O°sV 1 ey tert £9 na 09 6°11 96 = ga Q°a1 6‘vz psayjuvg
-9‘O1 —L‘0% =OFE 11 =G°f =1°G1 =a'L -9‘V1 | —2°S =s-O1 |) —GL, =S"o1 Vor —-L‘oz suey UIONPOW
Q‘rI VSS 6.081 Or HEI g‘L Z‘QI 9‘S P11 6.9 Bal 6°11 oS uvayy
Sron 03S ‘9 OOo! 2) | chee GOI OL L‘g 6G Q‘1I Cie) ONG) — a 00g! © Yoox] USsty
— — — Onn ite — — — — == = G°G1°2 Gen) GE1 G11 || JSoAIvET vos
"W Moped | TAL Moped | 2[qrpueur q I q I q I q I q I
o[qipuew | sfqipueu | jo wsuoy W "d *d 9) rd

jo yipeoig | Jo yidoq | [[e12A9

‘UOISII SITY} 0} JUDDe[pe suTe}UNOUI
wio1y spiedoa] U1apoUr Jo satias v Jo asoy) YIM poreduioo adery usI}s9M-YINOS DY} Woy snpsvg vsayjUYG ISSO} JO 2[qIPUeU PUL Y199} 24} Jo SUOISUDUTIG]

19 AIGV

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 3II

Discussion

The Fish Hoek specimen, which might well be very recent in age, appa-
rently represents the small variety of leopard which still occurs in the mountains
adjacent to the south-western Cape.

By contrast, the ? early Holocene Sea Harvest material represents a very
much larger variety.

ile il7 als

a

Fig. 71. Buccal views of the Panthera pardus M, (S 132) from Sea Harvest and the mandible
(Q 1800) from Fish Hoek.

Panthera pardus is an exceptionally widely distributed mammalian species
and a large number of subspecies have been named. Although the taxonomy
proposed by Pocock (1932) has been modified by more recent work, the wide
variation encountered within the species can be judged from Pocock’s observa-
tions. Amongst the measurements of modern leopard skulls given by this
author are the lengths of 112 upper carnassials and 102 lower carnassials. His
figures represent both males and females from throughout Africa and all these
specimens are exceeded in length by the Sea Harvest carnassials. Broom (1948)
gives the length of the largest modern leopard P* recorded by him as 28,2 mm,
which is still less than the 28,5 of the Sea Harvest specimen, S 131.

Smithers (1968: 12) discussed the taxonomy of modern P. pardus and
observed that, ‘In some cases attempts have been made to separate sub-
species into larger and smaller types on skull and tooth measurements of
comparatively few specimens, yet in some areas, e.g. Rhodesia, exceptionally

312 ANNALS OF THE SOUTH AFRICAN MUSEUM

large individuals do occur within populations of generally smaller size’. Since
the Sea Harvest material may all belong to a single individual, it may simply
be such an exceptionally large example of the species. However, other Sea
Harvest Carnivora, such as Hyaena brunnea and Herpestes pulverulentus, are also
larger than their modern counterparts in some respects, so it is more likely that
the leopard material represents a large variety rather than just an exceptional
individual.

The only other fossil leopard from South Africa which has been described
is the Makapanian P. pardus incurva (Ewer 1956a), a variety which is probably
a little smaller than the modern form from the Transvaal. The leopard is,
therefore, yet another carnivore which underwent size changes during the
Quaternary and, although their fossil record is poor, the Sea Harvest variety
may well be one of the largest which occurred.

Family Felidae
Subfamily Felinae
Panthera leo Linnaeus, 1758
(Figs 72, 73, 74; also Hendey & Hendey 1968: pl. 7A, B; Hendey 1968: pl. 10)

Present status

The lion is now extinct in the south-western Cape Province, but it was still
common in the region during the seventeenth century.

Material
(1) Elandsfontein

The only known cranial remains from this site, together with some asso-
ciated postcranial bones, are no longer in the South African Museum’s collec-
tions. A 2nd phalanx (9664) and parts of a radius (9054, 9632A), which appa-
rently belong to the same individual, are still available, as are the following:

Fore- and hindlimb:
8487A-D—Metacarpal, metatarsal and restored femur and tibia of a single
individual.

Forelimb:

16691A—C— Restored humerus, ulna and radius.

8377—Parts of left and right humeri, ulnae and radii.

20743 — Radius and part of an ulna.

11202B, 12781 —Humeri lacking proximal ends.

4331, 9115, 13715, 14763, 16670, 16795, 20216, 20217, 20416, 20422—Distal
ends of humeri.

273, 11156, 11202A, 20429— Ulnae lacking distal ends.

270, 5084—UlInae lacking distal ends and parts of proximal ends.

11154, 11155— Proximal and distal ends of a radius.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 313

5845, 11510, 20040, 20641, 21010— Proximal ends of radii.
20430—Shaft of radius.

8869 — Distal end of radius.

6510— Pisiform.

598, 3794, 8605, 9127—Metacarpals.

Hindlimb:
20284 — Proximal end of femur.
105 —Calcaneum.

(2) Melkbos

Mb 143—Left mandibular fragment with P, and M,.
Mb 603—Right mandibular fragment with part of P,.
Mb 561—Distal end of radius.

Mb 576, Mb 659—Incomplete metapodials.

(3) Swartklip

ZW 100 —Incomplete braincase.

ZW t1o1 —Incomplete braincase.

ZW 1447—Fragment of facial region with right I* and I, and left I! to I3, part
of CG, and P*.

ZW 3789—Right maxillary fragment with P%.

ZW 131 —Left maxillary fragment with dp*, dp* and M? just erupting, and
unerupted C, P* and P*.

ZW 106 —Left maxillary fragment with dp? and dp’, and unerupted P%.

ZW 122 —Left and right dp’.

ZW 1872—Left dp,.

ZW 1457 and others— Fragments of unerupted permanent teeth.

ZW 3492—Part of the shaft of a humerus.

ZW 2800— UlIna lacking distal end and part of proximal end.

ZW 350, ZW 526—Proximal and distal ends of a radius.

ZW 1040, ZW 1789, ZW 3240—Metacarpals or parts thereof.

ZW 3439 — Distal end of a tibia.

ZW 2533 —Incomplete metatarsal.

ZW 237, ZW 1775, ZW 2796, ZW 3276—I1st phalanges.

(4) Sea Harvest, Saldanha

S 129—Right maxillary fragment with alveoli of C, P* and M?, and P® just
erupting.

S 208—Incomplete right P,.

S 237—Metacarpal.

S 230—Calcaneum.

S 130, S 234, S 715—Metatarsals.

S 241—2nd phalanx.

314 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

The study of this material was hampered by the lack of adequate compara-
tive material. Most of the cranial specimens belong to juveniles and only a
single skull of a young modern lion was available, although 27 adult skulls
were examined in the course of the study. Only seven complete skeletons were
available, of which six belonged to zoo specimens which suffered from rickets.
The seventh belonged to a large male from Botswana (SAM 3985).

Sea Harvest

The Sea Harvest maxillary fragment, S 129, belongs to an individual
which was slightly older than that represented by the only juvenile skull in the
comparative series (SAM 36663). The fossil dp? had apparently just been shed,
while that of the skull of SAM 36663 is still in position, but in the process of

é

ecocemarrnantaee

sccsnsarormmnrce I meneetne

_erconeonanaeoeoe ‘covonsccesecerrcer
ooveencenmmnoancanenena,

Sesncoescoae, srosmamannnicecen @

Lx > 1 pense ease

naccinnneneee

womimercecooeccox f eomemccnancann

: ———

veneers S 6 aan

sensssintannreercce nvnnnevovce

eed _nooseeemmnnesscsce

2 oe eee

ceveroetoroeneenaionse:

corenccncsecocennn beaecrsmccanecperes

cocscaussssscsione nanan

——-| (200
sponcecccosecneen

rerrnecresnsoonins | emcee

ELLE, encrmereoreme

iccsmmsacssiaas “rreewonenmocaate:

fovea sccnnonsopopecocs cceceeeeeeaeaaal

oo = eet

z saneronermocorene

Rosa Receeeeeeeeceaae
eenorernnecsecceoetce

pomemnens a es

nSSSBRSSSOSRT sapere

od coweesconssssennn

| canal sapepmeeenceciee igescommonennatet

‘RCCOSSSSSR NOLES SEEEESISET pecsscecacsneseremeraced

eae COREELETOO besecnatssree

seonscsesaocoeen.

es | fronts sonesesesce

sressemmmammenoe Soe wascanaenenrees:

seenotensoceeaiser

eee sescserrrrococe
ssasopnariwsosccoen

rorronccnsemcsscseeer

ye = paca

SCE oevoocceenncesessen |

-wetsatassonecctate iver Comme

eRRSe NC NISSEES ™m RERSOGSSNESE

sressenenrcoscccee

j nemenereeeesoses

forte SISLLLES OOS SSSR

“torensencscetessct:

td esescenenaaoene: aguas

renacoserean coors seseemaoseoeoneeas:

SSCS ess Ropeeneenscceness
seeseerermenesusesaieee

ee tA eS ERERRRERONO CS

Z eonere cots

cael

SenccceSRSSOSS soessccenneasroneenmonces {

eonoeteessscssn

~~ amass

ed SEIS,

Ce saaseaiaaanaaneal poet csaameaenaamaanenll

nomnercessisine g

os > —<—)
ssccammmeowsssocs

Cees EC

poo ceessccaassonInICe ee ae

mcencesssscornoon ih

_sessenmecoseoeene

ES a | eecassssesaL SEES,

cocecassoppites commana

EELS ETE AT favor cobeaanereemmonance
nsanesccecanoeee

sseascaasesososonaniinte ae

ossccesssoscees A iS a | ET

RRO BIEBESO amenmaaens

_aaescemesesoetee STEER,

pons Iececcsssonecesntt ensecasiasosonoeetisssn

oeeneencesecesete:

pemecnasssecote

ecasssaoseccatc

pect ROR

mn
mM

Fig. 72. Occlusal views of the Panthera leo maxilla (S 129) from Sea Harvest and a modern
specimen (SAM 36663).

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 315

being forced out by the erupting P*. The fossil is similar in size to corresponding
parts of the modern specimen, but they differ in that the modern one has a
fully erupted P?, whereas in the fossil this tooth is completely absent. The fossil
dp? is lost, its alveolus partly overlapping the erupting P*. The post-canine
diastema is very short. The P® is similar to that of modern Panthera leo in all
observable respects and, judging from the size of the P* and M! alveoli, these
teeth were comparable in size to those of the moderns.

Most of the Sea Harvest postcranial bones are similar in size to those of
the large male from Botswana. The exceptions are a metatarsal II (S 715) and
the 2nd phalanx (S 241), which are a little larger than the corresponding bones
of SAM 3985.

On the basis of the few specimens available, it appears that the Sea Harvest
lion was comparable in size to modern P. leo, but differed in lacking P?. Although
the Sea Harvest fossils are regarded as Holocene in age, the possibility that the
lion from this site is the so-called “Cape lion’ (P. leo melanochaita H. Smith, 1842)
was dismissed, since a well developed P? was present in this variety (vide infra).

Swartklip

Some of the Swartklip lion specimens have already been discussed (Hendey
& Hendey 1968), and although it was concluded that they do not differ from
the modern form, a more detailed examination has shown that this is not the
case.

The two incomplete braincases, ZW 100 and ZW 101, belong to adults
and are similar in overall size to corresponding parts of the skulls of modern
females (Table 82). However, the more complete specimen, ZW 100, differs
from the moderns in having a broader occiput dorsal to the occipital condyles.
The occipital width expressed as a percentage of the braincase length is greater
in the fossil than in both males and females of the comparative series. The post-
orbital region is also relatively broad.

The snout fragment, ZW 1447, and the maxillary fragment, ZW 3789,
evidently belong to mature adults, since both have worn teeth. ZW 1447
compares in size to corresponding parts of the skulls of modern males and is
remarkable only because the P? is smaller than those of the comparative series
(Table 84). This suggests the possibility that the Swartklip lion was ancestral
to that from Sea Harvest, a variety in which P? is lost. These local fossils are
regarded as belonging in a lineage distinct from other lions and one which is
characterized by the reduction and eventual loss of P?.

The P? of ZW 378g is within the size range of variation observed in modern
males, while the P? alveolus is similar in size to that of ZW 1447.

The juvenile maxillary fragment, ZW 131, is larger than those of the
Sea Harvest specimen, S 129, and the modern juvenile, SAM 36663, although
it belongs to an individual which was ontogenetically younger than both these
specimens. The dp? is still in position and the P* is only just starting to erupt.
The P’, P* and M?, although unerupted or only partly erupted, are larger than

=
2)
fa
nm
5
=
Z,
<
S
o%
fy
<
e
=
°
nm
ta
se
=
fy
O
w
4
<
Z
Z
<

316

GobSE WS P
1Z1SE JWS 6

6zSL WS 6

9 a | | | |

“SUOI] UJOPOUL Jo satias e JO ssoy} YIM poieduiod ‘097 vsayjuD AipyIIVMG IY} JO VszoUIvIq 94} JO SUOISUIWICT

0a] DLIYJUDT

(Dj1py20UDJaU 097 * 7)
vor odep

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 317

TABLE 83

Dimensions of the deciduous teeth of the Swartklip Panthera leo, compared with those of a young
modern lion

dp? dp? dp* dp,
] b ] b ] b ] b
ZW 131 — — | 26,8 15,5 | 8,0 14,7 — _
ZW 106 3,8 2,9 | 27,0 15,0 | — — _— —
Swartklip —§ |_—A |] tA _ PTO _ _—_
ZW 122a = = = — | 84 14,2 — —
ZW 1872 — _— _ _— =. — | ¢. 20,1 8,0
Modern
P. leo SAM 36663 | — — | 24,5 13,8 | — _— 47,7 6,9

the corresponding teeth in the comparative series, while the dp? is larger than
that of SAM 36663. The same applies in the case of the dp? of ZW 106 and the
dp, of ZW 1872 (Table 83). Although ZW 106 belongs to a younger individual
than S 129, it has a longer post-canine diastema and an unerupted P? in the
maxilla, a further indication of the contrasting nature of this region of the skull
of the Swartklip and Sea Harvest lions.

To sum up, the Swartklip lion is apparently characterized by a relatiyely
short braincase and broad occiput, a reduced P?, enlarged posterior cheekteeth,
with a corresponding enlargement of the posterior part of the maxilla.

The postcranial skeleton is little different in size to that of the large modern
male, SAM 3983, although the phalanges, a metacarpal V (ZW 1040) and the
ulna (ZW 2800) are larger. If the larger specimens belonged to males and the
others to females, it would indicate that the overall size of the Swartklip lion
was greater than that of modern P. leo. This species would then have been like
others from Swartklip in that it was larger than its modern counterpart.

Melkbos

The Melkbos specimens, with the exception of Mb 659, have already been
described (Hendey 1968), and they too differ from the corresponding parts of
modern P. /eo in their larger size. The additional specimen, a metacarpal V, is
also larger than that of the modern comparative specimens (Table 86).

Elandsfontein

The Elandsfontein skull, which will presumably be described in detail
elsewhere, is appreciably larger than the skulls of modern lions and it has
already been suggested that there might ke a close relationship between this
form and that from Melkbos (Hendey 1968).

ANNALS OF THE SOUTH AFRICAN MUSEUM

318

‘P96] IOUS 5
‘TS6] WyoYypunT ,
"IJO9A[e JO O9ZIS UO Paseq so}eUII}se IO ‘Y1990} JUOUeUIOd poIdnisUy ,

a ee

2
0a]
DaayjuDg
usapoW

6
02]
psayjUuDg
ulopo|W

JSOAILH
BOS

Ol Ol Ol Ol 8 8 Il a II 8 U
6:7S ES eSOG Nt ESI OG EI rec) Ol 18, Le ciel Sie ey MN Oe 76 66,01 8.0% 8 6%
Sele |) SOS SKS |) SA SS || SES SQ SAO Sree —ZOl SSN OTE Sl Gee =S:9) =6'8 -1°8I -9°€Z osury
ITS L'vl 68% | OMI 627 | ZOl 16! 6LI €°SZ 6 II ‘6 L‘8€ €€l 797 Si, WG P61 8°97 uLoy
SPs 91 67 Isl O67 | ZII'? S‘6l O81 2 9°87 9 V7 761 €‘6€ Or 2 097 9L $6 T0@ 0872 Scie
TLS GS ical Srl Ose le SO) O07 | T9T < = L‘0@ 0°8E cel 0°97 a re Isl s‘sz eCCrl
MWHN
9 9 9 9 ¢ 9 8 8 L 9 L U
9'0S Sel SLT] OFT O8T] +6 Gl | wSl Lie | i) 3 Gel Ole i (EO! WW rxe
Ops OrGleaOrSe lc=O Mlealcec, | aes SSS || SSS Sal -r‘0I -L‘SI -O'VE O(N <=S 0K =Gi9 "68 -6'PI —O'I7Z osuey
CSP GE Oe | WE ESE || 8B yLI C‘rl O17 Eat TLI 6°SE 61 6°€7 OL £6 SG Wee uvoy
L‘vs Sil Gere | wets Gite | OG 2 “Gy?! TSI (CGC, DY 8°6 PL 9'rE SIT 9°7Z €L OO 2] LOL E€‘bz IZISE
Wvs
ASS SC Oe Il. Sil Sie || GSO! 0°61 = a = O'8I 0°6€ Oe! 5‘97 = guys — 7o10q
-SABLINIA
= ae = = = — = — = = = TET GiSG a as pS SS 68LEMZ
— = == Se ee a = — = E912 | O72 OEP? = 6‘87 ‘2 — in oe IE] MZ I
$$ | | ee] IBAA
= =) i ee = = = — = = == == SIS SO SL Ce; Levi MZ
=f Sa Se = i = = = a Sil @ || EELS Gveye~e = 9‘p7 9 Sys = a ee 1671S
q I q I q I q I q q I q I q I q I
1/001X9 ee
A TW val *"d 2) IW rd sd ed @)

“SUOT] UITIJY UIIYINOS 1UIIII Jo 9soy} YIM poredurod ‘souTAOIg ode U1d}saM-YINOS 9Y} WOLF 097 V1aYyJUD [ISSO} JO Y}99} OY} JO sUOISUIUI

Vg alav

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 319

In general, the postcranial bones from this site are as large or larger than
those of the available modern P. leo specimens, although an appreciable size
range of variation was observed in those skeletal elements which are represented
by several specimens (i.e. humerus, ulna, radius). ‘The best represented of the
bones is the humerus and fifteen specimens belonging to at least thirteen indivi-
duals are known. The size difference between the smallest and the largest of
the humeri appears to be rather more than would be expected in a single
population, although, in view of the inadequate comparative series, this is a
somewhat subjective judgement. However, the size difference between the
humeri of the largest male and smallest female in the comparative series is
only about 16%, while the difference between the largest and smallest humeri
in the Elandsfontein series is about 33%, or twice as much. Consequently,
it is possible that more than one form of the lion is represented at Elandsfontein
as is the case with other species, although in this instance the nature of the
preservation of the specimens did not suggest this. In fact, the preservation
of one of the smallest specimens (16670) and the largest specimen (16795)
would suggest that they both belong with the Cornelian element in the
fauna.

The material may actually include some specimens which belong to a
large felid other than the lion and some of the smaller bones approach those of
the Langebaanweg ? Machairodus in size (Table 85). Large machairodonts
are known from the Makapanian, namely the Homotherium sp. from Makapansgat
(Collings 1972; vide supra), Machairodus transvaalensis from Bolt’s Workings
(Broom 1939) and ? Epimachairodus sp. (? = Homotherium sp.) from Kromdraai
(Ewer 1955c), and one of these, probably a Homotherium, may have survived
into the Cornelian. The absence of cranial remains of a machairodont larger
than Megantereon is not necessarily significant, since the Elandsfontein Megante-
reon itself is apparently represented only by the incomplete mandible of one
individual.

It is possibly significant that the relatively small humerus referred to earlier
(16670) exhibits pathological conditions similar to those observed in other
machairodont bones from the south-western Cape (vide supra). The pathology of
16670 includes osteoporosus in the region of the olecranon fossa, bony ‘lipping’
of the articular surface and eburnation. No similar pathology was observed
in any of the other specimens referred to P. leo.

The identification of a large machairodont in the Elandsfontein assemblage
can clearly not be substantiated on such slender evidence, but there is at least
a possibility that a large machairodont was contemporary with the smaller
Megantereon. This possibility is not reflected in the list of carnivores identified at
Elandsfontein (Table 89), although a provisional listing might have been
warranted.

In order to illustrate the degree to which the lion postcranial bones from
the south-western Cape differ in size, measurements of one typical example, the
metacarpal V, are given in Tabie 86.

=
2)
ce
Ze)
=)
=
Z
<
5
o4
By
<
se
sal
2)
fe)
2)
a
s0
sal
eal
fe)
NM
4
<
Z
Z
<

320

OI o°GL—P‘ES 6°z9g lf 1°19-V'1S Q‘LS L‘&¥ G‘CY -9 puo ‘jsIp 1e ‘wieIp -ysod--jue “xeyy
II 6°LL—-S‘1G b‘zg L G‘E9-6‘0S 6°9S QGP 9‘6P uOT}E[NIIe “jsIp JO ‘WILIP “suvI} “xe[
6 bb11-6°VL L‘v6 [L o‘V6—S‘6L L‘Lg ot 1°€L pUo ‘jsIP Je “UIeIP ‘suvI] “xe]]
u asury ue u Bue Y uv 6986 ‘JT 66£9 ‘TJ

SPIpoy odie] 0a) DLaYIUD_ SNPOLWYIDIN é

UId}UOJspuely UIIpOyY Samueeqosue'y]

“Snpomysvy BamMuLEqosuLy] dy} PUL 077 DLIYZUDg UIOPOUI JO asoy} YIM poreduioo ‘ulsjuOJspuL[Y Woy TIUINY pray VdIV] Jo suoIsusUIC]

Cg alavy,

321

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE

L‘13-1°gt Foz 9‘0% O'1% — —

G‘0z-2‘gI 6°61 G61 0'3G — O'1% ‘9

L‘Go-P'1% 6'Ez Gas bbz 6‘gz g‘Lz

6°Gs-S‘1z L‘S% obs o'Sz L‘gz 0‘gz

£°G6—§ ‘Ee b‘o6 a‘66 G‘o06 — O%ZOI 9
LEzg 6S99 W

osury uvoy|y ysoarvpy | 6941 MZ | OFOI MZ | soqyla
vIG

0a] vsayUDg diyyaemsg

UIIPO|

Qs puso ‘ystp ‘wrerp ‘d—e ‘xepy
BGS puo “}sIp “UIeIp ‘sued) *xey
6°gz pus *xoid ‘uivrp ‘d-'v -xepy
L‘gz puo ‘xoid ‘wrerp ‘sues *xepy
S‘96 Yyysudg] [[e419AO
g6S

uloyUO}

-spue|y

eee
‘susurtoads UJapoUur Jo satias v YIM paiedusoo ‘souTAOIg edeD U19}s9M-YINOs dy) Woy A [edivovyoUI 04) vLaYJUDY JO SUOISUDUIIP OY J,

9g a1av

322 ANNALS OF THE SOUTH AFRICAN MUSEUM

Discussion

Although all the fossil lion specimens from the south-western Cape are
here referred to Panthera leo, it is evident that none of the forms represented is
identical to the modern varieties of this species.

The Swartklip and Sea Harvest lions apparently belong in a lineage
distinct from that which led to the modern South African P. leo and the large
Elandsfontein form might also be a member of this southern lineage. This
lineage may have become extinct quite early in the Holocene since it is last
recorded in the Sea Harvest assemblage and the lion present in the area in
historic times was apparently P. leo melanochaita, a skull of which is recorded from
Betty’s Bay in the vicinity of Cape Hangklip (Meester 1971).

An indication of the taxonomic status which should be afforded to a
population of lions characterized by the reduction and eventual loss of a
cheektooth is evident from a well-documented example of a similar development
_ in modern P. leo. Todd (1966: 520), in his study of the only surviving population
of Asiatic lions, found that the ‘fusion of the roots of Pmg is . . . a common trait
in Gir lions . . . [and the] absence of this tooth appears for the first time in a
skull of c. 1910 and appears to have increased in incidence among skulls of
animals which died between approximately 1953-1963’. The craniological
differences between the modern African and Gir lions are probably no greater
than those between the Swartklip and Sea Harvest lions and their more northerly
contemporaries. In order that the present study be consistent in respect of the
taxonomic recognition given an apparently major difference such as the pre-
sence or absence of a cheektooth with the consequent recognition of a distinct
lineage, the south-western Cape lion should be referred to a new species.
The example of the Gir lions shows that such a step is not necessarily
warranted. Furthermore, since the local fossil carnivores have not been
given new subspecies names either, the lion material is all referred simply
to P. leo.

Lions of Makapanian age are known in South Africa from Sterkfontein
(P. shawi Broom, 1948), Swartkrans and Kromdraai (Ewer 1956a). All are
poorly represented. The Sterkfontein lion is very large, that from Swartkrans
is comparable in size to modern P. leo, while that from Kromdraai is larger.
These size differences, coupled with those evident in the south-western Cape
fossils, indicate that the lion, like Canis mesomelas and other species, underwent
size Changes through much, if not all of the Quaternary.

The Cape lion

Mention was made earlier of a number of species of ‘Cape’ mammals such
as the quagga, Cape warthog and blaauwbok, the distributions of which were
largely confined to the more southerly parts of the African continent and which
became extinct during the historic period. The Cape lion, Panthera leo melano-
chaita, is the only carnivore included in this category of mammals and, as such,
it is relevant to the present study.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 323

Hamilton Smith (1842) described the Cape lion as follows:

‘The species is of the largest size, with a bulldog head; the facial line is much
depressed between the eyes; large pointed ears edged with black; a great
mane of the same colour extending beyond the shoulders; a fringe of black
hair under the belly; a very stout tail, and the structure in general propor-
tions lower than in other lions.’

Since the Sea Harvest lion is supposedly Holocene in age, it seemed
possible that it could perhaps have been the ‘Cape lion’ which is represented
at the site. The known postcranial bones suggest that it was large and it thus
conforms to the first of the statements in the diagnosis. In addition, the loss of
P? and very short post-canine diastema may well have fore-shortened the snout
so as to give it ‘a bulldog head’. However, the Sea Harvest lion is definitely
different from other specimens which are accepted as ‘Cape lions’.

Mazak (1964) listed eight specimens preserved in various collections,
while Meester (1971) described an additional skull and reviewed the craniologi-
cal characters of the subspecies on the basis of the three recorded skulls, the
other two having been described by Lundholm (1952) and Hemmer (1966).
There are three skulls in the South African Museum’s collections which must
now be added to the list of specimens.

The first, SAM 35121, is the skull of a female which was removed from a
mounted specimen in 1896. Its skin was destroyed. This specimen and another,
a male, were acquired from the South African Literary and Scientific Institution
when the collections of this organization were taken over by the newly founded
South African Museum in 1855. They were apparently part of the collection
made by Sir Andrew Smith prior to 1837 and are likely to have come from the
Cape Province. Smith did collect specimens from north of the Orange River,
but apparently none of these remained in South Africa. The possibility that
these two specimens were Cape lions has long been recognized (South African
Museum 1955), but the surviving skull has not hitherto been described. There
13 no record of the fate of the companion specimen, although it may have been
destroyed before 1896.

This skull is largely intact and well preserved, although the left P? and
some incisors are lost. At least three of the missing incisors were shed during life
and this, taken in conjunction with tooth wear and suture closure, indicates
that the skull belonged to a fairly aged individual. The braincase has been
damaged by a bullet and there is an entry wound about 15 mm in diameter in
the basi-sphenoid and an exit wound twice that size in the left parietal. It
conforms quite well in size with the two previously recorded female Cape lion
skulls (Table 87).

The second specimen, SAM 7529, also belongs to a female, but it is poorly
preserved and lacks the mandible. It was found on the farm Bergplaats near
Rooihoogte in the Beaufort West Division of the Cape Province and was
presented to the South African Museum in 1903. It lacks the right zygomatic

ANNALS OF THE SOUTH AFRICAN MUSEUM

324

p96] IOUS ; “TL6] 193S997A +

fi L L L 9 9 L L L il L u
L‘0@ O'6L 6:89 O'SLZ T'8€ O'LPI €°L7 6‘ZOl €°06 O'8rE 0'€0r
-€‘8I -0'89 —‘r9 -0‘SEZ =G 65 -0°9€1 “SST -0'L6 -1°98 —0°LZE -0°69€ osuey | * P oa) vsayiuvg UIDPO-!
r6l PPL S‘b9 O1SZ 6°9E O'7rI €‘97 6001 788 €‘SEE 9‘E8¢ ureW
8°07 pPl L‘89 0°97 O‘8E O9EI “2 9°LZ 8°86 €‘16 O*LZE O*8SE “2 StPEE INVS
—_ || | ——_—__—— ee : ; * ££ uol edeg
191 O*LS 6°L9 01 bZ GLE O?ET (S1ly6 0°66 1‘€6 S‘O€E O‘SSE z0Chl MWHN
9 9 ¢ ¢ 9 9 9 9 9 9 9 u
(We ‘OL S‘bL O'%PZ S‘6€ €'871 7°87 7:26 €'76 0‘00€ O‘LZE
-0'8I prs -$‘%9 -0'88I =o [KS -‘TII -0°97 —p8L -¢‘98 -0°S9Z -0'667 oduey | * & 0a) véayjubg UIepPOW
L‘6l €‘19 9‘L9 ZOIZ €‘8E 6811 €‘LZ 8‘b8 €‘68 O‘LLZ EOE uvoy
9°07 ‘2 P°€9 8°89 “2 OCIT 9 €‘LE ‘9 OST 2 9°87 ‘2 0°88 ‘2 8°€6 2 0°687 2 O*80€ 9 6¢SL WYVS
5 ‘07 TLO 8°7L O77 p6£ 76Cl 8°87 bb6 0°68 0°%67 O‘8ZE IZIS€ WVS
_ | |S _ | | [| | |: * 6 uol odeg
€‘07 6°79 ¢‘v9 0007 L‘Or 7971 0°62 006 L‘06 0187 OOTE Avg S.Anog
6 61 019 8°S9 0°707 1‘6€ 0071 0‘O€ 0°76 8‘b6 0167 O‘LOE ;319qsABIIN|]

T9/001 qipim 19/001 UtpIM 19/001 YIP 19/001 UPI 19/001 yisus] yysua]
x MT [e31g10 X MZ |oewoshZ| x MW | Prose | xX MOM | ouUTUKO x Td) [eseqo, | 3so}BOID
-19}U] oseg -Apuog

“SUOT] ULSTIFY UIIYINOS I9YJO JO satias 9sOYy} YIM pareduiod (nvYyI0UDJaU 097 DayJUDG) SUOT] Ide JO s{[N¥s 9Y) JO suOIsUdUITC]

Lg alavL

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 325

arch and all the teeth, while the right maxillary, frontal and occipital regions
are damaged. It is also similar in size to the previously recorded female Cape
lion skulls.

The third skull, SAM 33425, is that of a male and was found in a cave
near Studtis in the Willowmore district of the southern Cape Folded Mountains
in 1948. It is remarkably well preserved and still has dried-out soft tissue adher-
ing to it. This is not unexpected since Studtis is in an arid area and dry condi-
tions in sheltered situations can result in the good preservation of animal remains
through desiccation. The skull is that of a fairly aged individual. Parts of the
nuchal crest and left mastoid process, the left G and right C are damaged, while
the left P? and some incisors were lost during life. It is similar in size to the
previously described male Cape lion skull and is larger than those of the females.

The last lions in the Cape Province were killed in the eastern Cape between
1842 and 1858 (Harper 1945). Sir Andrew Smith did his collecting in South
Africa between 1821 and 1836, so that SAM 35121 must have been shot during
this period. Both the Rooihoogte and Studtis specimens probably date from the
early part of the nineteenth century as well, although this is by no means certain.
Studtis is in a remote area in the southern Cape mountains and it is possible
that some lions survived there later than elsewhere in the Cape Province.
It is in these mountains that the last surviving population of Equus zebra zebra
is found, which is an indication that the area offers the potential for
late survivals.

Meester (1971: 28) examined the craniological characters which have
been used to distinguish the Cape lion from those occurring further north and
found that ‘in view of the lack of agreement in skull features evident among the
only three specimens known . . . the discovery of further material might still
further complicate the task of diagnosis, and perhaps even render it altogether
impossible’. ‘The new specimens do not yet confirm Meester’s fears and they
too can be distinguished from the northern varieties.

Meester showed that the muzzle width (base canine width as a percentage
of greatest length) is one of the more useful distinguishing characteristics of
Cape lion, skulls and this is substantiated by the new specimens (Table 87).
The observed differences between the Cape lions and others are, however,
very small, especially in view of the number of specimens involved. The least
difference observed is only 0,6 % for females and 0,3 °% for males.

The male skull also differs slightly in occipital length (condylobasal length
as a percentage of greatest length), with the least difference being 1%. The
Roothoogte female is also outside the range observed in the comparative series,
but SAM 35121 is not.

In the study of the lion crania from Swartklip, standard skull measurements
could not be taken owing to the incompleteness of the fossils. One of the ratios
calculated was the occipital width expressed as a percentage of braincase
length, and it was found that both the South African Museum’s female Cape
lion skulls differed from the modern females in this respect (Table 82). Four

326

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 73. Ventral, dorsal and lateral views of the female Panthera
melanochaita skull SAM 35121 (Scale approximately 300 mm).

leo

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 327

Fig. 74. Ventral, dorsal and lateral views of the male Panthera leo
melanochaita skull SAM 33425 (Scale approximately 300 mm).

328 ANNALS OF THE SOUTH AFRICAN MUSEUM

of the male skulls compared closely to both male and female Cape lions, but
curiously they were the skulls of zoo specimens, while the skulls of the wild
males differed from the Cape lions. This indicates that in the Cape lion the
braincase is shorter relative to occiput width. This is probably just another way
of illustrating the sometimes distinctive occipital length of the subspecies.
Hemmer (1966) emphasized the value of M, breadth in identifying Cape
lions and these teeth in the new specimens are indeed broader than those of the
comparative specimens (Table 84), the difference being especially marked in the
case of the females.
Although Meester may ultimately be proved right and the Cape lion may
be found to be not readily distinguishable from other varieties in respect of its
skull characters, on the basis of the specimens presently known, the following
characters may be said to be indicative of P. leo melanochaita:
(1) The muzzle is relatively broad.

(2) The occiput is relatively short and broad.
(3) The lower carnassials are relatively broad.

The Betty’s Bay specimen described by Meester (1971) suggests that it was
the Cape lion which inhabited the south-western Cape during the latter part
of the Holocene, although since this specimen is from near the boundary,
between the south-western and southern Cape regions, it is possible that the
Cape lion was never present elsewhere in the former region. Neither the bonte-
bok nor the blaauwbok are recorded from the south-western Cape late in the
Holocene, so it is possible that the lion encountered locally by early European
settlers was a descendant of the Sea Harvest variety and not the Cape lion at all.

The situation in respect of the characteristics of the local Panthera leo,
Felis serval, Hyaena brunnea and Canis mesomelas populations will only be resolved
as more specimens of these species are recovered. There are, however, some
suggestions that locally endemic varieties were represented in the south-western
Cape.

Suborder FISSIPEDIA

Unclassified specimens

As with the material of Pliocene age, there were a number of specimens in
the Quaternary fossil assemblages which were not identified, although they
clearly belong to fissiped carnivores. The unclassified specimens are mainly
postcranial bones and fragmentary teeth and none suggested the presence of a
species other than those already listed.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 329

GENERAL DISCUSSION ON THE CARNIVORA

A striking feature of the fossil carnivore assemblages from the south-western
Cape is the variety of species represented (Tables 88, 89). In no instance, how-
ever, is the number of species in any one assemblage in excess of the number
which might be expected of a fauna in an African context. There are definite
records of at least 25 carnivore species from the historic period of the south-
western Cape and the most recorded in a fossil assemblage are the 20 from the
Varswater Formation at Langebaanweg.

TABLE 88

The late Pliocene Carnivora of the Varswater Formation, Langebaanweg, Cape Province.

Marine Estuarine Estuarine Faunal
Faunal Faunal Faunal Unit
Unit 2 Unit 1 Unit 2 unknown
(Bed 1) (Bed 2) (Bed ga)
SUBORDER PINNIPEDIA
Prionodelphis capensis. ; x x x
SUBORDER FISSIPEDIA

Canidae incertae sedis . : : *

Agriotherium africanum . : : x

Mellivora aff. punjabiensis. x

Enhydriodon africanus. : ‘ 4

Viverra leakeyi x x

Genetta sp. . : x

Herpestes Species A x x

Herpestes Species B x x

Percrocuta australis x

Hyaena abronia * ?

Hyaena Species B . : x

Hyaenictis preforfex : : 4

Hyaenidae Species E . x

Machairodus sp. . : : : x

cf. Homotherium sp. x

Felis aff. issiodorensts x

Felis obscura : P ' %

Dinofelis diastemata “ : x x

Carnivora incertae sedis ; : x

All extant families of African Carnivora are represented locally, while the
Ursidae are an addition to the list. Most of the extant African genera are
recorded, the most notable exceptions being Acinonyx and a number of mustelids
and viverrids. Although the absence of such forms may be due to inadequacies
in the fossil and recent records, it is much more likely that they were never
present locally.

The pattern of similarities and differences between the modern carnivores
and the various fossil assemblages follows an entirely predictable pattern, with
the resemblance at different taxonomic levels decreasing with the increasing

330 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 89

The Quaternary Carnivora of the south-western Cape Province.

Baard’s
Quarry, Sea
Lange- | Elands- Harvest, | Historic

baanweg*| fontein | Melkbos | Swartklip|Saldanha| Period
SUBORDER PINNIPEDIA
{Hydrurga leptonyx . ;
Lobodon carcinophagus x
Mirounga leonina ;
Arctocephalus pusillus : ? x
tArctocephalus gazella
+ Arctocephalus tropicalis
SUBORDER FISSIPEDIA
?Canis sp. ; : : x
Canis mesomelas
Canis terblanchet
Vulpes chama
Lycaon pictus
+ Otocyon megalotis
Ictonyx striatus . : : x
Mellivora cf. capensis : x
Mellivora capensis. ; x
Aonyx capensis . :
Viverra civetta . ; : x
+ Genetta genetta
Genetta tigrina :
Herpestes ichneumon ‘ x x
Herpestes pulverulentus
Atilax paludinosus
~ tCynictis penicillata . ..
Suricata major . : : x
Hyaena cf. bellax . . x
Hyaena brunnea a ; x x x
Crocuta crocuta ; ‘ x
+ Proteles cristatus :
Megantereon sp. : : x
Felts libyca
Felis serval
Felis caracal_ : ; x
Panthera pardus
Panthera leo

wu XK K XK XK

OS 1K OX
x xX x xX
x x x x
~S 2S x X XK XK

x
x xX KX XK XK X

x xX
x 5°) x XX
®— KA KK XK DX DK 2K

x

x x

x

* Pleistocene species only. + Not known locally in a fossil state.

age of the assemblages (Table 90). About 50% of the genera recorded from the
late Pliocene are now extinct, but only one of the 23 recorded Quaternary
genera is extinct. The relative ages of the principal local fossil mammal occur-
rences were already established prior to the commencement of the present study
and the carnivores have served to substantiate the earlier conclusions, while
in the case of the Elandsfontein fauna they provided the first definite faunal
evidence indicating that the assemblage is not homogeneous.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 331

Table 90

Faunal resemblances* between the fossil and modern fissiped carnivore assemblages from the
south-western Cape Province.

Elandsfontein
Langebaanweg| (Cornelian) Swartklip Moderns
Number of species . : 19 12 10 20
Number of genera . : 14 II 9 15
Number of families... 6 5 5 5

Langebaanweg/ Elandsfontein/ Swartklip/

Moderns Moderns Moderns
Species in common . : ; 0) 8 10
Faunal resemblance (%) . 5 : fe) 66 100
Genera in common . : : : 5 9 9
Faunal resemblance (%) . ; , 36 82 100
Families in common . : . : 5 5 5
Faunal resemblance (%) . ; : 100 100 100

Minimum number of genera and species are counted.
* Simpson 1967.

Before the significance of any element in a fossil mammal fauna can be
assessed, it is necessary that the identity and phyletic relationships of the indivi-
dual species be determined. In the case of most of the south-western Cape
carnivores this basic requirement was established and their relationships are
clear. In summing up the present state of knowledge of the local carnivores, it is
convenient to consider the pinnipeds and each of the fissiped families
individually.

PINNIPEDIA

The south-western Cape fossil record of the Pinnipedia is very incomplete
and only the Langebaanian Prionodelphis capensis and the Holocene Arctocephalus
pusillus are comparatively well represented. The local late Cenozoic history of
this group has apparently centred on the replacement of the Phocidae by the
Otariidae (Hendey 1972a). The former are now represented only by occasional
stray individuals of Antarctic and sub-Antarctic species, but the Otariidae are
represented by large permanent colonies of Arctocephalus pusillus, while indivi-
duals of sub-Antarctic species probably also stray onto the local coastline.

A. pusillus is known from a number of Holocene fossil occurrences, mainly
Late Stone Age coastal middens, while there is at least one record, that

332 ANNALS OF THE SOUTH AFRICAN MUSEUM

from the Saldanha Lime Quarry, which may date back to the Pleistocene.
There is a suggestion that species such as Mirounga leonina and Lobodon carcino-
phagus, which are today largely confined to higher latitudes, were more com-
monly represented earlier in the Holocene, and perhaps also the Pleistocene.
The local presence of these species in greater numbers than at present may be
related to the colder periods of the past.

The only phocid, and in fact the only pinniped other than Arctocephalus
which still has permanent colonies on the coast of Africa is the Mediterranean
monk seal, Monachus monachus, whose range also extends on to the west coast of
North Africa. This is a declining species and may be a relict population
descended from the generalized monachine which was probably ancestral to
all other members of the subfamily (Hendey 1972a). The Langebaanweg
Prionodelphis is in certain respects intermediate in character between Monachus
and the other extant monachines Leptonychotes, Ommatophoca, Hydrurga, Lobodon
and Murounga (Hendey & Repenning 1972). Perhaps significantly the South
African and Argentinian records of Prionodelphis are intermediate in age between
the ancestral monachine of the European Miocene and the extant species,
and they are intermediately situated between the surviving populations of
Monachus in low latitudes and the main area of modern monachine distribution
in southern high latitudes.

The successful radiation of the otariids at the expense of the phocids in
southern mid-latitudes is one of the more remarkable aspects of pinniped
evolution, since it was achieved in a relatively short space of time and it involved
the decline of an extremely successful family, being the one which completely
dominates the pinniped populations of the Northern Hemisphere and the high
latitudes of the Southern Hemisphere.

CANIDAE

Although the Canidae are a morphologically conservative group of carni-
vores, they have been very successful and the only continent on which they have
not become established in a natural state is Antarctica. In Africa today the
most commonly occurring canids are the jackals (Canis spp.), while foxes
(Vulpes spp., Fennecus sp., Otocyon sp.) and a hunting dog (Lycaon sp.) are, or
were, also widespread. Fennecus and several Canis and Vulpes species are north or
north-east African forms belonging in the Palaearctic region or only marginally
in the Ethiopian Region.

Canis mesomelas is an essentially southern African jackal whose range
extends also to East Africa, while C. adustus is a tropical species which only
occurs in the more northerly parts of southern Africa. The Asiatic jackal,
C. aureus, is found in Africa only in the north and north-east. These three species
may have had their origins in South Africa, East Africa and Asia respectively.

C. mesomelas is an extremely successful species and its earliest records date
back to the Makapanian. It manages to survive today even in areas where
persistent efforts are made to bring about its eradication. In this respect it is

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 333

similar to the North American coyote (C. latrans), a species which has actually
extended its range in recent years (Cahalane 1961).

The pattern of distribution of the true foxes (Vulpes spp.) is essentially
similar to that of the jackals. V. vulpes is a Palaearctic species with a limited
distribution in North Africa, V. pallida and V. rueppelli are distributed in the
more arid parts of tropical North Africa, while V. chama is a southern African
species. The foxes have more restricted ranges than the jackals and there is no
tropical equivalent of C. adustus. The absence of Vulpes from much of tropical
Africa may be due to its ecological niche being occupied by C. adustus. The
recorded distributions of Vulpes and C. adustus suggests that a mutually exclusive
inter-relationship exists between them (see Dorst & Dandelot 1970).

The highly specialized, insectivorous Otocyon megalotis has a discontinuous
distribution in sub-Saharan Africa, with one range centred in East Africa and
the other in the arid south-western parts of the continent. It is one of the species
whose distribution suggests the previous existence of a south-west to north-east
arid corridor in Africa (vide supra).

Lycaon pictus was once widely distributed in Africa, but in common with
other large predators its range has been considerably reduced as a result of
human activities.

Three of the five canid genera represented in Africa are endemic to the
continent (Fennecus, Otocyon, Lycaon), although Lycaon is also recorded in the
Pleistocene of Europe (Kurtén 1968). Eleven canid species are still extant, of
which only two (Canis aureus, Vulpes vulpes) are also recorded in Eurasia.

In general, the fossil record of the Canidae is good and this is essentially
true of the south-western Cape late Cenozoic, although one of the curious
features of the Langebaanweg fauna is the fact that this family is so poorly
represented. The poor representation in a Pliocene fauna of a highly successful
mammalian family which had its origins in the Eocene is obviously of some
significance and a possible explanation is that the Langebaanweg Viverra
leakeyt filled the ecological niche which was later to be occupied by the jackals
in this and other areas. V. leakeyi was similar in size to the modern jackals and
its dentition resembles that of Canis. In contrast to the Langebaanweg record,
medium-sized canids are particularly well represented in the Quaternary of the
south-western Cape, while Viverra is either extremely rare, as at Elandsfontein,
or not represented at all.

Canis and Vulpes apparently had their origins early in the Pliocene of
Eurasia and North America respectively, while Viverra was already established
in Europe in the Miocene (Romer 1966). Consequently, it would not be sur-
prising to find that Viverra was already established in Africa before either of the
canid genera and that the latter only reached the most southerly parts of the
continent late in the Pliocene.

The modern African civet, Viverra civetta, is apparently not an active
competitor with canids of comparable size. It is an omnivorous species with a
dentition modified accordingly, so that in this respect it is very different from

334 ANNALS OF THE SOUTH AFRICAN MUSEUM

Canis and Vulpes, as well as from the Pliocene V. leakeyi. The survival of Viverra
in Africa is therefore ascribed to the fact that it underwent a change in habits
during the Pleistocene, so removing itself from competition with the canids.

It is worth noting that in southern Asia there is a civet with a dentition
similar to that of V. leakeyi, namely, V. zibetha, which overlaps in its range with
the jackal, C. aureus. In this instance the suggested competition between the
civets and jackals must have been resolved in some other way, and it probably
illustrates the point that challenges and responses in mammalian evolution
are far more complex than would appear from the preceding speculations.
Nevertheless, the hypothesis presented here to account for the nature of the
local record of Canis, Vulpes and Viverra could offer at least the basis of an
explanation.

The generic identity of the Langebaanweg canid is not known, but it is
perhaps more likely to be Canis than Vulpes, partly because of its size and partly
because Vulpes is much less commonly represented in the local Quaternary
record. Both genera are recorded during the Makapanian, so both may also
have been present in South Africa during the Langebaanian.

Lycaon is a genus which is apparently confined to the Quaternary. The
absence of even an ancestral form at Langebaanweg may be the result of a
faulty record, since these animals are rare as fossils. On the other hand, the
suggestion was made earlier that the Langebaanweg Hyaenidae included
cursorial and more actively predaceous forms (Hyaena Species B, Hyaenictis
preforfex) and these species may have occupied the niche later taken over by
Lycaon.

The Pleistocene canids of the south-western Cape are all representatives
of genera recorded in the area in recent times, namely, Canis, Vulpes and Lycaon.
The Elandsfontein C. terblanchei is the only extinct species recorded, but it may
in fact simply be a primitive form of C. adustus. There is nothing, such as the
Makapanian C. brevirostris, which is certainly without an extant counterpart.
With the exception of the Elandsfontein Vulpes chama, none of the local fossil
canids is obviously and strikingly different from their extant counterparts,
although all the Pleistocene varieties do differ in some respects from those which
are still extant.

The three most commonly represented local species, V. chama, C. mesomelas
and L. pictus, form a complementary group in respect of size and habits. V.
chama is the smallest and the modern form feeds on small mammals, birds and
insects. C. mesomelas is a medium-sized canid and is capable of preying on small
antelope and, in recent times, also sheep. Apart from its predatory habits, it
will also scavenge the kills of other carnivores and its less discriminating diet
may have contributed to its success as a species. The large Lycaon hunts in packs
and preys on animals as large as the wildebeest and zebra. Although each of
these species underwent size changes during the Quaternary, the relative sizes
of the local fossil varieties remained the same.

The south-western Cape fossil canids shed little light on the evolution of

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 335

the genera represented. Significant developments within this family took place
largely during the Tertiary and the local record only reveals the later stages in
the evolution of extant species.

URSIDAE

Prior to the discovery of Agriotherium at Langebaanweg, the Ursidae had
not been recorded in sub-Saharan Africa, the only other members of this
family on the continent being representatives of the Ursus arctos group, which
are recorded from Pleistocene and Holocene contexts in North Africa (see
Erdbrink 1953). The U. arctos group are, in general, north-temperate forms
and they entered North Africa during the Pleistocene when conditions were
more temperate than they are today, and when there existed a land link across
what are today the Straits of Gibraltar in addition to that in the Suez region.
Ursids were not the only Eurasian species to become established in North
Africa and this region is part of the Palaearctic Region, with limited faunal
affinities with sub-Saharan Africa. In spite of the fact that the Ursinae have
proved a remarkably adaptable group, they were apparently unable to pene-
trate the arid barrier of the Sahara Desert.

The presence of Agriotherium in sub-Saharan Africa is not completely
unexpected, since during the late Tertiary when the Agriotheriinae were
radiating, there was far more faunal interchange between Africa and Eurasia
than was the case subsequently. Agriotherium apparently had its origins in
Eurasia in the mid Pliocene and it must have entered Africa at about this time.
Since it is recorded from the late Pliocene of the extreme south of the continent,
it must initially have been a widespread and successful immigrant.

It was suggested elsewhere that the principal cause of the extinction or
limitation of the Agriotheriinae was their unsuccessful competition with the
radiating Ursinae (Hendey 19724). However, since Agriotherium was apparently
a carnivorous rather than omnivorous ursid, its extinction cannot be accounted
for in this way. A. africanum was a very large and apparently carnivorous species
and if it was indeed actively predaceous, its prey is likely to have included the
larger contemporary herbivores such as the giraffids, so it cannot have been the
lack of suitable prey which led to the extinction of Agriotherium on this continent.
Bourliére (1963: 51) has stated that, “There is no doubt that the African savannas
represent an ideal type of habitat for any large-sized carnivore or omnivore with mainly
carnworous habits’. This conclusion was based on studies of present mammal
populations, but there is no reason to suppose that it would not have applied in
the late Tertiary and Pleistocene as well. a

While the extinction of a species may be caused by factors other than
competition (e.g. climatic changes), it may simply be that the large and proba-
bly clumsy Agriothertum was unable to compete successfully with other contem-
porary carnivores such as ‘lions’ (Panthera spp.), wnich must have made their
appearance in South Africa at about the time that Agriotherium became extinct.

Alternatively, Agriotherium may have been a scavenger which competed

336 ANNALS OF THE SOUTH AFRICAN MUSEUM

unsuccessfully with the radiating hyaenids of the late Pliocene, there having
been more larger hyaenid species during this period in time than there were
earlier in the Pliocene.

It is possible that in spite of the nature of its dentition, Agriotherium was
omnivorous or herbivorous like most other ursids. If this was the case, then its
extinction in other parts of the world could be ascribed to the rise of the Ursinae,
but this would not apply in the case of sub-Saharan Africa. However, its extinc-
tion could have been caused by unsuccessful competition with an entirely
unrelated mammalian group occupying the same ecological niche. In this
instance the most reasonable possibility would have been the terrestrial Cercopi-
thecoidea and/or Hominoidea, which were undergoing a radiation in Africa
during the late Tertiary.

Since Agriothertum was successfully established in Africa during the Pliocene,
it might be expected that the same would apply to the agriotheriine Indarctos,
the genus from which Agriotherium was apparently derived and one which was
also widespread during the Pliocene. Judging from its dentition, Indarctos was
more like the Ursinae in its habits and the fact that it has not been recorded
in Africa is not necessarily significant in view of the comparatively poor Pliocene
record on this continent. If it was indeed present in Africa, then its extinction
locally could also be ascribed to unsuccessful competition with the terrestrial
primates.

MUSTELIDAE

The Mustelidae are an essentially Holarctic and Neotropical family and
are not well represented in the Ethiopian Region. The extant genera recorded
in southern Africa are Ictonyx, Poecilogale, Mellivora, Aonyx and Lutra and each is
represented by only a single species. This family is also poorly represented in
the African fossil record and the mustelids have apparently never been a
significant element in the fauna of sub-Saharan Africa. The south-western
Cape fossil record is not exceptional in respect of this family, but three of the
extant genera, Ictonyx, Mellivora and Aonyx, are represented locally. In addition,
there is at least one extinct genus recorded at Langebaanweg, namely, Enhy-
driodon, while there might also be a second, as yet unnamed giant otter repre-
sented at the site. Only Mellivora is comparatively common as a fossil and
almost ail the material is referred to the extant species, M. capensis. An extinct
species which might be ancestral to M. capensis is known from Langebaanweg.

The ‘badgers’ (Mellivorinae and Melinae) are an unusual group of
animals which have no parallel in other carnivore families, and the two sub-
families are themselves a remarkable example of parallel evolution. Judging
from the fossil record, the Mellivorinae have been firmly established in Africa
and southern Asia since the late Tertiary. They are not as diverse a group as
the Holarctic Melinae and their evolution was apparently largely a matter of a
change along one principal lineage from purely carnivorous forms to the more
omnivorous M. capensis. The changes which were undergone are relatively

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 337

minor and Mellivora has apparently remained virtually unchanged for much
of the Quaternary.

The long-standing and widespread success of the mustelids as aquatic
carnivores is one of the more notable aspects of this family. The Lutrinae are
a more diverse subfamily than the Mellivorinae, but they are, nevertheless, by
no means as diverse as some other carnivore groups. The Langebaanweg
Enhydriodon has Aonyx as its local counterpart in the Quaternary, although the
former was a much larger form. The other species from Langebaanweg which
was tentatively identified as an otter is also a giant form and it has no counter-
part recorded from the Quaternary.

There are two African viverrids, Osbornictis piscivora and Atilax paludinosus,
which are adapted to life in or near water, but neither differs from other viver-
rids to the extent that the otters are differentiated from other mustelids. Osborn-
ictis has only a very limited distribution in Central Africa, and while Aizlax is
more widespread it is not quite so tied to aquatic environments. The indications
are that the mustelids adapted to aquatic habitats long before these two viver-
rids and the latter have so far been unable to establish themselves in this ecologi-
cal niche as successfully as the otters.

While the success in Africa of the Mellivorinae and Lutrinae is readily
explained by the nature of their specialized habits, this is not the case with the
smaller and more generalized mustelids such as Jctonyx, which co-exists with
the highly successful viverrids of comparable size and with comparable habits.
With the Elandsfontein record of Jctonyx, it is now known that this genus has
been a part of the African fauna at least since the Cornelian. Its local history is,
however, probably far longer and, just as Mellivora has paralleled the evolution
of the Melinae since the late Tertiary, so Jctonyx has paralleled the New World
Mephitinae.

The co-existence of Jctonyx striatus and Herpestes pulverulentus in the south-
western Cape may simply be explained by the fact that the former is essentially
nocturnal, while the latter is diurnal. The same explanation may apply in other
parts of Africa where Jctonyx is found in association with other herpestine species.

VIVERRIDAE

The Viverridae are the most diverse of the African carnivores and they
are the dominant small carnivores of the Ethiopian and Oriental Regions,
with about 80 extant species having been recorded (Morris 1965). Six species
have been recorded in the south-western Cape in recent times, namely, Genetta
genetta, G. tigrina, Herpestes ichneumon, H. pulverulentus, Atilax paludinosus and
Cynictis penicillata. Only H. pulverulentus still survives in appreciable numbers.
In addition to these smaller species, some of which are known locally as fossils,
large civets have been recorded from Langebaanweg and Elandsfontein,
although they are not known from a post-Cornelian context.

A feature of the South African Viverridae is that while the smaller Herpesti-
nae apparently underwent a radiation during the Pleistocene, the larger

338 ANNALS OF THE SOUTH AFRICAN MUSEUM

Viverrinae went into a decline. The latter development has already been
accounted for by the suggestion that the civets gave way when they came into
competition with canids of comparable size and that Viverra civetta survived by
becoming progressively more omnivorous. Since this species now manages to
survive elsewhere in Africa in the same areas as foxes and jackals, the apparent
decline in its range in South Africa during the Pleistocene is still not accounted
for, although mention was made earlier of possible environmental controls.
Carnivores are not usually as influenced by environmental changes as herbivores,
but since berries and fruits form part of the diet of V. civetta (Dorst & Dandelot
1970), it is possible that vegetation changes in parts of South Africa reduced
the viability of the civet in these regions. In addition, there is little, if any,
overlapping in the ranges of V. civetta and Vulpes chama and since these
species probably do compete for certain foods such as insects and small verte-
brates, they may well have a mutually exclusive relationship.

The apparently rather sudden radiation of the Herpestinae is also not
readily accounted for, although it may be that the habitats presently occupied
by the various species were simply unexploited in the late Tertiary. For example,
the Makapanian form of Atzlax, described by Ewer (1956c) as Herpestes mesotes,
can only at that time have begun to exploit an environment (i.e. near water)
which had obviously existed for far longer than the history of the family.

The local fossil record of the smaller Viverridae is not good, but apparently
only Genetta and Herpestes were present in the south-western Cape during the
Langebaanian. The first record of Suricata anywhere is that from the Cornelian
element of the Elandsfontein fauna, so this is another genus which was most
probably differentiated during the Pleistocene. The earliest record of Cynictzs
is that from Makapansgat (Ewer 1956d), so that this genus, and perhaps also
Paracynictis, may well have had their beginnings in the Makapanian. The
earliest record of Mungos is from the early Pleistocene of East Africa (Petter
1969), while Crossarchus is known from the Transvaal Makapanian (Broom
19376). The other southern African herpestine genera (Bdeogale, Helogale,
Rhynchogale, Ichneumia) have no local fossil record, but they too are likely to
have originated during the Pleistocene (Petter 1969).

The north-temperate Mustelidae, which are ecological vicars of the
Viverridae, apparently underwent a similar and comparatively recent radiation.
For example, Anderson (1970) concludes that five of the seven extant species of
Martes were derived from the ‘early middle Pleistocene’ M. vetus. The spread of
Martes during the Pleistocene, and therefore also its taxonomic differentiation, was
largely determined by the climatic changes of this epoch. Although such changes
were not as extreme in Africa, they did influence the environment and may have
made some contribution towards the diversification of the smaller viverrids as well.

HYAENIDAE

The diversification of the Hyaenidae during the late Tertiary and early
Pleistocene and their subsequent decline is one of the more striking aspects of

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 339

the Old World fossil carnivore record. As many as six hyaenid species are now
definitely or tentatively recorded from the South African Langebaanian, while
seven species are known from the Makapanian (Ewer 1967). This is in marked
contrast to the two species which are still extant in South Africa today.

Ewer (1967: 120) has discussed this phenomenon and concluded that the
rise and decline of the hyaenids was determined by its relationships with the
machairodonts, since it was with these ‘primary’ predators that the ‘niche for
bone-crushing specialists (stood) invitingly wide open’ and ‘hyaenas and sabre-
tooths . . . form an ideal... combination’ with ‘the existence of the latter ...a
precondition for the evolution of the former’.

At Langebaanweg the hyaenids were associated with two large machairo-
donts and the ‘false sabre-tooth’, Dinofelis. By the Makapanian the smaller
machairodont, Megantereon, was also present, but so too were large felines
(Panthera spp.) so that the Makapanian is the transitional age in respect of local
hyaenid evolution. By the Cornelian the sabre-toothed cats are recorded for
the last time and, judging from the Elandsfontein record, large felines (Panthera
leo) became the dominant predators. The final disappearance of the machairo-
donts during the Cornelian ‘corresponds with the shrinking of the hyaenid fauna
to those few species which have succeeded in surviving today, in association
with modern Felinae’ (Ewer 1967: 121).

The South African Hyaenidae are, therefore, a group which began their
decline during the Makapanian and by the Cornelian had stabilized at the two
species which are still extant, namely, Hyaena brunnea and Crocuta crocuta.

The pattern of local hyaenid evolution is now so well documented that
they have become important from the point of view of the relative dating of
late Cenozoic faunas and deposits. Kurtén (1957a) indicated the possible
usefulness of Crocuta for the purposes of dating South African and other fossil
faunas and since then this, and other hyaenid genera have been similarly
employed (see Ewer 1967). Relative dating using the hyaenids has not been
without its problems and there are still crucial issues, such as the absolute dating
of at least some hyaenid occurrences, which must be settled. It is in this respect
that the East African hyaenid record is so important and if the situation is already
promising, then it will become even more so as the East African material is studied.

The distribution of modern hyaenids bears some similarity to that of other
carnivore families. One species, C. crocuta, is, or was widely distributed through-
out sub-Saharan Africa and in this respect is similar to other large carnivores
such as Panthera leo and Lycaon pictus. By contrast, H. brunnea is a southern African
species, its distribution early in the historic period probably having been much
like that of Vulpes chama. H. hyaena is an East and North African species whose
range extends into Asia and is thus similar to that of Canis aureus.

FELIDAE

The modern fauna of the south-western Cape included five felid species
ranging in size from the wildcat (Felis libyca) to the lion (Panthera leo). The felids

340 ANNALS OF THE SOUTH AFRICAN MUSEUM

are thus reminiscent of the canids, which were also represented by a series of
species of differing sizes. wo felids which are recorded elsewhere in South
Africa, the black-footed cat (F. nigripes) and cheetah (Acinonyx jubatus), were
apparently never present in the south-western Cape. All the locally occurring
modern species are recorded from Cornelian and/or Florisian contexts in the
south-western Cape and it is likely that the whole series were represented from
the Cornelian to early in the historic period.

Although there is no local record of Makapanian felids, inferences may be
drawn from the local Langebaanian and Transvaal Makapanian records.
During the Langebaanian the large felids were machairodonts and a false
sabre-toothed cat, while smaller lynx-like felines were also present (Felis spp.).
The representation of machairodonts continued throughout the Makapanian,
but large felines (Panthera spp.) also appeared. The lynx-sized felines no doubt
occurred during the Makapanian as well. It is by no means certain from the
fossil record when the wildcat first appeared in the south-western Cape, but
although it is first recorded from the Florisian, it is likely to have been present
for far longer. The final change from the archaic to modern felid faunas was
completed during the Cornelian and has already been referred to in connection
with the Hyaenidae.

The felids are perhaps the most highly specialized and actively predaceous
of the fissiped carnivores and the modern forms may be conveniently categorized
according to their size and locomotory adaptations. The first category includes
the smallest forms, of which F. libyca is the only local representative. The second
includes F. caracal and F. serval and they may perhaps be distinguished by the
former being an ambulatory and the latter a more cursorial form. This distinc-
tion is, however, not clear and both may be equally adaptable in as far as
locomotion is concerned. A more definite distinction along the same lines is
made in the third category, which includes P. pardus and A. jubatus. The former
is an ambulatory form which is also well adapted to arboreal locomotion, while
A. jubatus is a highly specialized cursorial form. The final category includes the
largest of the extant felids and P. /eo is the African representative. The felids are,
therefore, so represented that the constituent species can prey on animals
from very small to very large. In those categories in which there are two similar
sized species, these may have distinct locomotory adaptations which enable
them, if necessary, to favour different sorts of prey so that they need not be in
active competition with one another.

Apart from their obvious dental specializations, the machairodonts also
included locomotory adaptations not found in the felines mentioned above. The
last definitely recorded South African machairodont, Megantereon, which had
relatively short and heavy limbs, was evidently an ambulatory form quite
distinct from the partly arboreal P. pardus and cursorial A. jubatus in respect of
locomotion. While this should, theoretically, have enabled it to refrain from
active competition with contemporary felids of similar size, it is obviously a
disadvantage for a highly predaceous carnivore to be a relatively slow and

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 341

cumbersome animal and this may have contributed to its ultimate extinction.

In an African context, the Suidae are perhaps the herbivores most likely
to have been the preferred prey of a carnivore such as Megantereon. It may
therefore be significant that the local decline of the machairodonts went together
with the decline of the Suidae, which had undergone a relatively brief, but never-
theless spectacular radiation in Africa during the Makapanian and Cornelian.

These, and earlier speculations on the inter-relationships and habits of
South African late Cenozoic carnivores give an insight into the enormously
complex arrangement of the eco-systems of which these animals were a part.
While any single statement on this topic would obviously oversimplify the
realities of the situation, there are grounds for believing that the decline of the
African Suidae in the face of competition with the Bovidae, contributed to the
decline of the Machairodontinae, which in turn led to a decline in the Hyaeni-
dae. At the same time the rise in the Bovidae favoured the success of their most
common predators, the Felinae. Any one development in mammalian evolution
is likely to have wide repercussions and there is ample scope for more detailed
studies on the ecology of the local late Cenozoic Carnivora.

Although some aspects of the taxonomy, phylogeny and zoogeography of
the south-western Cape Carnivora have been summarized, there are still two
topics relating to this order which may be enlarged upon.

AGE AND RELATIVE DATING

Repeated references have been made to the age of the carnivore species
dealt with in this report, and to the application in the relative dating of faunas
and deposits. The use of the fossil carnivores in dating is directly dependent
upon the extent to which their phyletic relationships are understood. It follows
that some species are more useful than others in this respect, but no single
species can yet be used as an entirely reliable age indicator on its own and the
need to view a fauna as whole is an obvious necessity (see Ewer 19570).

The accompanying illustrations (Fig. 75) summarize the recorded time
spans of some of the species which have been described or referred to in this
report. The carnivores are actually more useful for relative dating purposes than
these text figures would suggest, because even though a single species may span
most, or all of the Pleistocene ages, temporal variants are recognized. For exam-
ple, comparisons between undated Canis mesomelas remains with those of the
recorded Makapanian, Cornelian, Florisian and Holocene varieties could result
in its identification with one of the forms and its age may be determined in this
way. Clearly, the more material that is described, the more useful the age
categorizations will become.

SIZE CHANGES

In most of the descriptions of the Quaternary fossil species mention was
made of the size differences between them and their modern counterparts.
Similar observations have previously been made in reference to a variety of

342 ANNALS OF THE SOUTH AFRICAN MUSEUM

LANGEBAANIAN | MAKAPANIAN | CORNELIAN | FLORISIAN | HOLOCENE

ES Ee Ef

Canis mesomelas
C. terblanchei ?
? C. adustus
V. pulcher
Vulpes chama
Lycaon pictus
Agriotherium
Ictonyx Striatus
M. aff punjabiensis
Mellivora capensis
Enhydriodon
Aonyx capensis
V. leakeyi
Viverra civetta

Fig. 75a. The recorded time spans of some South African Canidae, Ursidae, Mustelidae and
Viverridae.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 343

LANGEBAANIAN | MAKAPANIAN | CORNELIAN

FLORISIAN |HOLOCENE

H. abronia

gs ed

yenere rete ar
a < 24 Percrocuta

- z Crocuta
e “A i Machairodus

eee

‘ % ¥ Homotherium a 7 sal
a : ks Dinofelis

ite Panthera

ra Panthera

Fig 75b. The recorded time spans of some South African Hyaenidae and Felidae.

344. ANNALS OF THE SOUTH AFRICAN MUSEUM

vertebrate species from many parts of the world.

More than twenty years ago, Hooijer (1950: 360) wrote: ‘In current
literature on Quaternary faunas, both in the Old and in the New World,
we find quite a few scattered notes as to the large average size of a given fossil
or subfossil relative to recent material of the same species used for comparison.
It is, however, only during the last few years that statements as to this Quater-
nary size decrease apparently representing a general evolutionary trend appear
in the literature. It is considered as probable by some authorities that this
decrease in general size is to be correlated with the warming-up of the world’s
climates since the Ice Age, which would be the working of Bergmann’s Principle
in a temporal rather than a geographic way.’ He continued, noting that,
‘there can be no doubt that the fossil or subfossil animals are con-specific
with the recent, and that we are consequently dealing with subspecific advances
only. No extinction nor even migration is involved; it is evolution in sttu. The
means and modes of the various metrical characters are shifting in the course
of time: we have temporal clines, chronoclines.’

More recently, Kurtén (1968: chapter 17) has discussed the question of
Quaternary size changes and shown that they were not necessarily synchronous
in carnivore species from the same parts of the world and that species in different
parts of the world were not necessarily of uniform size at any given time. The
latter is perhaps fairly obvious in view of the size differences between different
populations of species which are still extant.

The reasons that species have undergone size changes in the course of their
evolution are almost certainly complex and varied, but in the higher latitudes
at least climatic fluctuations apparently played a significant role. Since Quat-
ernary climatic changes in South Africa were probably not as extreme as those
in the temperate latitudes of Europe, it is less likely that body size in relation
to heat regulation (Kurtén, 1968: 245) was a significant mechanism influencing
size changes. However, local environmental changes resulting from climatic
fluctuations may have had some effect on the average size of individuals in the
various species which occurred in the most southerly parts of Africa.

Apart from the problem of the causes of size changes, there are also diffi-
culties in analysing the actual nature of the changes. For example, in South
Africa there is an almost complete lack of chronometrically dated) fossil faunas,
so that even though their relative ages may be established, the absolute time
differences between them are not. Consequently, rates of size change cannot
be accurately determined. A second problem relates to the fact that size changes
in different skeletal elements of a single species were not necessarily constant.
This difficulty is clearly illustrated by the fossil Vulpes chama from the south-
western Cape. The limb bones of the Cornelian variety were only a little longer
than those of modern V. chama, but they were much stouter, while the skull was
much larger. Some of the teeth of the two varieties were little different in size,
but others, notably the posterior cheekteeth, were very much bigger in the
Cornelian variety.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 345

The species dealt with in this report are represented by a variety of skeletal
remains which are not always directly comparable. In order that some com-
parison of the relative sizes of local Quaternary species be made, the mean
lengths of upper and lower carnassials of some species are illustrated (Fig. 76).
It is important to emphasize that the changes in the size of carnassial teeth do
not necessarily reflect equivalent changes in other skeletal elements or overall
body size. For example, it was shown that when the Cornelian Canis mesomelas
from Elandsfontein is considered in relation to the modern form, the skull size
is greater than would have been expected from the size of its lower carnassial.

130

120

118
Relative lengths

of

188 ps oor M,

Elandsfontein

@ Canis mesomelas M,

(Cornelian)

Swartklip 4 at
2 (Florisian) * Vulpes chama P
3 Sea Harvest e Mellivora capensis p4

e (early Holocene)
4 Moderns O Hyaena brunnea My |gg
O Felis serval My,
e

Fig. 76. The mean lengths of P* or M, of some South African Carnivora.

The selected examples show clearly that the size changes in the carnassials
were not constant or synchronous in different species. In general, however, the
largest sizes were attained in the Florisian and early Holocene. An apparent
exception in the illustrated examples is Mellivora capensis, but the Elandsfontein
Bone Circle representative of this species, which is not shown and which
probably postdates that from Swartklip (vide supra), was in fact larger than the
modern variety. A real exception to the general rule is Vulpes chama, the largest
local representative being Cornelian in age. In this instance there is also a
correlation between overall skull size and the size of the carnassials.

As far as actual body sizes are concerned, the situation is obscure. There
are indications that some species such as Vulpes chama and Panthera leo underwent

346 ANNALS OF THE SOUTH AFRICAN MUSEUM

overall decreases in size between the Cornelian and the present. Most species
probably did fluctuate in size, but generally the local fossil varieties appear to
have been larger than their extant counterparts. The only species in which there
might have been varieties which were appreciably smaller than the modern
form is Mellivora capensis.

Perhaps the only confident statement which can be made at present on
the subject of size changes in the Quaternary Carnivora of the south-western
Cape is that such changes did take place and they were apparently of the same
nature as those reported in other parts of the world.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 347

THE ROLE OF CARNIVORA IN THE ORIGIN OF BONE ACCUMULATIONS

In recent years there has been an increasing interest in the manner in
which bones may become accumulated under natural conditions and as a result
of the activities of animals, including man. The principal reason for this interest
in Africa has been the controversy about the origin of the bone assemblages
associated with early hominids (e.g. Dart 1957; Brain 1969). It is now evident
that there has often been more than one factor involved in the accumulation
of any single assemblage.

Animal activity is frequently the cause of bones being accumulated in
particular areas and, since animal behaviour is so variable, it is inadvisable to
generalize about the activities of a particular species in this regard. Each
occurrence being investigated should be considered individually. For example,
the contention that hyaenas are not ‘bone collectors’ (Dart 1956, 1957) has
been shown to be erroneous (Sutcliffe 1970). While it might be expected that
carnivores would be the mammals most likely to be responsible for bone
accumulations because of their predatory and scavenging habits, herbivorous
mammals may also be ‘bone collectors’. In Africa the most obvious example of
this kind is the porcupine (Hystrix), which is known to accumulate large quanti-
ties of bones and other hard materials in its lairs. The bones are gnawed by the
porcupines to keep their constantly growing incisors sharpened and/or at the
correct length, something which cannot be achieved by gnawing on its normally
soft food. It is also possible that they derive some advantage from the pieces of
bone which are ingested. As a result of the physical damage done to bones by
porcupine gnawing, the association of this animal with bone accumulations is
usually readily evident.

This may also be true of assemblages resulting from carnivore activity,
since the teeth of carnivores may leave characteristic marks and fractures on
bones. This has been clearly demonstrated by Brain (1969, 1970), who has
shown that leopards were responsible for at least a part of the fossil accumulation
at Swartkrans. Similarly, hominid activity may leave characteristic imprints
on bone assemblages which represent food residue or raw materials for artefacts
(e.g. Hendey & Singer 1965).

Two kinds of carnivores which have been responsible for bone accumula-
tions have now been mentioned, namely, hyaenas and leopards. In each case
the assemblages investigated differed in composition and situation. The spotted
hyaenas (Crocuta crocuta) studied by Sutcliffe on the Serengeti Plain carried
parts of carcasses which they had scavenged back to their lairs in order to
avoid the loss of this food supply to other carnivores, notably the lion. Brain’s
example of the Swartkrans leopard (Panthera pardus incurva) is similar, although
in this instance bones accumulated in a cavern fortuitously situated beneath
trees into which the leopards had carried their prey.

The second example illustrates the fact that it is not necessary for carnivores
to occupy a sheltered lair in order that they be responsible for the accumulation
of bones. It is to be expected, however, that carnivores which do occupy such

348 ANNALS OF THE SOUTH AFRICAN MUSEUM

lairs are more likely to give rise to concentrations of bones. Consequently,
although it is theoretically possible that any carnivore might be responsible
for bone accumulations, in practise concentrations will only build up when a
carnivore habitually returns to the same place with its kills and scavengings.
Although a study of the habits of extant species will give an indication of those
species which behave in this way, it does not necessarily follow that their extinct
counterparts would have behaved in a similar fashion. For example, the modern
lion is a free-ranging species and has not been reported to be responsible for any
bone accumulations, but the extinct variety of the European Pleistocene (P. leo
spelaea) apparently did occupy caves and might therefore have contributed
some of the bones which are found in caves in Europe today.

Bone accumulations may also be only indirectly associated with the activi-
ties of carnivores. For example, predation and scavenging in any one area may
be confined to a place where animals congregate for a special reason, such as at a
waterhole. In these instances the remains of kills may accumulate in significant
quantities and if they are fairly quickly covered by sediments and the conditions
for bone preservation are good, then they will eventually appear as a relatively
high concentration of fossils in a limited area.

Carnivore activity might also be only a primary factor contributing towards
a bone accumulation. The remains of kills may be transported by some other
agency and eventually be concentrated elsewhere. River transport is an obvious
example of this kind and another is the concentration of fossils on wind deflated
surfaces as a result of the removal of the deposit in which they were originally
incorporated.

In the south-western Cape carnivore activity has apparently contributed
towards the accumulation of some of the local fossil assemblages. This has
already been suggested in the case of Swartklip (Hendey & Hendey 1968),
while at Langebaanweg many of the fossils show signs of physical damage done
by carnivores (Hendey & Repenning 1972). Langebaanweg differs from the
situation at sites such as Swartkrans and Swartklip in that the carnivore kills
were not concentrated in a limited area.

Since the south-western Cape assemblages differ in composition and origin,
it is convenient to consider the more significant assemblages individually and
to examine the evidence for and against carnivore activity in each instance.

There are three contrasting situations evident at the fossil sites of the south-
western Cape, namely:

(1) Accumulation of bones in, or adjacent to, a river estuary (Langebaanweg).

(2) Accumulation in the vicinity of an inland water source (Elandsfontein).
(3) Accumulation in a rock shelter (Swartklip and Sea Harvest).

LANGEBAANWEG

Reference has already been made to the post-mortem damage to certain
fossils recovered at Langebaanweg which results from their having been chewed
by carnivores. Circular depressed fractures (punctate marks), striations on bone

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 349

surfaces, spiral fractures of long bones and other signs of artificial interference
(Fig. 77) are ascribed to activities of those predators and scavengers which are
themselves part of the Langebaanweg assemblage. Some of the damage to bone
is undoubtedly made by the teeth of carnivores. Other is in itself less con-
vincingly so, but when it is associated with definite tooth marks it is also taken
to be the result of carnivore activity.

In the current investigations at Langebaanweg particular attention is
being paid to the condition of zn sztu fossils and also to their mode of occurrence.
There is a great deal of accidental damage done to fossils in the course of the
mining operations at the site, but it is clear that many of the fossils were damaged
at or about the time they were incorporated in the accumulating sediments.
It is well known that the remains of animals which die naturally, or which are
killed by predators, are soon dispersed by the predators themselves, or by
scavengers. The nature of the occurrence of many of the fossils at Langebaanweg
suggest that they are the remnants of carcasses which have suffered in this
manner. Incorporation into the sediments must have been at a variable rate,
since the degree of bone destruction and dispersal of skeletal elements is itself
variable.

Reference has already been made to the inferred nature of the environment
at Langebaanweg at the time that the fossils were being accumulated (vide
supra), and this has in part been determined by the nature of the fossil occur-
rences themselves. In order to illustrate this point, and also to indicate the
manner in which some of the fossils came to be in their present condition and
position, several contrasting examples are examined.

Fossil occurrences in Bed 2

Parts of a Mammuthus subplanifrons skeleton (L 12723) exposed at the north
end of East Stream is a good example of the occurrence of a large mammal in
Bed 2. An upper molar, mandible, some vertebrae, an innominate and some
limb bones were found scattered over an area of several square metres. Although
it is certain that some material was lost during mining operations, and that
other bones might remain in unexcavated deposit, this specimen illustrates
quite clearly than even skeletons of large mammals suffered post-mortem dis-
turbance and were not necessarily preserved in their entirety. In this instance
dismemberment of the skeleton was at least partly due to the activities of
carnivores. A crushed but otherwise largely intact tibia has had parts of both
extremities chewed away and there are marks left by large canine teeth on
adjacent surfaces of the bone. The damage to this bone, and others from the
same skeleton, was almost certainly done by hyaenas. Hyaena abronia, Hyaena
Species B and Percrocuta australis remains are recorded from the same level
of Bed 2, some in the immediate vicinity of East Stream.

The second example is that of a medium-sized herbivore, Nyanzachoerus
(L 14429), the partial skeleton of which was exposed on the bed of East Stream
about 70 metres south of the Mammuthus skeleton. The skull, vertebral column,

ANNALS OF THE SOUTH AFRICAN MUSEUM

30°

“SYIVUI JOO} IIOATUIVD SUIMOYS s}UdWIseIy JUDG ‘q ‘5D
‘QIOATUIVI B JO soUTUvD dy} Aq IPEU d}VUTUIOUUT Uv UO syIvUI 9}vI]9UNg “gq
"1/6961 AAGT UONeAvoXe oy} SulInp potoAodeI VE pog WoIy s}UUIseIy DUOG JO soduIexo [eoIdAT, “Vv
*“S9IOATUIVO JO SUCTION 94} UIOIJ SUT}[NsoI SomMUvEqosULT UOJ SoUOg 0} OSeUIeG “LL “SI

Silmeeerlh SIT

6st @

IT

8

TTT wi mm

6 8 Vv

|

™ : [ltt i inl nil

git ST van! ¢\I 0 6 8

Zit ran Wi i
HUNT a PATA ;

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 351

innominates and a hindlimb lacking only phalanges were recovered. The skull
was badly fragmented before being incorporated into the deposits and there
was no pattern to the distribution of individual skull parts, just as elements of
the postcranial skeleton were completely disarticulated and disordered. Never-
theless, the fact that the remaining parts of this skeleton were found almost in
their entirety over a limited area, indicates that after the loss of three of the
limbs only minor post-mortem disturbance took place. A notable feature of this
skeleton is that parts of the remaining hindlimb and adjacent skeletal elements
are burnt. Fire damage to the sacrum, innominate and femur suggests that
this could have occurred when these elements were still in articulation. How-
ever, damage to the distal extremity of the hindlimb is such that this could
only have occurred after the individual elements had been disarticulated.
The sequence of events must therefore have been the death of the animal,
followed by at least partial dismemberment, then damage to the remains by
fire, and finally incorporation in the accumulating sediments. Damage to some
of the bones and the removal of three of the limbs may well have resulted
from the actions of hyaenas. Following the fire there must have been little or
no further disturbance, otherwise the bones would not have been found in so
close an association. The implications are that this animal died on a land surface
which must have been vegetated, the plant cover then having been burnt. The
remains could not have been transported by water before being covered by
sediment. Five other Nyanzachoerus individuals have so far been recorded in the
vicinity of L 14429. The occurrence of the remains of fish in association with
these specimens suggests that the land surface on which they occur was subse-
quently inundated and buried beneath subaqueously accumulated sediment.

A comparable example of another medium-sized herbivore is the partial
skeleton of a boselaphine (L 13101) which was found near the northern end of
Main Stream. As with the yanzachoerus, the skull was fragmented, the vertebral
column was largely complete, but with this specimen only one scapula and part
of one innominate was otherwise present. Only proximal parts of ribs were
represented. No two elements of the skeleton were found in articulation,
although the remaining parts were found over a very limited area. A large area
was opened up around the skeleton, but no trace of the missing limbs was
found. It is likely that L 13101 represents the remains of a carnivore kill and
that after it had received the attentions of both predators and scavengers just
about all that remained was the skull, vertebral column as far as the sacrum
and the proximal parts of some ribs. A small part of the right innominate was
found a little distance from the vertebrae and this shows very clear tooth marks
left by a large carnivore (? hyaena). The dorsal spines of the thoracic vertebrae
are scarred by the toothmarks of a small carnivore (? viverrid). In this instance
none of the remaining bones had been burnt.

The skeleton of the Hyaena abronia holotype (L 14186) is unusually complete
for a specimen from Langebaanweg. Since it was found only about 15 metres
from the Nyanzachoerus discussed above, and since its occurrence contrasts in

352 ANNALS OF THE SOUTH AFRICAN MUSEUM

many ways with that of the pig, it is worth recording. The skull and about
80 bones of the postcranial skeleton were recovered, with the thorax the only
part not represented. Most of the postcranial bones are complete, but some
have suffered damage which is clearly not recent. For example, both femora
lack their distal ends, both tibiae their proximal ends and the ulnae their
olecranon processes. Neither of the patellae were found. The absence of the
missing skeletal elements, and the damage referred to, could be explained if
the newly-dead animal lay on its stomach with the back arched and the limbs
drawn up. Partial burial leaving the now missing parts above ground surface
would have left the exposed parts in a position to be removed or destroyed by
scavengers, weathering or erosion. Thereafter there must have been some
localized disturbance of the remains, since although they were found over an
area of less than one square metre, the only parts still articulated were the
skull and mandible, and the mandible was in fact slightly displaced.

It is possibly significant that quicksand conditions may develop in water-
logged areas of ‘E’ Quarry, and if such conditions also existed at the time
Bed 2 was being laid down, animals could have been trapped in such areas.
This might have happened in the case of the H. abronia holotype.

There are no known examples of complete or partial skeletons of smaller
vertebrates occurring in the Bed 2 deposits. This is possibly due to the fact that
such skeletons are more readily dispersed by predators, scavengers and other
agencies. Vertebrate microfauna is quite common in Bed 2, in which three small
viverrid species are also recorded. It is likely that these small carnivores were
responsible for the presence of at least some of the small vertebrate remains
being added to the accumulating deposits, in the same way that the larger
carnivores contributed to the presence of the preserved remains of large animals.

One of the most notable features of the deposits in certain areas of ‘E’
Quarry is the abundance of the remains of fossorial species. For example, in
the vicinity of East Stream one chrysochlorid and two bathyergid species are
represented in appreciable numbers. Their relative abundance can be explained
by the fact that death of individuals underground virtually ensures preservation
if soil conditions are favourable, and this was certainly the case with the Bed 2
deposits. The presence of these species is taken as a further indication that there
was a land surface in the East Stream area at the time the deposits were being
laid down.

In addition to the fish remains in the East Stream area mentioned earlier,
a shark, frogs, a penguin and a seal are recorded in association with the remains
of terrestrial vertebrates. They indicate the existence of an aquatic environment
close to the East Stream land surface and the later inundation of this surface.

From the nature of the occurrences described above, it is deduced that at a
certain time, or times, during the deposition of Bed 2, predation and scavenging
of animals took place on the fringes of an estuary and that the activities of
carnivores were a major factor involved in the mortality of the animals now
preserved as fossils. In addition, the actual nature of the fossil occurrences was

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 353

at least partly determined by the post-mortem disturbance of animal carcasses
by carnivores. Although much of the damage to, and dismemberment of
skeletons in this horizon can be attributed to hyaenas, other carnivores were
also involved, as the evidence of toothmarks shows.

Another point of minor significance arises in this connection. Associated
with a Percrocuta specimen from Bed 2 (L 13033) were a series of bone fragments
the edges of which are apparently abraded. These specimens may owe their
condition to erosion in the digestive tract of the Percrocuta, rather than having
been transported by water or weathered on a land surface. The ability of hyaenas
to crack and swallow bones is well known (see Dart 1956: 43), and the bone
fragments which are swallowed are eventually completely reduced by digestion
and the residue excreted. If the bone fragments associated with the Percrocuta
were indeed stomach contents, then their digestion was obviously not far
advanced. The significance of this particular discovery is that other, apparently
water-rolled or surface weathered bone fragments may actually be stomach
contents of hyaenas and possibly erroneous conclusions could be drawn from
their presence in the deposits.

Fossil occurrences in Bed 3a

There appears to be a higher proportion of very fragmented bone in Bed
ga than is the case in Bed 2. As before, tooth marks indicate that fragmentation
was caused by carnivores. Most often the broken edges are fresh and sharp,
indicating that the fragments were never swallowed. In controlled excavations
carried out in Bed ga, it was noted that many of the bone fragments stand
vertically, or near vertically in the deposits, which suggests that they were
dropped or moved in soft waterlogged or subaqueous deposits. If they were
transported by water, the movement must have been very limited because the
edges of the fractures are still remarkably fresh in appearance.

Hyaenid remains are not uncommon in these deposits, so it is likely that
once again they were to a large extent responsible for the condition of the
in situ fossils of Bed 3a. However, seals are also very common in Bed 3a and
perhaps they too played a role in the dismemberment and fragmentation of
skeletons by scavenging floating or stranded carcasses of terrestrial mammals.
No record could be located of this type of behaviour in seals, so that this
opinion is pure speculation based only on the fact that the seal is perhaps the
most commonly occurring carnivore in Bed ga. It is perhaps more likely that
the seal remains themselves owe their condition partly to the shoreline scaven-
ging of hyaenids.

Ecology of the Langebaanweg Carnivora

Since the Langebaanweg Carnivora are the only local fossil series which
differs appreciably from the modern carnivore fauna, it is of interest to consider
their ecology in some more detail. In order to facilitate comparisons between
the fossil and modern representatives of the group, they were categorized in the
arbitrary manner illustrated in Figure 78.

354 ANNALS OF THE SOUTH AFRICAN MUSEUM

AQUATIC forms
(A)

MARINE FRESH-WATER
(Al) (A2)
TERRESTRIAL forms
(T)

Miro

SMALL MEDIUM LARGE
(TI) (T2) (T3)

Fig. 78. Categories of Carnivora represented in the south-western Cape Province.

The marine aquatic group (Ar) is the only one in which the Langebaanweg
and modern faunas are directly comparable. A single resident pinniped species
is represented in both faunas, the principal difference being that the Pliocene
one is a monachine, while the modern species is an otariid. Although there is
no certain evidence that the Langebaanweg Prionodelphis was not an exclusively
piscatorial species, it has been suggested that it was an estuarine scavenger.
Similar behaviour is not recorded for the extant Arctocephalus pusillus.

The fossil and modern representatives of the freshwater group (Ag) differ
in that the former includes one, and possibly two giant species, while there is
only a single, relatively small otter in the modern fauna. Modern Aonyx capensis
includes fish, frogs, crabs and molluscs in its diet. Although crabs have not been
recorded from Langebaanweg, it is very likely that they were present, so the
diet of the fossil otter(s) may have been essentially similar to that of the modern
species. The size differences are, however, unexplained and it may well be that
the Pliocene otters were different to modern Aonyx in their habits.

If the unclassified carnivore from Langebaanweg is indeed an otter, then
it and not Prionodelphis may have fulfilled the role of an aquatic scavenger. ‘The
very large canine teeth of this species would have been admirably suited to
tearing flesh from carcasses and its posterior cheekteeth, which are not known,
may have been adapted for bone-crushing. Both it and the Enhydriodon may have
contributed to the accumulation of terrestrial vertebrate bones in the Lange-
baanweg deposits.

There is a marked difference between the modern and fossil small terrestrial
carnivores (group Tr) in that the modern fauna includes a much wider variety
of species. This group is comprised mainly of viverrids and it has already been
indicated that their radiation was a feature of the Pleistocene. The rather gene-
ralized Langebaanweg Genetta and Herpestes species may have occupied a
variety of ecological niches which are today filled by more specialized forms.
The three small carnivores recorded from Langebaanweg were probably
responsible for the accumulation of many of the vertebrate microfaunal bones
at the site. The absence in the Langebaanweg fauna of a small felid comparable

4 + >, 2
Se :
natneomtitiene tend Oi ae ee ans Sel

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 355

to Felis libyca may simply be due to an inadequacy in the fossil record.

There is a less marked difference in the number of species making up the
group of medium-sized terrestrial carnivores (T2), although the highly special-
ized Otocyon and Proteles have no recorded counterparts in the Langebaanweg
fauna. The early mellivorine from Langebaanweg is balanced by an advanced
form in the modern fauna, although it is likely that the habits of the two species
differed, modern Mellwora capensis being a more omnivorous form. The smaller
of the Langebaanweg felids (Felis aff. issiodorensis, F. obscura) probably have the
caracal and serval as their modern counterparts.

Comparisons between the modern and fossil faunas of group T3 can be
made in three categories. Firstly, there are those species which prey on small
vertebrates and scavenge from the kills of large predators. The fossil representa-
tives are Viverra leakeyi and perhaps the unidentified canid, while Canis mesomelas
is their modern counterpart. Secondly, there are the large predators which are
represented at Langebaanweg by the two machairodonts and Dinofelis and in
the modern fauna by Panthera leo and P. pardus. In addition, the modern fauna
includes the cursorial Lycaon pictus which has no definite counterpart at Lange-
baanweg, while the giant Agriotherium has no modern counterpart. Finally
there are the large scavengers, the Hyaenidae. They are well represented and
complementary to the sabre-toothed cats in the Langebaanweg fauna and less
diverse and complementary to the large predators of the modern fauna. Hyaena
Species B and Hyaenictis preforfex may have been more actively predaceous than
modern hyaenids. It is this group which was probably the most important
contributor to the accumulation of fossils at Langebaanweg and whose activities
determined the nature of many of the fossils which have been preserved.

On the basis of the available record, the following major changes were
undergone by the south-western Cape carnivore fauna between the Lange-
baanian and the historic period:

(1) One family of seals was replaced by another, but in each instance only one resident species
is represented.

(2) Giant fresh-water carnivores were replaced by a smaller form.

(3) There was a diversification among the smaller predators.

(4) Highly specialized insectivorous and omnivorous carnivores were evolved.

(5) There was a change in the representation of large terrestrial carnivores, including the

replacement of a civet by a jackal, the replacement of sabre-toothed cats by ordinary
forms, and a reduction in the variety of large scavengers represented.

To conclude the account of the Langebaanweg Carnivora, they are com-
pared and contrasted with those of the North American Hagerman Local Fauna
(Bjork 1970). The two assemblages are broadly contemporaneous and while
that from Langebaanweg is from a coastal estuarine environment, ‘the environ-
ment of deposition at Hagerman during the late Pliocene was a broad flood
plain with trees and grassland adjacent to the tributary streams of Lake Idaho’
(Bjork 1970: 51). The recorded vertebrate faunas from the two occurrences
are essentially similar in composition (i.e. carnivores, herbivores, reptiles, birds,
etc.), although the species represented are, of course, quite different.

wenusueam ££

ait ae

3 56 ANNALS OF THE SOUTH AFRICAN MUSEUM

The two assemblages include a similar number of fissiped carnivore species,
but there is a taxonomic correspondence only at the family level (Table 91).
Only one of the families represented at Langebaanweg, the Viverridae, is not
recorded at Hagerman and this gap is filled by the Mustelidae, which make
up 50% of the species recorded in the Hagerman carnivore fauna. At Lange-
baanweg the Mustelidae make up at most only 16% of the carnivore species
represented, although the Mustelidae and Viverridae together make up about
45% of the total. Only one hyaenid is doubtfully recorded from Hagerman on
the basis of a single deciduous tooth, whereas the five Langebaanweg species are
represented by numerous specimens. ‘There is, however, a large canid from
Hagerman (Borophagus) which evidently fulfilled an hyaenid role in North
America. Only in the case of the Felidae is there any marked resemblance in
the nature of the species represented in the two assemblages.

TABLE QI

The representation of fissiped carnivores in the late Pliocene
occurrences at Langebaanweg, compared with that of the
Hagerman Local Fauna (Bjork 1970)

Number of species

Langebaanweg Hagerman
Canidae . : : I 2
Ursidae . : : I 2
Mustelidae_ . ‘ 2 Or 3 9
Viverridae . : 4 )
Flyacnidae = sa. 25 PI
Felidae . ; : 5 4

Langebaanweg Hagerman

No. of species 19 18
No. of genera c. 14 16
No. of families 6 4 0r 5

Langebaanweg/
Hagerman
Species in common : é : , O
Faunal resemblance (%) : ; Oo
Genera in common : ‘ : : I
Faunal resemblance (%) : 5 7
Families in common. : : : 4 0r 5

Faunal resemblance (%) , : 100

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 357

It is remarkable that two carnivore faunas, which fulfil the same ecological
role in essentially similar environments at the same period in time, should
differ in so many ways. The indications are that even in the late Tertiary the
faunas of South Africa and North America had long since been developing
independently from one another.

ELANDSFONTEIN

The Elandsfontein fossil exposures are situated close to one of the few
permanent waterholes anywhere in the Sandveld area of the south-western
Cape. The river nearest to the site is the Sout, some 10 km distant, which under
existing climatic conditions has water in it for only a few months each year and
even then it is seldom, if ever flowing strongly. The water table in the Elands-
fontein area is close to the surface and this, coupled with the presence of the
existing waterhole, suggests that it might have been a local source of fresh water
for a long time. Consequently, it is an area where animals are likely to have
concentrated at least as far back as the Cornelian, to judge from the fossil record.

The situation is, therefore, one where there is likely to have been concen-
trated carnivore activity over prolonged periods and many of the fossils at the
site probably represent the remains of carnivore kills. The Elandsfontein fossils
have not been studied with the view to establishing evidence of carnivore activity
in the same detail as those from Langebaanweg. In addition, there are some
complicating factors which would have to be taken into consideration in any
detailed analysis of the nature of the occurrences.

Many of the fossils from this site have been gnawed by porcupines (Singer
1956), and some of the recorded fossil concentrations may represent now exposed
remnants of the subterranean lairs of these rodents. Porcupines still occupy such
lairs in the area today, although none has been excavated and examined.

A second difficulty is that there has clearly been some contribution to the
fossil occurrences as a result of hunting in the area by primitive man. For
example, an Early Stone Age ‘living floor’ uncovered at the site revealed an
association of artefacts with a variety of mammalian remains (Singer & Wymer
1968). Since there is evidence also of Middle Stone Age, Late Stone Age and
historic period hominid activity, it follows that each of these periods of occupa-
tion may have added faunal remains to the deposits.

There are, however, some bone accumulations which might have resulted
from the activities of carnivores. One of these is the Bone Circle occurrence,
which was repeatedly referred to earlier in connection with the Florisian
Carnivora from this site. Some accounts of this occurrence have previously been
given elsewhere (Inskeep & Hendey 1966; Singer & Heltne 1966). The earlier
opinions concerning the possible origin of this occurrence are here rejected.

There are marked similarities between the nature of this occurrence and
those at Swartklip, the most obvious and perhaps also the most misleading
difference between them being that whereas those at Swartklip are vertical
exposures, the Bone Circle occurrence was a horizontal exposure. The Swartklip

358 ANNALS OF THE SOUTH AFRICAN MUSEUM

sites are more obviously in the form of subterranean animal lairs, but were
they to have been exposed horizontally they might well have taken on the
appearance of the Bone Circle occurrence. If it is assumed that carnivores such
as Canis mesomelas made use of aardvark (Orycteropus) burrows, it follows that
bones might become accumulated in the lair by its new occupants and might
subsequently be exposed by the very active wind erosion which is prevalent
at the site. C. mesomelas is quoted as an example because a number of specimens
of this species were the only carnivore remains associated with a miniature
‘bone circle’ at the northern end of the site. The original Bone Circle also
included Hyaena, Crocuta and Mellivora remains and in this instance these animals
might also have been one-time occupants of the hypothetical lair. In neither
of the two occurrences referred to was there evidence of porcupine activity.

It was suggested earlier in connection with the Elandsfontein Suricata that
those carnivores which occupy subterranean lairs might be more commonly
represented as fossils than those which do not. Mellivora capensis was mentioned
as an obvious example in addition to the Suricata. ‘The fact that C. mesomelas,
and to a lesser extent also Hyaena brunnea, is so abundantly represented at
Elandsfontein at least suggests the possibility that it did frequently, or habitually,
occupy such lairs.

There is clearly a potential for a more thorough investigation of bone
concentrations at the Elandsfontein site, but they are here regarded as the
results of carnivore activity rather than ‘some freak of deflation’ (Singer &
Wymer 1968: 64), although deflation is the mechanism by which they have
been exposed.

There is other more direct evidence of carnivore activity at Elandsfontein.
The machairodont humeri 3058/9 show signs of tooth marks which were
definitely not made by porcupines, but which could have been made bya
carnivore. More convincing still is the Mellivora braincase 20981, which has two
circular holes 4 mm in diameter and 21 mm apart on its dorsal side and a
similar hole in the basi-occipital. The spacing and size of these holes suggests
that they were made by the canines of a carnivore of about the size of Mellivora
itself, although the canines of C. mesomelas also fit them quite well. This specimen
is reminiscent of the leopard-punctured australopithecine skull described by
Brain (1970).

An aspect of the Elandsfontein fossil occurrences which might be an indica-
tion of carnivore activity is the fact that there are remarkably few recorded
instances of skeletal elements having been found in articulation. This indicates
a great deal of post-mortem disturbance and dispersal of skeletons of the kind
which results from the actions of both predators and scavengers when animal
remains are not fairly quickly buried. Exceptions to this general rule on skeletal
dispersal are most commonly recorded in the ‘bone circle’ occurrences. Actual
articulations are known, but more commonly individual skeletal elements were
slightly separated from one another. This is also a feature of the Swartklip
occurrences and in both instances it appears to be skulls and cervical vertebrae

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 359

or the distal extremities of limbs which are found in the closest association.

One of the very few recorded instances of dispersal of a single individual
outside of the ‘bone circles’ is that of a Pelorovis skeleton (16710). The almost
complete skull was found ventral side uppermost a few metres away from both
halves of the mandible and most of the anterior part of the vertebral column.
There are no obvious tooth marks on any of the skeletal elements, but the
disturbance of the skeleton is of the kind which might result from the actions of
an hyaenid. On the other hand, this and other instances of disturbance might
also be the result of butchering by hominids.

SWARTKLIP AND SEA HARVEST

Some evidence supporting the conclusion that the Swartklip assemblages
accumulated in carnivore lairs has already been discussed elsewhere (Hendey
& Hendey 1968: 48-51), but this may now be qualified and enlarged upon as a
result of more recent observations.

Firstly, it was stated that the Swartklip assemblages did not include
porcupine gnawed bones. These have in fact now been found at Site 1, but
the numbers are so small that it is unlikely that the porcupine ever played a
significant role in the accumulation of bones at these sites.

The presence of appreciable amounts of broken ostrich eggshell and at least
two complete ostrich eggs was a problematical aspect of the assemblages,
although it was suggested that these eggs formed part of the diet of one of the
occupants of the lairs. Recently a photograph appeared in a Cape Town news-
paper (The Argus, 19 April 1971) showing lionesses in a Kenyan wildlife park
eating ostrich eggs and since it has been suggested that Panthera leo was one of
the occupants of the Swartklip lairs, it is possible that this species was responsible
for bringing in and breaking up the eggs found in the deposits. It was mentioned
earlier that lions in Africa are not known to be occupants of caves and rock
shelters, although this does not apply in the case of the Pleistocene lions of
Europe. The Swartklip fossils date from the late Pleistocene when conditions
in the south-western Cape were colder than they are today, so it is possible that
the prevailing climate of that time prompted the local lions to make use of
sheltered lairs, even though they do not do so today.

Just as porcupine gnawmarks on bones from Swartklip have now been
recognized, so have tooth marks and fractures caused by carnivores. Bone
fragments with spiral fractures and punctate marks, identical in appearance
with those from Langebaanweg, have been recovered.

Although the Swartklip lairs were probably occupied by a variety of species
over a period in time, the ones which are regarded as being responsible for
most if not all of the bones introduced into the lairs are Canis mesomelas, Hyaena
brunnea, Panthera leo and Hystrix africaeaustralis.

Essentially similar observations and conclusions apply in the case of the
Sea Harvest site, although in this instance both Crocuta crocuta and Panthera pardus
are added to the list of possible bone collectors.

360 ANNALS OF THE SOUTH AFRICAN MUSEUM

CONCLUSION

For a region which is so limited in extent, the south-western Cape Province
is providing a remarkable record of animal life during the late Cenozoic. The
existing local record, taken in conjunction with that from the rest of southern
Africa, has already given a fairly clear indication of the nature and evolution
of higher forms of life in the subcontinent during this period in time. In general,
the fossil occurrences of comparable age in East Africa are more significant
than those to the south, but the records from the two areas are complementary
to one another. The same applies in the case of the fossil faunas of the south-
western Cape in relation to those from the rest of southern Africa. They are
an integral part of the record as a whole and are, therefore, of more than just
local interest.

In the present report the tendency was to emphasize the occurrences at
Langebaanweg. This was done because the fauna from this locality is unique
in so many ways and it is regarded as the most important of the local fossil
faunas. .

Many notable discoveries have already been made at Langebaanweg.
For example, the fauna includes one of the earliest of the true elephants, an
early ancestor of the white rhinoceros, the first fossil penguin recorded in
Africa, the first bear from sub-Saharan Africa and what is perhaps the last of
the African boselaphine antelopes. New or unusual records of this kind are of
interest in themselves, but their real significance must be assessed against a
broader background. Thus while the seal, Prionodelphis capensis, is only the second
record of the genus and one which allows adequate definition of the genus for
the first time, its significance lies chiefly in the additional insight it has given
on the possible origin and evolution of the Antarctic seals. Previously little was
known about this subject, even though surviving populations of these seals
may be counted in millions and their area of distribution covers millions of
square kilometres of the earth’s surface.

In addition, the Langebaanweg fossils have given the first real indications
of the nature of the local environment in late Pliocene times. It was evidently
very different from that of the present and it is expected that geological and
other studies stimulated by the palaeontological investigations at Langebaan-
weg will contribute more to the knowledge of the late Pliocene environment.

The local Quaternary record is of interest since it includes assemblages
ranging in age from Cornelian to Holocene, which makes it possible to trace
in some detail the final stages in the evolution of the modern fauna of the
region. This part of the local fossil record is less remarkable than that from
Langebaanweg, but this does not diminish its importance. Attempts to deter-
mine the character of the historic period fauna, which is now much depleted
as a result of human activity, have been undertaken in conjunction with the
Quaternary palaeontological investigations. One of the significant aspects of
the recent fauna of the region is the fact that it was from the south-western
Cape that many of the species which characterize the Ethiopian faunal region

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 361

were first described.

Additional interest in the late Cenozoic fauna of the south-western Cape
stems from the fact that this region is atypical of sub-Saharan Africa as a whole,
since it is situated at the southern continental extremity and is climatically
differentiated. The biological implications of its geographical location are
significant in both palaeontological and neontological studies.

Although many investigations have already been carried out on the fossil
mammals from the south-western Cape, the research potential in this field is
still good. Boné & Singer (1965: Table 1) recorded 13 carnivore species from
two sites in the south-western Cape, while the present report records at least
45 species from more than half a dozen occurrences. To a greater or lesser
extent a species list increase can be expected in other mammalian orders as well.

It is striking how little information on the late Cenozoic mammals of
Africa is included in a recent book on the fossil mammals of the world (Kurtén
1971), yet this was the time when the modern African fauna, which by any
standard is exceptional in quantity and variety, went through the final stages
in its development. In his review of the Pliocene, Kurtén (1971: 152) wrote
that this epoch was ‘something of a paradise lost, a climax of the Age of Mam-
mals before the coming of the cold; a time when life was richer, more exuberant
than ever before or after’. He goes on to say that conditions which prevailed
at this climax are now found only in a few areas, and those are mainly in Africa.
We are thus faced with the curious situation that the continent in which the
modern mammals represent what may well be the acme of mammalian evolu-
tion is also the one in which the fossil mammal record is perhaps least well
known.

The situation is, however, improving and the recent investigations into
the origin and evolution of the Hominidae has brought about an increased,
although still largely incidental interest in other African fossil mammals. It is
probably true to say that some of the more significant advances in recent years
in the field of palaeomammalogy have been made in Africa. The traditional
view that Africa played a somewhat subsidiary role to Eurasia in the origins
and evolution of its own fauna is now being modified. It had also become
fashionable to regard Africa as a refuge for archaic forms and, while this is
not without foundation, there has perhaps been a tendency to underemphasize
the role this continent has played in the later history of mammals.

With these points in mind, it is hoped that the report is concluded on a
note of expectation rather than achievement. There is much which has still to
be done.

SUMMARY

A general account is given of the recorded fossil Mammalia of the south-
western Cape Province and also of the more important late Cenozoic fossil
occurrences in this region. A formal nomenclature for the Pliocene deposits at
Langebaanweg is suggested and a new system of age names for the South

362 ANNALS OF THE SOUTH AFRICAN MUSEUM

African late Cenozoic, based on fossil mammal faunas, is proposed.
A more detailed account is given of the species of Carnivora known from
the local late Cenozoic and the following new species are described:

Percrocuta australis, Hyaena abronia, Hyaenictis preforfex, Felis obscura and Suricata
major.

Promellivora Pilgrim, 1932 is rejected and the material identified as such is
referred to Mellwora. Vishnuictis Pilgrim, 1932 is reduced to the status of a sub-
genus of Viverra and Pseudocwetta Petter, 1967 is included in Viverra (Civettictis).
It is suggested that the Hyaena hyaena and Hyaena brunnea groups be subgeneri-
cally distinguished, with the new name, Parahyaena, being applied to the latter
group. Megantereon problematicus Collings, 1972 is provisionally listed as Homo-
therium sp. indet. Three previously unrecorded skulls of the Cape lion, Panthera
leo melanochaita, are described.

Some aspects of the zoogeography of local mammals are considered and the
role of carnivores in the origin of bone accumulations is discussed. Brief com-
ments on topics such as the local non-mammalian fossils and size changes in
Quaternary carnivores are also included.

ACKNOWLEDGEMENTS

The current investigation into the late Cenozoic mammals of the south-
western Cape Province was initiated in 1965, while the systematic study of the
Carnivora was undertaken intermittently between 1967 and 1970 and then
more continuously during the succeeding two years. During this time many
persons contributed directly and indirectly to the investigation. ‘They are too
numerous to mention individually, but being aware of the assistance they have
given, they will also, I hope, be aware of my regard and gratitude.

The present report was the direct result of encouragement received from
Professor Lester C. King of the University of Natal and I am greatly indebted
to him for his interest and assistance in the undertaking. I also wish to express
my deep appreciation to Dr C. K. Brain of the Transvaal Museum, who super-
vised the research.

This research is being supported by the South African Council for Scientific
and Industrial Research, Chemfos Ltd. (a subsidiary of the African Metals
Corporation) and Shell South Africa (Pty.) Ltd. The Wenner-Gren Foundation
for Anthropological Research, New York, provided the vehicle used in the
field work (Grant no. 2752-1834). :

Finally I wish to acknowledge the assistance and encouragement given to
me by my wife; hers was the most difficult task of all.

This work was submitted in partial fulfilment of the requirements for the
degree of Doctor of Philosophy in the Department of Geology of the University
of Natal.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 363

REFERENCES

Acocks, J. P. H. 1953. Veld types of South Africa. Mem. bot. Surv. S. Afr. 28: 1-192.
ALLEN, J. A. 1925. Carnivora collected by the American Museum Congo Expedition. Bull.
_ Am. Mus. nat. Hist. 47: 73-281.

ANDERSON, E. 1970. Quaternary evolution of the genus Martes (Carnivora, Mustelidae).
Acta zool. fenn. 130: 1-132.

ARAMBOURG, C. 1959. Vertébrés continentaux du Miocéne supérieur de |’Afrique du Nord.
Publs Serv. Carte géol. Algér. Paléont. Mem. 4: 1-159.

Bjork, P. R. 1970. The Carnivora of the Hagerman Local Fauna (Late Pliocene) of south-
western Idaho. Trans. Am. phil. Soc. 60: 1-54.

Bone, E. L. & SinceEr, R. 1965. Hipparion from Langebaanweg, Cape Province and a revision
of the genus in Africa. Ann. S. Afr. Mus. 48: 273-397.

Boniray, M. F. 1969. Etudes francaises sur le Quaternaire. Faunes Quaternaires de France.
Bull. Ass. franc. Etude Quatern. 1969 (suppl.): 127-142.

BouriERE, F. 1963. Observations on the ecology of some large African mammals. Jn: HOWELL,
F. C. & BOURLIERE, F., eds. African ecology and human evolution. Publs Anthrop. Viking Fund 36:
43-54:

Brain, C. K. 1969. Who killed the Swartkrans ape-men? Bull. S. Afr. Mus. Ass. 9: 127-139.

Brain, C. K. 1970. New finds at the Swartkrans australopithecine site. Nature, Lond. 225:
III2—-111Q.

Brain, C. K. & MEEsTER, J. 1964. Past climatic changes as biological isolating mechanisms in
southern Africa. In: DAvis, D. H. s., ed. Ecological studies in southern Africa: 332-340. The
Hague: Junk.

Broom, R. 190g. On the evidence of a large horse recently extinct in South Africa. Ann. S. Afr.
Mus. 7: 281-282.

Broom, R. 1937a. On some new Pleistocene mammals from limestone caves of the Transvaal.
S. Afr. F. Sct. 33: 750-768.

Broom, R. 19376. Netices of a few more new fossil mammals from the caves of the Transvaal.
Ann. Mag. nat. Hist. (10) 20: 509-514.

Broom, R. 1939. A preliminary account of the Pleistocene carnivores of the Transvaal caves.
Ann. Transv. Mus. 19: 331-338.

Broom, R. 1946. The South African fossil ape-men. The Australopithecinae. Transv. Mus.
Mem. 2: 1-153.

Broom, R. 1948. Some South African Pliocene and Pleistocene mammals. Ann. Transv. Mus.
21: 1-38.

BROTHWELL, D. R. 1963. Digging up bones. London: British Museum (Natural History).

BuTLer, P. M. 1969. Insectivores and bats from the Miocene of East Africa: new material.
In: LEAKEY, L. S. B., ed. Fossil vertebrates of Africa. 1: 1-37. London: Academic Press.

Butter, P. M. & Hopwoop, A, T. 1957. Insectivora and Chiroptera from the Miocene rocks
of Kenya Colony. Fossil Mammals Afr. 13: 1-35.

ButzeEr, K. W. 1961. Paleoclimatic implications of Pleistocene stratigraphy in the Mediterranean
area. Ann. N. Y. Acad. Sci. 95: 449-456.

CAHALANE, V. H. 1961. Mammals of North America. New York: Macmillan.

CuHuRCHER, C. S. 1956. The fossil Hyracoidea of the Transvaal and Taungs deposits. Ann.
Transv. Mus. 22: 477-501.

CuurcHER, C. S. 1970. The fossil Equidae from the Krugersdorp caves. Ann. Transv. Mus.
26: 145-168.

Ciark, J. D. 1957. Prehistory: Third Pan-African Congress Livingstone 1955. London: Chatto &
Windus.

Cope oF STRATIGRAPHIC NOMENCLATURE. 1961. Bull. Am. Ass. Petrol. Geol. 45: 645-665.

CorEtTzeEE, C. G. 1967. Preliminary identification manual for African mammals. 7. Carnivora. Washing-
ton: Smithsonian Institution.

Cotuincs, G. E. 1972. A new species of machaerodont from Makapansgat. Palaeont. afr. 14:
87-92.

Cooke, H. B. S. 1947. Some fossil hippotragine antelopes from South Africa. S$. Afr. 7. Sct.
43: 226-231.

Cooke, H. B. S. 1950. A critical revision of the Perissodactyla of southern Africa. Ann. S. Afr.
Mus. 21: 393-479.

a
i
:

P nm Y

364. ANNALS OF THE SOUTH AFRICAN MUSEUM

Cooks, H. B. S. 1955. Some fossil mammals in the South African Museum collections. Ann. S.
Afr. Mus. 42: 161-168.

Cooke, H. B. S. 1963. Pleistocene mammal faunas of Africa, with particular reference to southern
Africa. In: HOWELL, F. C. & BOURLIERE, F., eds. African ecology and human evolution. Publs
Anthrop. Viking Fund 36: 65-116.

Cooke, H. B. S. 1964. The Pleistocene environment in southern Africa. In: DAVIS, D. H. S., ed.
Ecological studies in southern Africa: 1-23. The Hague: Junk.

Cooke, H. B. S. 1967. The Pleistocene sequence in South Africa and problems of correlation.
In: BISHOP, W. W. & CLARK, J. D., eds. Background to evolution in Africa: 175-184. Chicago:
University Press.

Cooke, H. B. S. 1968. Evolution of mammals on southern continents. II. The fossil mammal
fauna of Africa. Q. Rev. Biol. 43: 234-264.

Cooke, H. B.S. & Mactio, V. J. 1972. Plio-Pleistocene stratigraphy in East Africa in relation
to proboscidean and suid evolution. Jn: BISHOP, W. W. & MILLER, J. A., eds. Calibration of
hominoid evolution: 303-329. Edinburgh: Scottish Academic Press.

CoprEns, Y. 1972. Tentative de zonation du Pliocéne et du Pléistocéne d’Afrique par les grands
Mammiferes. C. r. Acad. Sci. Paris 2°74: 181-184.

DARLINGTON, P. J. 1957. <oogeography. The geographical distribution of animals. New York:
Wiley.

DaRLINGTON, P. J. 1965. Biogeography of the southern end of the world. Cambridge: Harvard Uni-
versity Press.

Dart, R. A. 1956. The myth of the bone-collecting hyena. Am. Anthrop. 58: 40-62. ~

Dart, R. A. 1957. The osteodontokeratic culture of Australopithecus prometheus. Transv. Mus.
Mem. 10: 1-105.

Davis, D. D. 1964. The giant panda. A study of evolutionary mechanisms. Fieldiana, ool.
Mem. 3: 1-339.

DE Graaf, G. 1960. A preliminary investigation of the mammalian microfauna in Pleistocene
deposits of caves in the Transvaal System. Palaeont. afr. 7: 59-118.

DE HEINZELIN, J. 1969. Le groupe de l’?Omo et l’4ge du Pleistocene. Bull. Soc. belge Géol., Paléont.,
Hydrol. 78: 1-5.

Dietricu, W. O. 1942. Altestquartare Saugetiere aus der stidlichen Serengeti, Deutsch-Ost
Afrika. Palaeontographica (A) 94: 43-133.

Dorst, J. & DANDELOT, P. 1970. A field guide to the larger mammals of Africa. London: Collins.

Dreyer, T. F. & Lye, A. 1931. New fossil mammals and man from South Africa. Bloemfontein:
Nasionale Pers.

Du Torr, A. L. 1917. The phosphate of Saldanha Bay. Mem. geol. Surv. Un. S. Afr. 10: 1-31.

ELLERMAN, J. R., Morrison-Scott, T. C. S. & Hayman, R. W. 1953. Southern African mammals
1758-1951: a reclassification. London: British Museum (Natural History).

Ennoucui, E. 1953. Un nouveau genre d’Ovicapriné dans un gisement pléistocéne de Rabat.
C. r. somm. Séanc. Soc. géol. Fr. 8: 126-128.

ERDBRINK, D. P. 1953. A review of fossil and recent bears of the Old World. Deventer: De Lange.

Ewer, R. F. 1954. The fossil carnivores of the Transvaal caves. The Hyaenidae of Kromdraai.
Proc. zool. Soc. Lond. 1242 565-585.

Ewer, R. F. 1955a. The fossil carnivores of the Transvaal caves. The Hyaenidae, other than
Lycyaena, of Swartkrans and Sterkfontein. Proc. zool. Soc. Lond. 124: 815-837.

Ewer, R. F. 19550. The fossil carnivores of the Transvaal caves. The Lycyaenas of Sterkfontein
and Swartkrans, together with some general considerations of the Transvaal fossil hyaenids.
Proc. zool. Soc. Lond. 124: 839-857.

Ewer, R. F. 1955c. The fossil carnivores of the Transvaal caves: Machairodontinae. Proc.
zool. Soc. Lond. 125: 587-615.

Ewer, R. F. 1956a. The fossil carnivores of the Transvaal caves: Felinae. Proc. zool. Soc. Lond.
126: 83-95.

Ewer, R. F. 1956. The fossil carnivores of the Transvaal caves: Canidae. Proc. zool. Soc. Lond.
126: 97-119.

Ewer, R. F. 1956c. The fossil carnivores of the Transvaal caves: two new viverrids, together
with some general considerations. Proc. zool. Soc. Lond. 126: 259-274.

Ewer, R. F. 1956d. Some fossil carnivores from the Makapansgat valley. Palaeont. afr. 4: 57-67.

Ewer, R. F. 1957a. A collection of Phacochoerus aethiopicus teeth from the Kalkbank Middle
Stone Age site, Central Transvaal. Palaeont. afr. 5: 5-20.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 365

Ewer, R. F. 1957). Faunal evidence on the dating of the Australopithecinae. Jn: CLARK, J. D.,
ed. Prehistory: Third Pan-African Congress Livingstone 1955. London: Chatto & Windus.
Ewer, R. F. 1962. A note on some South African fossil otters. Navors. nas. Mus., Bloemfontein

1: 275-279.

Ewer, R. F. 1967. The fossil hyaenids of Africa—a reappraisal. Jn: BISHOP, W. W. & CLARK, J. D.,
eds. Background to evolution in Africa: 109-123. Chicago: University Press.

Ewer, R. F. & Cooke, H. B. S. 1964. The Pleistocene mammals of South Africa. In: Davis,
D. H. S., ed. Ecological studies in southern Africa: 35-48. The Hague: Junk.

Ewer, R. F. & Sincer, R. 1956. Fossil Carnivora from Hopefield. Ann. S. Afr. Mus. 42: 335-347.

FIcARELLI, G. & Torre, D. 1970. Remarks on the taxonomy of hyaenids. Palaeoniogr. ital.
46: 13-33.

Firzsimons, F. W. 1919. The natural history of South Africa. Mammals. 2, London: Longmans,
Green.

Fuint, R. F. 1971. Glacial and Quaternary geology. New York: Wiley.

Fiower, W. H. 1885. Osteology of the Mammalia. London: Macmillan.

Forster, J. R. 1781. Tyger-cat of the Cape of Good Hope. Phil. Trans. R. Soc. Lond.
41: 4-6.

GapsuniA, L. & RusinsteIn, M. 1968. On the correlation of the Cenozoic deposits of Eurasia
and North America based on fossil mammals and absolute age data. Jnt. geol. Congr. 23
(10): 9-17.

Gazin, C. L. 1933. New felids from the Upper Pliocene of Idaho. 7. Mammal. 14: 251-256.

Gentry, A. W. 1970. The Bovidae (Mammalia) of the Fort Ternan fossil fauna. Jn: LEAKEY,
L. S. B. & SAVAGE, R. J. G., eds. Fossil vertebrates of Africa 2: 243-323. London: Academic Press.

Grecory, W. K. & HELLMAN, M. 1939. On the evolution and major classification of the civets
(Viverridae) and allied fossil and recent Caner: A phylogenetic study of the skull and
dentition. Proc. Am. phil. Soc. 81: 309-392.

Harper, F. 1945. Extinct and vanishing mammals of the Old World. Spec. Publs Am. Comm.
int. wild Life Prot. 12: 1-850.

Harrison, D. L. 1968. The mammals of Arabia. 2: London: Ernest Benn.

HaucurTon, S. H. 1932a. On the phosphate deposits near Langebaan Road, Cape Province.
Trans. geol. Soc. S. Afr. 35: 119-124.

HaucutTon, S. H. 1932b. The fossil Equidae of South Africa. Ann. S. Afr. Mus. 28: 407-427.

Hemmer, H. 1965. Zur Nomenklatur und Verbreitung des Genus Dinofelis Zdansky, 1924
( Therailurus Piveteau, 1948). Palaeont. afr. 9: 75-89.

Hemmer, H. 1966. Mitteilung iiber ein weiteres Exemplar des Kapl6wen— Panthera leo melano-
chaita (Smith, 1842). Z. Saugetierk. 31: 57-61.

Henpey, Q.B. 1967. A specimen of ‘Archidiskodon’ cf. transvaalensis from the south-western Cape
Province. S. Afr. archaeol. Bull. 22: 53-56.

HEnvbeEy, Q. B. 1968. The Melkbos site: an Upper Pleistocene fossil occurrence in the south-
western Cape Province. Ann. S. Afr. Mus. 52: 89-119.

HENbDEyY, Q. B. 1969. Quaternary vertebrate fossil sites in the south-western Cape Province.
S. Afr. archaeol. Bull. 24: 96-105.

HENpDEY, Q.B. 1970a. A review of the geology and palaeontology of the Plio/Pleistocene deposits
at Langebaanweg, Cape Province. Ann. S. Afr. Mus. 56: 75-117.

HeEnpey, Q.B. 1970). The age of the fossiliferous deposits at Langebaanweg, Cape Province.
Ann. S. Afr. Mus. 56: 119-131.

HENDEY, Q.B. 1972a. The evolution and dispersal of the Monachinae (Mammalia: Pinnipedia).
Ann. S. Afr. Mus. 59: 99-113.

HENbEy, Q. B. 1972b. A Pliocene ursid from South Africa. Ann. S. Afr. Mus. 59: 115-132.

HeEnpDeEy, Q.B. 1972c. Further observations on the age of the mammalian fauna from Lange-
baanweg. Cape Province. Palaeoecol. Afr. 6: 172-175.

HeEnDEy, Q. B. 1973a. Carnivore remains from the Kromdraai australopithecine site. Ann.
Transv. Mus. 28: 99-112.

HeEnbeEy, Q. B. 19736. Fossil occurrences at Langebaanweg, Cape Province. Nature, Lond.
244: 13-14.

HENDEY, OB. & Henpey, H. 1968. New Quaternary fossil sites near Swartklip, Cape Province.
Ann. S. Afr. Mus. 52: 43-73.

HeEnpey, Q.B. & REPENNING, C. A. 1972. A Pliocene phocid from South Africa. Ann. S. Afr.
Mus. 59: 71-98.

— “SSeS 2 OR a

4, BSE

St hr

Ji,

Ps A

366 ANNALS OF THE SOUTH AFRICAN MUSEUM

HENDEY, Q.B. & SinceER, R. 1965. The faunal assemblages from the Gamtoos Valley shelters.
S. Afr. archaeol. Bull. 20: 206-213.

Hooyer, D. A. 1950. A study of subspecific advance in the Quaternary. Evolution 4: 360-361.

Hooyer, D. A. 1972. A late Pliocene rhinoceros from Langebaanweg, Cape Province. Ann. S.
Afr. Mus. 59: 151-191.

Hooyer, D. A. & SINGER, R. 1961. The fossil hippopotamus from Hopefield, South Africa.
Kool. Meded. Leiden 37: 157-165.

Hopwoop, A. T. 1929. New and little known mammals from the Miocene of Africa. Am. Mus.
Novit. 344: 1-9.

Hopwoop, A. T. & HOLLyYFIELD, J. P. 1954. An annotated bibliography of the fossil mammals of
Africa (1742-1950). Fossil Mammals Afr. 8: 1-194.

Howe Lt, A. B. 1944. Speed in animals. New York: Hafner.

Hux ey, T. H. 1880. On the cranial and dental characters of the Canidae. Proc. zool. Soc. Lond.
1880: 238-288.

INsKEEP, R. R. & HENDEy, Q.B. 1966. Actas del V Congreso Panafricano de Prehistoria y de
Estudio del Cuaternario. 2. An interesting association of bones from the Elandsfontein
fossil site. Publnes Mus. arqueol. Santa Cruz de Tenerife 6: 109-124.

Keen, E. N. & Sincer, R. 1956. The fossil Suidae from Hopefield. Ann. S. Afr. Mus. 42: 350-360.

KENSLEY, B. F. 1972. Pliocene invertebrates from Langebaanweg, Cape Province. Ann. S. Afr.
Mus. 60: 173-190.

Kino, J. E. 1964. Seals of the world. London: British Museum (Natural History).

Kine, L. C. 1951. South African scenery. Edinburgh: Oliver & Boyd.

Kuen, R. G. 1972. The Late Quaternary mammalian fauna of Nelson Bay Cave (Cape
Province, South Africa): Its implications for megafaunal extinctions and environmental
and cultural change. Quatern. Res. 2: 135-142.

KurTEN, B. 1953. On the variation and population dynamics of fossil and recent mammal
populations. Acta zool. fenn. 76: 1-122.

KurtTEn, B. 1954. The type collection of Ictitherium robustum (Gervais, ex Nordmann) and the
radiation of the ictitheres. Acta zool. fenn. 86: 1-26.

KurTEN, B. 1956. The status and affinities of Hyaena sinensis Owen and Hyaena ultima Matsumoto.
Am. Mus. Novit. 1764: 1-48.

KurtTEN, B. 19572. Mammal migrations, Cenozoic stratigraphy, and the age of Peking man and
the australopithecines. 7. Paleont. 31: 215-227.

KurtTEN, B. 1957). Percrocuta Kretzoi (Mammalia, Carnivora), a group of Neogene hyenas.
Acta zool. cracov. 2: 375-404.

KurTEN, B. 1957c. The bears and hyenas of the interglacials. Quaternaria 4: 69-81.

KurtTEN, B. 1963a. Return of a lost structure in the evolution of the felid dentition. Commentat.
biol. 26: 1-12.

KurtTEN, B. 19636. Notes on some Pleistocene mammal migrations from the Palaearctic to the
Nearctic. Eiszeitalter Gegenw. 14: 96-103.

KurTEN, B. 1968. Pleistocene mammals of Europe. London: Weidenfeld & Nicolson.

KurTEN, B. 1971. The Age of Mammals. London: Weidenfeld & Nicolson.

LEAkEy, L. S. B. 1965. Olduvai Gorge 1951-196]. 1. Cambridge: University Press.

LEAKEY, L. S. B. 1967. Notes on the mammalian faunas from the Miocene and Pleistocene of
East Africa. Jn: BISHOP, W. W. & CLARK, J. D., eds. Background to evolution in Africa: 7-29.
Chicago: University Press.

Leakey, M. D. 1971. Discovery of postcranial remains of Homo erectus and associated artefacts in
Bed IV at Olduvai Gorge, Tanzania. Nature, Lond. 232: 380-383.

Levyns, M. R. 1962. Past plant migrations in South Africa. Ann. Cape prov. Mus. 2: 7-10.

Linton, D. L. 1969. Evidences of Pleistocene cryonival phenomena in South Africa. Palaeoecol.
Afr. 5: 71-89.

LuNDHOLM, B. 1952. A skull of a Cape lioness (Felis leo melanochaitus H. Smith). Ann. Transv. Mus.
22: 21-24.

LyDEKKER, R. 1884. Indian Tertiary and post-Tertiary Vertebrata. Siwalik and Narbada
Carnivora. Mem. geol. Surv. India Palaeont. indica (10) 2: 178-354.

Mactio, V. J. 1970a. Early Elephantidae of Africa and a tentative correlation of African Plio-
Pleistocene deposits. Nature, Lond. 225: 328-332.

Mactio, V. J. 19705. Four new species of Elephantidae from the Plio-Pleistocene of north-
western Kenya. Breviora 341: 1-43.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 367

Mactio, V. J. 1971. Vertebrate faunas from Kubi Algi, Koobi Fora and Ileret areas, East
Rudolf, Kenya. Nature, Lond. 231: 248-249.

Mactio, V. J. & HENpDEy, Q.B. 1970. New evidence relating to the supposed stegolophodont
ancestry of the Elephantidae. S. Afr. archaeol. Bull. 25: 85-87.

Mason, R. J. 1962. Prehistory of the Transvaal. Johannesburg: Witwatersrand University Press.

MatruHew, W. D. 1929. Critical observations upon Siwalik mammals. Bull. Am. Mus. nat. Hist.
56: 437-560.

Mazak, V. 1964. Preliminary list of the specimens of Panthera leo melanochaitus Ch. H. Smith,
1842, preserved in the museums of the whole world in 1963. <. Saugetierk. 29: 52-58.

MEESTER, J. 1955. Fossil shrews of South Africa. Ann. Transv. Mus. 22: 271-278.

MEEsTER, J. 1965. The origins of the southern African mammal fauna. Zoologica afr. 1: 87-93.

MEEsTER, J. 1971. An additional skull of the extinct Cape lion, Panthera leo melanochaita (H. Smith,
1842) (Mammalia: Carnivora). Ann. Transv. Mus. 27: 27-33.

Morris, D. 1965. The mammals. A guide to the living species. London: Hodder & Stoughton.

Net, J. A. J. 1971. The mammals of Africa: an identification manual. g. Order Pinnipedia. Washing-
ton: Smithsonian Institution.

OAaAKkLey, K. P. 1964. Frameworks for dating fossil man. London: Weidenfeld & Nicolson.

PATTERSON, B. 1965. The fossil elephant shrews (family Macroscelididae). Bull. Mus. comp. Zool.
Harv. 133: 295-335-

PATTERSON, B. 1966. A new locality for early Pleistecene fossils in north-western Kenya. Nature,
Lond. 212: 577-578.

Petrer, G. 1963. Etude de quelques viverridés (Mammiféres, Carnivores) du Pleistocéne
inferieur du Tanganyika (Afrique orientale). Bull. Soc. géol. Fr. 5: 265-274.

Petrer, G. 1967. Problemes actuels de paléontologie: Evolution des vertébrés. Petits carnivores
Villafranchiens du Bed I d’Oldoway (Tanzanie). Collogues int. Cent. natn. Res. scient. 163:
529-538.

PEeTTER, G. 1969. Interpretation evolutive des characteres de la dentures des viverridés africains.
Mammalia 33: 607-625.

Pitcrim, G. E. 1931. Catalogue of the Pontian Carnivora of Europe. London: British Museum (Natural
History).

Pitcrim, G. E. 1932. The fossil Carnivora of India. Mem. geol. Surv. India Palaeont. indica (n.s.)
18: 1-232.

PIvETEAU, J. 1948. Un felidé du Pliocéne de Roussillon. Annls Paléont. 34: 99-124.

Pocock, R. I. 1932. The leopards of Africa. Proc. zool. Soc. Lond. 1932: 543-591.

Pocock, R. I. 1951. Catalogue of the genus Felis. London: British Museum (Natural History).

Poynton, J. C. 1960. Preliminary note on the zoogeography of the Amphibia in southern
Africa. S. Afr. F. Sct. 56: 307-312.

Poynton, J. C. 1964. The biotic divisions of southern Africa, as shown by the Amphibia. Jn:
DAVIS, D. H. S., ed. Ecological studies in southern Africa: 206-218. The Hague: Junk.

Ranp, R. W. 1956. The Cape fur seal Arctocephalus pusillus (Schreber) ; its general characteristics
and moult. Investl Rep. Div. Fish. Un. S. Afr. 21: 1-52.

RAvEN-Hanrrt, R. 1967. Before Van Riebeeck. Cape Town: Struik.

REPENNING, C. A. 1967. Palearctic-Nearctic mammalian dispersal in the late Cenozoic. In:
HOPKINS, D. M., ed. The Bering Land Bridge: 288-311. Stanford: University Press.

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. Antarctic Res. Ser. Washington 18: 1-34.

Roserts, A. 1951. The mammals of South Africa. Johannesburg: Central News Agency.

Romer, A. S. 1966. Vertebrate paleontology. Chicago: University Press.

Ross, G. J. B. 1969. The southern elephant seal, Mirounga leonina, on the South African coasts.
Ann. Cape prov. Mus. 6: 137-139.

Savace, D. E. & Curtis, G. H. 1970. The Villafranchian Stage-Age and its radiometric dating.
Spec. Pap. geol. Soc. Am. 124: 207-231.

ScHEFFER, V. B. 1958. Seals, sea lions and walruses. A review of the Pinnipedia. Stanford: University
Press.

Scnutze, B. R. 1965. Climate of South Africa. 8. General Survey. Pretoria: Government Printer.

ScLaTeR, W. L. 1900. The fauna of South Africa. Mammals. London: Porter.

Scorr, W. B. 1907. A collection of fossil mammals from the coast of Zululand. Rep. geol. Surv.
Natal Zululand 3: 252-262.

ee ae ae

368 ANNALS OF THE SOUTH AFRICAN MUSEUM

SEELEY, H. G. 1891. On Bubalus bainii (Seeley). Geol. Mag. 3: 199-202.

SHORTRIDGE, G. C. 1934. The mammals of South West Africa. London: Heinemann.

Simpson, G. G. 1941. Large Pleistocene felines of North America. Am. Mus. Novit. 1136: 1-27.

Simpson, G. G. 1967. Evolution and geography. Jn: simpson, G. G. The Geography of evolution:
71-131. New York: Capricorn Books.

Simpson, G. G. 1971. Fossil penguin from the Late Cenozoic of South Africa. Science 171: 1144-
1145.

SINGER, R. 1956. The ‘bone tools’ from Hopefield. Am. Anthrop. 58: 1127-1134.

SINGER, R. 1957. Investigations at the Hopefield site. In: CLARK, J. D., ed. Prehistory: Third
Pan-African Congress Livingstone 1955: 175-182. London: Chatto & Windus.

SINGER, R. 1962. Simopithecus from Hopefield, South Africa. Biblthca primatol. 1: 43-70.

SINGER, R. & Bone, E. L. 1960. Modern giraffes and the fossil giraffids of Africa. Ann. S. Afr.
Mus. 45: 375-548.

SINGER, R. & FuLLerR, A. O. 1962. The geology and description of a fossiliferous deposit near
Zwartklip in False Bay. Trans. R. Soc. S. Afr. 34: 205-211.

SINGER, R. & HELTNE, P. G. 1966. Actas del V Congreso Panafricano de Prehistoria y de Estudio
del Cuaternario. 2. Further notes on a bone assemblage from Hopefield, South Africa.
Publnes Mus. arqueol. Santa Cruz de Tenerife 6: 261-264.

SINGER, R. & Hooyer, D. A. 1958. A Stegolophodon from South Africa. Nature, Lond. 182: 101-102.

SINGER, R. & INSKEEP, R. R. 1961. A complete fossil equid skull from Hopefield. S. Afr. archaeol.
Bull. 17: 23.

SINGER, R. & KEEN, E. N. 1955. Fossil suiformes from Hopefield. Ann. S. Afr. Mus. 42: 169-179.

SINGER, R. & Wyner, J. 1968. Archaeological investigations at the Saldanha skull site in South
Africa. S. Afr. archaeol. Bull. 25: 63-74.

SmitH, C. H. 1842. The Naturalist’s library; ed. by W. Jardine. 35. Introduction to Mammalia.
London.

SouTH AFRICAN MusEuM. 1955. The South African Museum, Cape Town, 1855-1955. Cape Town:
The Trustees.

STEPHENS, J. 1959. A new Pliocene cat from Kansas. Pap. Mich. Acad. Sci. 44: 41-46.

SMITHERS, R. H. N. 1968. Preliminary identification manual for African mammals. 25: Carnivora:
Felidae. Washington: Smithsonian Institution.

STROMER, E. 1926. Reste land- und siisswasser bewohnender Wirbeltiere aus den Diamanten-
felden Deutsch-Siidwestafrikas. In: KAISER, E., ed. Die Diamantenwiiste Siidwestafrikas. Berlin:
Reimer.

STROMER, E. 19314. Reste siisswasser- und land bewohnender Wirbeltiere aus den Diamanten-
feldern Klein-Namaqualandes (Siidwest-afrika). Sher. bayer Akad. Wiss. 1931: 17-47.

STROMER, E. 1931). Palaeothentoides africanus, nov. gen., nov. spec., ein erstes Beuteltier aus
Afrika. Sber. bayer Akad. Wiss. 1931: 177-190.

SutTcuirFE, A. J. 1970. Spotted hyaena: crusher, gnawer, digester and collector of vones. Nature,
Lond. 2273 1110-1113.

Ta.sot, W. J. 1947. Swartland and Sandveld. Oxford: University Press.

TEDFORD, R. H. 1970. Principles and practices of mammalian geochronology in North America.
Proc. N. Am. Paleont. Conv. 1969: 666-703.

TuHENtus, E. 1966. Zur Stammesgeschichte der Hyanen (Carnivora, Mammalia). <. Sdugetierk.
31: 293-300.

TuENtus, E. 1967. Zur Phylogenie der Feliden (Carnivora, Mamm.). &. zool. Syst. Evolforsch.
5: 129-143.

Topp, N. B. 1966. Metrical and non-metrical variation in the skulls of Gir lions. 7. Bombay
nat. Hist. Soc. 62: 507-520.

ToOERIEN, M. J. 1952. The fossil hyenas of the Makapansgat valley. S. Afr. F. Sci. 48: 293-300.

TuRNBULL, W. D. 1970. Mammalian masticatory apparatus. Fieldiana, Geol. 18: 149-356.

Van Couverinc, J. A. 1972. Radiometric calibration of the European Neogene. Jn: BISHOP,
W. W. & MILLER, J. A., eds. Calibration of hominoid evolution: 247-271. Edinburgh: Scottish
Academic Press.

Van Noten, F. 1967. Inventaria archaeologica africana; ed. by J. Nenquin. SA 1-SA 5. Excavations
at Gordon’s Bay. Tervuren: Musée Royal de l’Afrique Centrale.

VAN ZINDEREN Bakker, E. M. 1967. Upper Pleistocene and Holocene stratigraphy and ecology
on the basis of vegetation changes in sub-Sharan Africa. In: BISHOP, W. W. & CLARK, J. D.,
eds. Background to evolution in Africa: 125-147. Chicago: University Press.

LATE CENOZOIC CARNIVORA OF SOUTH-WESTERN CAPE PROVINCE 369

VAN ZINDEREN BAKKER, E. M. 1969. The ‘arid corridor’ between south-western Africa and the
Horn of Africa. Palaeoecol. Afr. 4: 139-140.

VirRET, J. 1954. Le loess a bances durcis de Saint-Vallier (Drome) et sa faune de mammiféres
Villafranchiens. Nouv. Arch. Mus. Hist. nat. Lyon 4: 1-200.

Versa, E. S. 1971. A new fossil alcelaphine (Artiodactyla: Bovidae) from Swartkrans. Ann.
Transv. Mus. 27: 59-82.

Wa ker, A. 1969. Lower Miocene fossils from Mount Elgon, Uganda. Nature, Lond. 223: 591-
593:

WELLs, L. H. 1959. The nomenclature of South African fossil equids. S. Afr. 7. Sci. 55: 64-66.

WELLs, L. H. 1969. Faunal subdivision of the Quaternary in southern Africa. S. Afr. archaeol.
Bull. 24: 93-95.

WELLs, L. H. 1970. A late Pleistocene faunal assemblage from Driefontein, Cradock District,
C.P. S. Afr. F. Sci. 66: 59-61.

WELts, L. H. & Cooxg, H. B.S. 1956. Fossil Bovidae from the Limeworks Quarry, Makapans-
gat, Potgietersrus. Palaeont. afr. 4: 1-55.

WituraMs, J. G. 1967. A field guide to the national parks of East Africa. London: Collins.

Woop, H. E. et al. 1941. Nomenclature and correlation of the North American continental
Tertiary. Bull. geol. Soc. Am. 52: 1-48.

Youne, C. C. & Liu, P. T. 1948. Notes on a mammalian collection probably from the Yiishé
Series (Pliocene), Yiishé, Shansi, China. Contr. natn. Res. Inst. Geol. Shanghai 8: 273-291.

ZDANSKY, O. 1924. Jungtertiare Carnivoren Chinas. Palaeont. sin. (C) 2: 1-149.

ZumpT, I. F. 1969. Factors influencing rabies outbreaks: The age and breeding cycle of the
yellow mongoose, Cynictis penicillata (G. Cuvier). Jl S. Afr. vet. med. Ass. 40: 319-322.

3
4

SAAS 28! a>

. pee \ Ye

a... ss ae ae

“wae

panne

INSTRUCTIONS TO AUTHORS

Based on

CONFERENCE OF BIOLOGICAL EDITORS, COMMITTEE ON FORM AND STYLE. 1960.
Style manual for biological journals. Washington: American Institute of Biological Sciences.

MANUSCRIPT

To be typewritten, double spaced, with good margins, arranged in the following order:
(1) Heading, consisting of informative but brief title, name(s) of author(s), address(es) of
(authors), number of illustrations (figures, enumerated maps and tables) in the article.
(2) Contents. (3) The main text, divided into principal divisions with major headings; sub-
headings to be used sparingly and enumeration of headings to be avoided. (4) Summary.
(5) Acknowledgements. (6) References, as below.

Figure captions and tables to be on separate sheets.

ILLUSTRATIONS

To be reducible to 12 cm X 18 cm (19 cm including caption). A metric scale to appear
with all photographs.

All illustrations to be termed figures (plates are not printed; half-tones will appear in their
proper place in the text), with arabic numbering; items of composite figures to be designated
by capital letters (A, B, C etc.).

REFERENCES

Harvard system (name and year) to be used: author’s name and year of publication given
in text; full references at the end of the article, arranged alphabetically by names, chronologi-
cally within each name, with suffixes a, b, etc. to the year for more than one paper by the same
author in that year.

For books give title in italics, edition, volume number, place of publication, publisher.
For journal articles give title of article, title of journal in italics (abbreviated according to the

World list of scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses,

volume number, part number (only if independently paged) in parentheses, pagination.

Examples (note capitalization and punctuation)

ButLoucu, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FiscHeR, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. 7. Conch., Paris
88: 100-140.

FiscHER, P.-H., Duvat, M. & Rarry, A. 1933. Etudes surles échanges respiratoires des littorines.
Archs Zool. exp. gén. 74: 627-634.

Koun, A.-J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region
of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320.

Koun, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the
Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: scHULTZE, L.
Koologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Stid-
Afrika. 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

ZOOLOGICAL NOMENCLATURE

To be governed by the rulings of the latest International code of zoological nomenclature issued
by the International Trust for Zoological Nomenclature (particularly articles 22 and 51).
The Harvard system of reference to be used in the synonymy lists, with the full references
incorporated in the list at the end of the article, and not given in contracted form in the synonymy
list.
Example
Scalaria coronata Lamarck, 1816: pl. 451, figs 5 a, 6; Liste: 11. Turton, 1932: 80.

Q. B. Hendey

THE LATE CENOZOIC CARNIVORA
OF THE

SOUTH-WESTERN CAPE PROVINCE

_ sw AR — - a i wo, * bh NP hk, / Se. a NS WEN. = A £7 ane fo)

—T wie

VNVIWWUVDTILIVS SAITAaAVAAaTT CIBRARIFS

NOILNLILSN

STITUTION NOILNLILSNI

SAJIYVYUEIT LIBRARIE:

INSTITUTION

WANN
Qos
SS
STITUTION NOILNLILSNI

Yo
Yl
NVINOSHLIWS

SMITHSONIAN
f

NOILNLILSNI

LIBRARIES

JiuvVYUYdIT LIBRARIES SMITHSONIAN

Salevia a
INSTITUTION

“STITUTION

N¥INOSHLIWS
SMITHSONIAN

qiuvugir

NOILALILSNI

YSTITUTION NOILNILILSNI

INSTITUTION

NVINOSHLINS SHI1YVYdIT LIBRARIES

me

=

=

oO

w~)

zi Ss

ie AN

= “AS

. x
NOILNLILSNI

2) Zz

Lid Ww

_ —

me =

< c

a +

m Oo

un Ps

34iMvyuNRgIT LIBRARIES SMITHSONI

NOILALILSN

NVINOSHLINS S3I1Y¥Vvu9i
a |

INSTITUTION

LIBRARIES SMITHSONIAN

NVINOSHLINS. S31LYVUGIT

SAIYVYGIT LIBRARIES SMITHSONIAN

NOILALILSN!I NVINOSHLIWS

NVINOSHLIWS

LIBRARIES SMITHSONIAN

NOILNLILSNI

NVINOSHLIWS

X

INSTITUTION

SMITHSONIAN

NVINOSHLIWS

LIBRARIES

INSTITUTION

INSTITUTION NOILNLILSNI

SZIYVYSIT LIBRARIE:

NOILNLILSNI NVINOSHLINS

INSTITUTION

NOILALILSNI

NOILALILSNI

LIBRARIES

INSTITUTION NOILALILSNI

S3SIYVYUSIT LIBRARIES

Fi
es
bike. 2
Oe \;
S)
Bw
=
=
YW) 2
INSTITUTION NOILONLILSNI
WY
uJ
[ae
<C
ao
mM
J
SAJIYVYSIT LIBRARIES
ir
a
Pe)
Y>
a
mM
w
INSTITUTION NOILNLILSNI
“ee Ww
Y a
NRO \ Se
AS 2.
NY S
S3IMVYUGIT

NVINOSHLINS S31u¥vuaagl LIBRARIES SMITHSONIAN

= y ”

Re ae.

&@ NX =

2 BS =

XK <

2 ™ :

z =: - AR

SMITHSONIAN

Pa ices

O ZG wo
ke

5 ‘ae

= =

a 5

z ae:

NVINOSHLINS

=

=

=e

Ww

oO

Zz

=)

rd

SMITHSONIAN

LIBRARIES SMITHSONIAN. INSTITUTIO

a VU?
a
a ac
_
e sf A 4
a co (XO Ne
re) _ a
TIC |
= ah ;
Oo — s f be
= 0 ;
= < .
ra ye
S 7 = 7
NVINOSHLIWS Sa IYVUgl
‘DM Bs
fe) hy Fi
=f
= .
_INSTITUTIOF
Ss ,
no
‘a
ion
=
a
oO
=

NVINOSHLINS S3IYVUdIF

INSTITUTION

SMITHSONIAN INSTITUTIO

a

NVINOSHLINS S3INVUa

SMITHSONIAN

LIBRARIES
NOILNLILSNI

NVINOSHLINS S3IYWYE

eo peg > ce ary Bit are wo
= .O eu re AS us & ey
oc. Z ff, i o = WAS fae “a e oe
ac yy, 4 = 4 NS se = B43 G;
fea) j = = . ea) = 7a
my fo O = Ss) = O a
zZ os Zz . af = 5 =<
| NVINOSHLIWS SMITHSONIAN INSTITUTION NOILNLILSNI NVINI
ee Ne & ie Sy A a z= een, ©
es az i O = xX
oe. 7 w — tL w — wo X
F- 5 x 5 £%: E al
> = > = Ui pi > = = ‘
ea ES ae eee yar 2 i: a ae
- 2a .
I Sa fi z oe Z D
RIES SMITHSONIAN INSTITUTION NOILALILSNI NVINOSHLINS sa lyvugiq Oi. BRARIES SMITI
: es = . = = ee =
nS X 4 a ase =f = a
a S A. i 8 Z : 3
S$ \ e WW: : : : :
ae ae Ss = = = =
is i S = fas, > = > = >
oh ee “”) cae Zz 7) Zz ” 2
HSNI_NVINOSHLINS S31yvygIy LIBRARIES SMITHSONIAN INSTITUTION NOILALILSNI_ NVING
is 2 a = us sa
iz) oa na WW a WW Ss
2 ow ce. Pe , i 4)
= : =|
ce yh = : < e 5 tN
so Tk 3 mo. = mn a ae
ey ES __ SMITHSONIAN _ INSTITUTION NOILNLILSNI_ Saluvudi7_ LIBRARI ES SMIqi
iv za
a © Bs oO 2 Oo ots oO
. = “Oo _ ow a w _
eae ze) te Ps) = a 5 £y
Ps : : = : : E bye
. “Gijiy ce = ia c. si - Ge
ag Oo - a = v Z
SNI_NVINOSHLINS S314YVYSIT LIBRARIES INSTITUTION, NOILNLILSNI NYING
aes w z ngs 2 ” z a w z
= eS = as Ws = < ‘ 2 ts
? =f ; Ea cae ae — > aa z
: $G# FSS 2 g :
= 2 Vp = WS 2 F zZ E
a > | = > = > =
ae Ps ” © Le ina ” z 7)
1ES SMITHSONIAN INSTITUTION NOILMLILSNI NVINOSHLIWS (Sa1YV¥I7_ LIBRARIES SMITH
i. a z Z x z
a <” - MH wr. vi n am
ze: “3 : ESN 2 g 2
van “I a QQ N 4 wf,
a = m — N am — jee) eA
. i ‘a O —_— Oo ai O ce
| a ay - eit z ay
SNINVINOSHLIWS | $3 1YVYGi7 LIBRARI ES_ SMITHSONIAN _ INSTITUTION _, NOILA.LILSNI_NVINO
. > ~—~
= S) — S g = O
ow = w = Yf,; w = w
2 Wa 5 : 5 £45 : :
- bea, co > K Lf — xy
= \S" 5 2 - YAP 2 i 2
a Foe, 2 “a n° . es ee e
[9] = ep) — w oe
IES SMITHSONIAN INSTITUTION NOMOLIISNI_NVINOSHIINS, SAluvugi7 LIBRARIES SMITH
. = ore w x
. = < = < = Wy), < =
a > ae > a = =
\ = 3 = = s GUY. z =
DB a ® LD) n fa O
\ ce 3 : S) bp Ap Ye = 9° Se 7: x s S
: a = = = = = “SN >
2 wn 2 7) 2 ” =
SNI_NVINOSHLIWS LIBRARIES SMITHSONIAN INSTITUTION ee Ne
= am e
& " % tw & Ww O
> a = oe . a - =
~ a , ~—
oO a a = « =
. 5 ae S = 3
4 = oe) Zz acy =

LICOVAILLA AIL

{AIC TITTITICNAL

WA