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Page
AverY, D. M.
Micromammals as palaeoenvironmental indicators and an interpretation of the Late
Quaternary in the southern Cape Province, South Africa. (Published January
ee a en ee Tee ee AS sie Sc.kn Qe ne PEG b a APT es beds wale 183

DINGLE, R. V.
The Campanian and Maastrichtian Ostracoda of south-east Africa. (Published
eee SPM ee eye neg aivghs Se yt es he M's Et pes oa ote ne oe 1

NEW GENERIC NAMES PROPOSED IN THIS VOLUME

Dutoitella Dingle, 1981 ......
Klingerella Dingle, 1981 .....

Ponticulocythere Dingle, 1981

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oo A a ea,

VOLUME 85 PART 1 OCTOBER 1981 ISSN 0303-2515

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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

FiscHer, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.

Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
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THE CAMPANIAN AND MAASTRICHTIAN
OSTRACODA OF SOUTH-EAST AFRICA

By

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THE CAMPANIAN AND MAASTRICHTIAN OSTRACODA OF SOUTH-
EAST AFRICA

By

R. V. DINGLE
Department of Geology, University of Cape Town

(With 81 figures and 20 tables)
[MS. accepted 6 November 1980]

ABSTRACT

78 species, representing 46 genera of ostracoda are recorded from the Campanian and
Maastrichtian strata of south-east Africa, where rocks of this age have been sampled in the
Natal—Zululand Basin, on the Transkei Swell, and in the Outeniqua Basin (on the Agulhas
Bank). 35 of the species are new, 34 have previously been described from south-east Africa, 1
previously described from Australia, and 8 are left in open nomenclature. 1 new subfamily
(Unicapellinae), and 3 new genera (Klingerella, Ponticulocythere, and Dutoitella) are erected.
Of the 35 new species, the following 23 are formally described: Cytherelloidea mfoloziensis,
Platella africana, Pontocyprella nibelaensis, Pariceratina hirsuta, Eucytherura? pyramidatus,
Cytheropteron brenneri, Pedicythere fragilis, Apateloschizocythere mclachlani, Amphicytherura
armatus, Klingerella aranearius, Krithe nibelaensis, Pondoina igodaensis, Xestoleberis luciaensis ,
Ponticulocythere biremis, Dutoitella dutoiti, D. mimica, Haughtonileberis nibelaensis, Oertliella
maastrichtia, Hermanites? arcus, Parvacythereis monziensis, Curfsina monziensis, Cativella?
dubia, and Australileberis stangerensis.

Population analyses on the faunas from Zululand lead to the recognition of six distinctive
ostracod assemblages, that in turn can be related to various palaeosedimentary environments.
All were considered to have been low-energy, open-water, normal marine situations. Further
consideration of these data, including the use of a Cytheracea—Cytherellidae—Bairdiacea/
Cypridacea triangular diagram (CCBC plot), allows a reconstruction of the Campanian-—
Maastrichtian palaeogeography of Zululand, and the recognition of local late Cretaceous
sea-level fluctuations. A comparison with other Gondwanide localities reveals a relatively close
relationship at the generic level between south-east Africa and western Australia and east
Africa, and only weak links with west Africa and South America. Finally, an ostracod zonal
scheme is proposed, based on the Zululand faunas.

CONTENTS

PAGE
a It ee is ae doe Gk bceed neki ahdoesvnaw ep Zz
Regional stratigraphy and sampling localities..................... 3
PSP TIMING OO SCHIPIONS,. ... oc De eet ccs cere have essn eats 12
UBT Meng 131
TONS sc PEs gs PEGs Ve ahd od 4d cade ge wae 132
Me MR Re ae bes Shi psa sv an SATA ae SD 132
BH-9 borehole and Zululand outcrops .............. 132
MOR ONS Sin bbe Mics on coe whe Pe apd SOs een PH 146

Campanian-—Maastrichtian palaeogeography of
southern Natal—Zululand Basin................. 149
ESS oS cn ok eee eee ae 152
Pe NE Oe Le it qa Oe te a aa wiht gha me fat Slaw 153
BES A CANT Ca EA ees) Se nee ogee ee gn ee ee 154
SUSANNE PERG, Ee cae a ae 8 Go that ors uw cxie are BP oe 156

1

Ann. S. Afr. Mus. 85 (1), 1981: 1-181, 81 figs, 20 tables.

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

Southern area: Aguilas Banker ta Gap ise cd eens 156
SRNR os sas ewes ck ee wee ee aan pees gin edaene gl ae 159
ByGstratigrapny 5. < cAues » kaso es wih Does car sane nes ete 160
SOMES SE UTICA id el orate wiawiran eRe Rane an Pee Roan eens 160
Phylogeny—species distribution (appearances) ....... 160
Osteace@ ronal scheme 3.6608 on eee ode es pasien dent 163
Phyloseny=—hisher tata. ey BME oleh ee beak 4 166
Phylogeny—species distribution (extinctions)......... 170
Comparison with other Gondwanide localities............ 170
Wester AUStraltal oc caters «nit Grn e ye Oe ae 12
POS PUI oer e inte afta anaes > pit th Oakes tema eee 174
WEST NIRA fice. ahead x, cad re ore ae oe cota 174
OMEN AMMOINGA Sy. oo 5. GO tereea aks oe eee eee COR eee 17s
INTRODUCTION

The purpose of this contribution is to document and discuss as fully as
possible the taxonomy, palaeoecology and stratigraphic distribution of the
Campanian and Maastrichtian ostracod faunas of south-east Africa.

Marine sediments of this age in south-east Africa occur in two large
continental margin basins (Outeniqua and Natal—Zululand) which are separated
by the generally positive Transkei Swell region on which rest small, thin patches
of marine Mesozoic strata (Dingle 1978) (Fig. 1). The St Johns Basin on the
narrow continental shelf off the Transkei coast may contain uppermost Cre-
taceous sediments. Campanian—Maastrichtian times were marked by relatively
high sea-levels as the Upper Cretaceous marine transgression reached its peak,
with the result that whilst the basinal facies are merely the youngest strata in a
thick Mesozoic pile, the thin deposits over the Transkei Swell rest directly on
pre-Mesozoic basement (Figs 1, 3).

The first studies on Campanian—Maastrichtian marine ostracods from south-
east Africa were made by Chapman (1916) when he described the microfauna of
the late Campanian/early Maastrichtian limestones from the ‘Lower’ quarry at
Needs Camp. (There is no evidence to indicate that in his 1904 or 1923 papers
Chapman sampled horizons as high as Campanian at the Umzamba cliff sections
in Transkei.) Chapman (1916) recorded three species of ostracod from Needs
Camp, and since then the only other descriptions of ostracods of this age from
south-east Africa were by Dingle (1971, 1980) on Maastrichtian from the
Outeniqua Basin, and Santonian—Campanian from the Richards Bay BH-9
borehole, Zululand, respectively.

The present study, together with a reassessment of the earlier work, has
resulted in the recognition of 78 species of ostracods in the Campanian and
Maastrichtian sediments of south-east Africa (Table 1). These have been allo-
cated to 46 genera, with 8 types in open nomenclature. In the Natal—Zululand
Basin, where ostracods are abundant, the faunas can be grouped into 4 distinct
assemblages, 2 of which can be further subdivided. This allows the recognition
of 6 palaeosedimentary environments which are shown to have alternated during
Campanian—Maastrichtian time. Sparser faunas from the other areas are com-

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 3

NATAL-
ZULULAND
BASIN

“TUGELA
CONE

ST JOHNS
BASIN

Port
Elizabeth

basins

- 2kmisobath

Fig. 1. Distribution of Campanian—Maastrichtian sediments in south-east Africa (shaded).
Numbers refer to sampling localities: 1l—sample 818, Agulhas Bank; 2—Igoda estuary;
3—Lower or East Quarry, Needs Camp; 4—Umzamba Cliff; 5—JC-1 borehole; 6—BH-9
borehole, Richards Bay; 7—Monzi and Mfolozi River outcrops (Kennedy & Klinger 1975
localities 20 & 21); 8—Nibela Peninsula (Kennedy & Klinger 1975 localities 110 & 113).

pared with the better-known ostracod assemblages from Natal—Zululand, and
suggestions for their palaeoenvironments are advanced.

The temporal distribution of the various ostracod species in Zululand,
Richards Bay BH-9 borehole, and the Agulhas Bank is shown in Table 2 and is
related to the ammonite zonation of Kennedy & Klinger (1975). A consideration
of the age ranges of some of the diagnostic, and less environmentally-bound
species, leads to a proposed ostracod zonation scheme for the Zululand Cam-
panian—Maastrichtian succession. 4 zones and 6 subzones (a total of 8 separate
periods) are recognized.

REGIONAL STRATIGRAPHY AND SAMPLING LOCALITIES

Figures 1-2 show the approximate extent of the Campanian and Maastrich-
tian sediments in south-east Africa and the localities from which samples were
available for study. Measured sections and sampled horizons are shown in
Figure 3, and where they coincide with localities examined by Kennedy &
Klinger (1975), the original notation system has been retained to allow correla-

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE |

Geographical distribution of ostracods in Campanian and Maastrichtian strata of
south-east Africa

Zulu- Um- Needs Agulhas
land BH-9 JC-\ zamba Igoda Camp Bank

Cytherella sp... ; ; x x x x x
Cytherelloidea contorta . A : x x
C. umzambaensis : , ‘ : x x
C. griesbachi é : se
C. mfoloziensis . : ' . . <
Platella africana : : , x 4
Bairdoppilata andersoni . ; : x x Vx x * x
B. andersoni aequalis x
B. africana . . : x
B. cf. africana. ‘ A : : x
Bosp. A” 3 : : : x
Bythocypris richardsbayensis ‘ : x x x x
BT sp. . . : : ; : me
Paracypris eee ; x x x

P. zululandensis . ; : : : x x x
P.sp.A : : : ‘ : : Xx
(ape) ae 2 : q x
Pontoc) prelia Ge ee ‘ x
PISpe . ; : : : : x
Pariceratina ener i xX
Eucytherura? pyramidatus x
Cytheropteron brenneri x
C. cf. westaustraliense <
Pedicythere fragilis x
Apateloschizocythere iachiens ‘ x
A. laminata ' : : x
Amphicytherura jae : : : ~ x

A. zululandensis ; : : : 54
- armatus . : 3 ; : ‘ x
A.sp. A : : : x
Klingerella aranearius : 3 ' x
Hutsonia? sp... : : : : x
Krithe nibelaensis : 3 : : x x
K. sp. A : : : ; : Xx
Pondoina igodaensis : x
Xestoleberis luciaensis : ‘ : x x x
Buntonia? sp. : x
Brachycythere sicarius x %
B. longicaudata * < x x
Pterygocythere aoa ~
Ponticulocythere biremis ; : x
Agulhasina quadrata . é : ‘ x
Unicapella sacsi ; : } : * * x
U. reticulata : : : ‘ : x
Dutoitella dutoiti ' : : : x
D.mimica . : : x x
Haughtonileberis haughtoni ; , x
H. vanhoepeni_. : ; 5 : x x
H. fissilis ; : : x x x
H.nibelaensis . : : : ; x x
Rayneria nealei x

CAMPANIAN AND MAASTRICHTIAN OSTRACODA a

85(1) Zulu- Uim- Needs Agulhas
land BH-9 JC-\1 zamba Igoda Camp Bank

Cythereis klingeri , 4 x *
C. transkeiensis . ; : 3 : x x
Oertliella sp. A. ‘ ; . : x
O. pennata . ; : : : ; x
O. africana : : : : : x *
O.maastrichtia . : x
Gibberleberis ie ; ; ; x
G. sp. A : : : : r x
Trachyleberis minima ; ; x
T. zululandensis . ‘ : . 4 x
T. schizospinosa 3 ; ; x
Hermanites kennedyi : : x < Me
H2arcus . : : : x
Hf? cf. arcus : ; : x x
Parvacythereis monziensis 4 : x
P. spinosa . ; y ; ; : x
Curfsina monziensis . s : ; x
Cativella? dubia : : ; : x
Phacorhabdotus? anomala : x x
P?sp. A : F : x
Australileberis pmamerersts ; ; x
Paraplatycosta reticulata . 3 x
Indet. sp. 1 x
Indet. sp. 2 x
Indet. sp. 3 x
Indet. sp. 4 ~
Indet. sp. 5 x
Indet. sp. 6 *
Indet. sp. 7 : : : : x
Imget.sp. . ; : ; ‘ : x

Totals : : : ‘ 44 28 it 5 15 6 es

78 species, 46 genera

tion with their ammonite zonation. These workers recognized eight subdivisions
in the Campanian—Maastrichtian of Zululand based on ammonite assemblages,
and although these were intended to be provisional upon the establishment of a
more sophisticated scheme, details of their zonation are given in Table 3 to
allow the extension of our ostracod ranges outside the south-east African region.

OUTENIQUA BASIN (locality 1 on Figs 1, 3).

A single sea-floor sample from the Alphard Formation consisting of light
Olive, clayey sand containing abundant glauconite grains, pyrite and shell
fragments, and occasional fish teeth. Position: 35°20,0'S 23°17,0’E from a
sea-floor depth of 1 203 m. The rich and well-preserved microfauna of this
sample was originally described by Dingle (1971). It contains, amongst others,
the following planktonic foraminifera: Gublerina reniformis (Marie), Rugoglo-
bigerina rugosa (Plummer), R. rugosa rotundata Brénnimann, Globotruncana
arca (Cushman), G. stuarti (de Lapparent), and G. aegyptiaca Nakkady, which
indicate a Middle-Upper Maastrichtian age (Postuma 1971). This is corrobo-

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x
x

87.0
88.0

xX Xo ces

GPS

me PS PS 2S OS

x

Cll

89.0

BH-9

TABLE 2

92.3

Xe OG OOK, BS

x
x

Dies)
100.0
102.2
106.0

110.0

Distribution of ostracod species in samples from the Campanian and

x

DR OR PK SEXO: ew

x

Maastrichtian of Zululand (outcrops at Monzi, Mfolozi River, and Nibela,

Cl

and borehole BH-9 at Richards Bay) and Agulhas Bank (sample 818).

OSES OSES PSS OEE

xX X X

x

Pe OC oe EO NSS

xXx X

x

SANTONIAN

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

EAST LONDON

etagieh Si NIBELA
<) PENINSULA

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Fig. 2. Main map: details of sampling localities in Zululand. BH—9 is locality 6 in Figures 1 and

3; 20 and 21 are locality 7 in Figures 1 and 3; and 110 and 113 are locality 8 in Figures 1 and 3.

Geology is after Kennedy & Klinger (1975). Insert: localities in the vicinity of East London.

Lower Quarry Needs Camp, and Igoda estuary are localities 3 and 2, respectively, in Figures 1
and 3.

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s1oquinu AjITRIO] UTeYL “ROLY 3Sed-YINOS UT Sar}TTedO] UeNYOIseepPy—ueiueduey pajdwies wioI sassauyory] sJoys1og puke suUOT}IOaS poInsKap, “¢ “BLY

ajdwes =

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1D ooo telaag th

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CAMPANIAN AND MAASTRICHTIAN OSTRACODA

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10 ANNALS OF THE SOUTH AFRICAN MUSEUM

rated by McLachlan & McMillan’s (1979) recognition of the R. rugosa and G.
arca zones on the Agulhas Bank, and the G. stuarti zone in Zululand.

A sample of calcareous sandstone containing Eubaculites latecarinatus,
Gunnarites sp. cf. G. kalika, inoceramids, and other indeterminate bivalves has
been dredged from Alphard Formation outcrops at 34°08,25’S 25°10,45’E on the
Agulhas Bank (Sample 1336; Dingle 1973; Klinger et al. 1980). Klinger et al.
(1980), who described the molluscan fauna, have dated this sample as Maastrich-
tian I. No ostracods have been recovered from the lithified matrix.

EASTERN CAPE (localities 2-3 on Figs 1, 3)

Small outcrops of late Campanian/early Maastrichtian limestones occur at
Igoda (locality 2) and Needs Camp (Lower Quarry) (locality 3).

At Igoda, on the coast (Fig. 2), Klinger & Lock (1978) have described 20 m
of glauconitic arenaceous limestones and calcareous sandstones with a basal
conglomerate resting on Permo-—Triassic Beaufort Group. There is a meagre and
poorly preserved invertebrate macrofauna with Baculites subanceps Haughton,
Eupachydiscus? sp. and Saghalinites sp. cf. S. cala (Forbes) that indicates a late
Campanian/early Maastrichtian age.

The Lower Quarry at Needs Camp lies about 13 km inland from Igoda (Fig.
2), and consists of friable, polyzoa-rich limestone with poorly preserved micro-
and macrofauna. On the basis of the common presence of the brachiopod
Eolacazella affine (Bosquet), Klinger & Lock (1978) consider the Lower Quarry
and Igoda strata to be the same age, a view corroborated by Siesser & Miles
(1979: 148) using calcareous nannofossils. McGowran & Moore (1971) had
previously suggested a Campanian to Maastrichtian age for the Needs Camp
strata on the presence of two specimens of Rugoglobigerina aff. R. rugosa
(Plummer). Chapman’s (1916) type material from Needs Camp (South African
Museum slide SAM-2736) has been examined and the specimens re-illustrated
herein.

Limestones from both Igoda and Needs Camp are recrystallized, so that the
preservation of microfossils is generally poor.

UMZAMBA (locality 4 on Figs 1, 3)

Numerous publications have described the succession and fauna of the
coastal cliffs around the Umzamba River mouth in northern Transkei, the most
comprehensive and up to date for our purposes being Klinger & Kennedy (1977,
1980), and Makrides (1979). The Campanian I-Santonian III boundary occurs in
the cliff section between beds 7 and 8 (within bed 7 according to Makrides 1979)
and the section extends into Campanian II (with Baculites sulcatus). Chapman
(1904, 1923) and Dingle (1969) have described the ostracods from the basal
(Santonian) part of the Umzamba succession, but no work has been published
on the Campanian ostracod faunas. The Campanian part of the Umzamba cliffs
is badly decalcified and only two samples from this section contained ostracods.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 11

OFF-SHORE NATAL, TUGELA CONE (locality 5 on Figs 1, 3)

This is the SOEKOR borehole JC—1 on the continental shelf at 29°27,69'S
31°35,66’E in 72 m of water. The borehole penetrated about 2 300 m of Tertiary
and Upper Cretaceous strata before entering Palaeozoic quartzites, and the
section between 1 935 m and 1 560 m was reckoned to be Campanian (94 m) and
Maastrichtian (281 m) in age (Du Toit & Leith 1974). Eighteen samples
containing ostracods were available for study.

ZULULAND (localities 6-8 on Figs 1, 3)

This is the area from which Kennedy & Klinger (1975 et seq.) have
collected extensively during the course of their revision of the Cretaceous
ammonite faunas of south-east Africa. Here, the Campanian—Maastrichtian
strata constitute the upper part of the St Lucia Formation. The southernmost
section is in the Richards Bay borehole (BH-9) (28°48,65'S 31°57,75’E, Fig. 2,
locality 6 on Figs 1, 3) where approximately 50 m of Campanian I and II overlie
45 m of Santonian which in turn rests on granitic basement. The ammonite
stratigraphy of BH—9 has been described by Klinger & Kennedy (1977) and the
ostracods by Dingle (1980), who examined ten samples in the 32 m of core
available to him from the Campanian I and II. The excellent preservation of the
fossils and completeness of the sequence allow BH—9 to be used as a standard
for biostratigraphic and palaeoecological comparisons in Zululand and else-

where.
TABLE 3

Kennedy & Klinger’s (1975) ammonite subdivision of the Campanian and
Maastrichtian stages in Zululand.

MAASTRICHTIAN

MAASTRICHTIAN I. No ammonites present. Inoceramid debris is abundant.

MAASTRICHTIAN II. Coarsely ornamented baculitids of the Eubaculites ootacodensis type are
abundant. Pachydiscids are also present.

MAASTRICHTIAN I. Feebly ornamented to smooth Eubaculites are common. Other ammonites
include Saghalinites sp., Pachydiscus (Neodesmoceras), Menuites, ‘Epiphylloceras’ and
Hoploscaphites. The local base is drawn below the appearance of abundant Eubaculites.

CAMPANIAN

CAMPANIAN V. Giant Bostrychoceras are abundant, with scarcer Saghalinites and compressed
pachydiscids.

CAMPANIAN IV. Saghalinites cala and Pachydiscus (P.) sp. are common. Other ammonites include
Gunnarites antarcticus, Nostoceras? sp., Pachydiscus (Neodesmoceras) sp.

CAMPANIAN III. Faunas are sparse but distinctive. A feebly nodose Baculites is abundant, and
giant (1 m) pachydiscids (probably Eupachydiscus) are very common.

CAMPANIAN II. Menabites (Australiella) is abundant in the lower part of this division but species
including M. (A.) australis and M. (A.) besairei, together with Bevahites spp range through-
out. Baculites sulcatus is abundant throughout, whilst pachydiscids become common in the
higher parts: e.g. Anapachydiscus subdulmensis, A. wittekindi, A. arrialoorensis and Pachy-
discus manambolensis. Other ammonites include Hoplitoplacenticeras plasticum plasticum,
Maorites sp., Neogaudryceras sp., Gaudryceras sp. and Bostrychoceras sp.

CAMPANIAN I. Submortoniceras woodsi (Spath) and related forms are common; other ammonites
include Bevahites spp. and Menabites spp, Hauericeras gardeni, Pseudoschloenbachia,
Bostrychoceras spp, Vendegiesiella sp. cf. spinosa, V. trituberculata, Karapadites, and
diplomoceratids. The local base is drawn below the level of abundant Submortoniceras.

12 ANNALS OF THE SOUTH AFRICAN MUSEUM

Upper Campanian II to Campanian IV sections are exposed in extensive
cliffs along the southern part of the Nibela Peninsula in the St Lucia Game Park
(locality 8 on Figs 1, 3; sites 110, 113 on Fig 2). Upper Campanian II to
Campanian III material was collected at Kennedy & Klinger’s (1975) locality 110
(27°59,17'S 32°24,57'E), whilst the Campanian IV was sampled at their locality
113 (27°58,20'S 32°26,95'E). Farther south, upper Campanian to Maastrichtian
sections are exposed in the vicinity of the village of Monzi (locality 7 on Figs 1,
3). Campanian V was sampled in a road cutting to the north of the village
(Kennedy & Klinger’s (1975) locality 21, 28°25,00'S 32°18,58’E) (Fig. 2), whilst
Maastrichtian I and II occurs in a complete sequence in low cliffs on the north
bank of the Mfolozi River, and in a small quarry 200 m to the east (Kennedy &
Klinger’s (1975) locality 20, 28°26,98’S 32°16,60’E) (Fig. 2). Although the upper
parts of some sections are heavily decalcified, the preservation of microfossils in
the unweathered sections is generally moderate to good.

A total of 53 fossiliferous samples (including Chapman’s (1916) types from
Needs Camp) were available for study. Microfossils were extracted by washing,
and photographed with a Cambridge 180 Stereoscan. Specimens were mounted
on aluminium stubs using double-sided Sellotape and were coated with a gold
palladium mixture. Types and illustrated material are deposited in the South
African Museum, Cape Town.

SYSTEMATIC DESCRIPTIONS

The classification used here is based mostly on the Ostracod Treatise
(Moore 1961), with various additions necessitated by recent work. Morphological
terms have been supplemented by those introduced to cover features visible at
high magnifications (e.g. Sylvester-Bradley & Benson 1971). In addition to
taxonomic notes, age range, geographic distribution and palaeoecology for each
species are discussed, with ecological preferences expressed in terms of ostracod
assemblages 1 to 7, and the sedimentary environments that they are thought to
represent. These assemblages are defined in the discussion section.

Abbreviations: RV = right valve, LV = left valve, MPC = marginal pore
canals, SCT = subcentral tubercle, ATE = anterior terminal element.
PTE = posterior terminal element, ME = median element, AM = anterior
margin, PM = posterior margin, DM = dorsal margin, VM = ventral margin, é
NPC = normal pore canals, and MA = marginal areas.

Subclass OSTRACODA Latreille, 1806
Order PODOCOPIDA Miller, 1894
Suborder PLATYCOPINA Sars, 1866
Family Cytherellidae Sars, 1866

Members of this family constitute one of the main elements of the Cam-
panian—Maastrichtian ostracod faunas of south-east Africa, particularly in the

13

CAMPANIAN AND MAASTRICHTIAN OSTRACODA

UONRIAIP Plepueys BdIe] — 4 JUSWIUOIIAUD PotIajoId — + yussoid — *

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Og OT* CT* Cx L* CT# : eset) Yo Het) Seni, S149GI] O]SAX
sok otk rf SHIDLUAD “py

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ek * * SISUAIZUOU SIAAAYJAIDAADG

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ok * * ** # DIJYIIAISDDUL *C

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eaoepiidaéy

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(BpOdvsjSO ]e10} 94 UBB) SIsUadvgspadyoiA stAdAI0YIAg

eooripsreg

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poqo1ysa
i
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p dave

(soujaw) syidap 19}eM poyeUiysa

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

areas of Zululand north of the Richards Bay BH-9 borehole. In the Campanian
I, cytherellid populations make up 10-20 per cent of the total fauna and this
rises in Campanian II: 20-35 per cent (Richards Bay borehole) to ~20 per cent
(at outcrop farther north). Campanian III ostracod faunas are also about 20 per
cent cytherellid, but this increases rapidly in Campanian IV, V, and lower
Maastrichtian I to reach levels up to 50 per cent, falling to 20-30 per cent in
Maastrichtian II. As will be noted in the discussion section, these fluctuations
are directly related to changes in the palaeosedimentary environments.

assembl.

locality
ostr.

Tc
a CYTHERELLA

morphotypes

80 120 160

Fig. 4. Distribution of Cytherella sp. in Campanian—Maastrichtian strata of Zululand and
Agulhas Bank. A. Total number of valves. B. Percentage of total ostracod population.
C. Morphotypes 1-4 as percentage of total Cytherella sp. plotted by ostracod assemblage.

The family is represented in the Campanian—Maastrichtian by 3 genera:
Cytherella, Cytherelloidea, and Platella with 5 designated species and 4 mor-
photypes. 2 of the species and all 4 morphotypes are inherited from the
Santonian, whilst 1 species is restricted to the Campanian (Cytherelloidea
griesbachi), and 2 are restricted to the Campanian—Maastrichtian (Cytherelloidea
mfoloziensis and Platella africana).

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 15

Genus Cytherella Jones, 1849
Cytherella sp.
Fig. SA-F
Cytherella sp. 1, 2, 3, 4, Dingle, 1980: 5-7, fig. 2A-F.

Remarks

Dingle (1980) recognized four morphotypes of Cytherella in the Richards
Bay BH-9 borehole and it was hoped that with the addition of more material
formal descriptions of these might be possible. Whilst all four have been
recognized in younger samples, and estimates of their relative importance have
been made (Fig. 4C), it has not been possible confidently and consistently to
discriminate to the extent of formally describing new species. ‘End members’ of
each morphotype can easily be recognized but within any population there is
usually a considerable residue of intermediate forms. For the purposes of
population counts, these can be placed in categories, but the decision is
sometimes arbitrary. Consequently no formal subdivision within the genus
Cytherella is yet possible.

Age, distribution, palaeoecology

Santonian II to Maastrichtian III (Zululand outcrops, BH-9 Richards Bay,
Igoda, JC-1 borehole, Agulhas Bank). Although Cytherella sp. occurs in ostra-
cod assemblages 1-3, 4a, 4b, 5a, 5b, 6-7, it is only consistently present in
numbers above trace in 4~7, reaching a maximum (>40% total ostracod popula-
tion) in 6 (Fig. 4): deep (>500 m, outer shelf/upper slope) oceanographically
unstable environment.

In terms of the four morphotypes recognized by Dingle (1980), Figure 4C
shows mean percentage of total Cytherella plotted against the various ostracod
assemblages. Several points emerge:

(i) Morphotype 2 (ovate form) dominates throughout (48-84%) but is most
prominent in assemblages 6, 7, and 5a (>60%).
(ii) Morphotypes 3 (elongate, parallel sided form) and 4 (large plump form) in
combination are most common in assemblages 4a and 4b.
(iii) Morphotype 1 (asymmetric form) is consistently most abundant in the
ostracod assemblages 5a and 5b.

Based on their preferred habitat, therefore, it is possible to recognize the
following approximate groupings: 100-200 m—morphotypes 3 and 4;
200-500 m—morphotype 1; >500 m—morphotype 2. :

Cytherella sp. occurs in small numbers (dominantly carapaces) in the
Maastrichtian part of borehole JC—1 (between levels 1 835 and 1 652 m) (Table
10). Morphotypes 2 and 3 have been recognized, but because of their scarcity no
environmental significance can be placed on their presence. It is probably
significant, however, that Cytherella sp. is virtually absent from samples in which
charophytes has been found, and this is thought to reflect the species’ intoler-

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 5. Cytherella. A. Cytherella sp. morphotype 1, SAM-—K5568, BH9 82,03 m, RV,
Campanian II. B. Cytherella sp. morphotype 2, SAM-—K5569, BH9 82,03 m, RV, Campanian
II. C. Cytherella sp. morphotype 3, SAM-K5570, BH9 82,03 m, RV, Campanian II.
D. Cytherella sp. morphotype 4, SAM-—K5571, BH9 82,03 m, RV, Campanian II. E. Cytherella
sp. morphotype 3, SAM—K5662, JC-1 1811 m, LV, Maastrichtian. F. Cytherella sp. morpho-
type 2, SAM-K5663, JC-1 1835 m, RV, Maastrichtian.
Scale bars all 100y.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA ly

ance to large influxes of freshwater-derived detritus. A similar antipathetic
relationship has been noted for Bythocypris? sp. in borehole JC-1.

Genus Cytherelloidea Alexander, 1929

This is an important subsidiary genus in some of the Upper Cretaceous
rocks of south-east Africa. It is recorded sporadically throughout the Santonian
II to Maastrichtian II of Zululand and Richards Bay BH-9 borehole where it
generally constitutes 1-5 per cent of the total ostracod population, but has not
been found in the following areas: the Maastrichtian III of the Agulhas Bank;
the late Campanian/early Maastrichtian of Igoda and Needs Camp; Campanian—
Maastrichtian of the JC—1 borehole.

In the Campanian—Maastrichtian sequences of Zululand, Cytherelloidea is
represented by four species: C. umzambaensis, C. contorta, C. griesbachi, and

ostr.
assembl.

& CYTHERELLOIDEA PLATELLA

2
a
=
o

2
E
N
E
3

uo

contorta
griesbachi
africana

mfoloziensis

CAMPANIAN

Fig. 6. Distribution of Cytherelloidea and Platella in Campanian—Maastrichtian strata of

Zululand and Agulhas Bank. A. Total number of valves of C. umzambaensis. B. Total

number of valves of C. contorta. C. Total number of valves of C. griesbachi. D. Total number

of valves of C. mfoloziensis. E. C. mfoloziensis as percentage of total Cytherellidae. F. Total

valves of genus Cytherelloidea as percentage of Cytherellidae (dash line, bottom scale) and

percentage total ostracods (solid line, top scale) plotted by ostracod assemblage. G. Total
number of valves of P. africana.

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

C. mfoloziensis. The genus is always numerically subordinate to its relation
Cytherella, but reaches its maximum importance in terms of percentage of the
family Cytherellidae in ostracod assemblage 4a (28%), whilst constituting <10
per cent in assemblages 5a, 5b and 6 (Fig. 6). These data indicate that in
south-east Africa Cytherelloidea is generally more tolerant of shallower water
conditions than is Cytherella, although its preferred environment is also deep
water (>500 m, outer shelf/upper slope), where its mean percentage of the total
ostracod population reaches 3-4 per cent, compared to 1-2 per cent in the
shallower areas.

Cytherelloidea umzambaensis Dingle, 1969
Figs 7A, 9C

?Cytherella williamsoniana Jones, 1849, Chapman, 1904; 236.
Cytherelloidea umzambaensis Dingle, 1969: 351-353, fig. 3. Dingle 1980: 7, figs 3A, 4A-B.

Remarks

The range of C. umzambaensis overlaps with that of C. mfoloziensis in
Campanian IV and it would seem that the latter evolved from the former under
the stress of the establishment of deep-water environments in lower Campanian
IV times. In late Campanian IV, C. umzambaensis declines in importance
before it is finally replaced (Fig. 6).

Age, distribution, palaeoecology

Santonian II to Campanian IV (Umzamba, Richards Bay BH-9 borehole,
and Nibela Peninsula). C. umzambaensis is an environmentally tolerant species
that occurs sporadically and in relatively small numbers in the Richards Bay
BH-9 borehole and at outcrops in the Nibela Peninsula. It has been found in
ostracod assemblages 1-3, 4a, 4b, 6-7 but preferred the moderate depth en-
vironments represented by assemblages 4a, 4b and Sa: quiet, 100-300 m, inner-
outer shelf environments (Table 4).

Cytherelloidea contorta Dingle, 1980
Fig. 7B
Cytherelloidea contorta Dingle, 1980: 11-12, figs 3D, 4E.

Remarks

This rare species has been recorded from single horizons in the Richards
Bay BH-9 borehole and at outcrop in the Nibela Peninsula (Fig. 6B).

Age, distribution, palaeoecology

Campanian I and Campanian II (Richards Bay BH-9 borehole and Nibela
Peninsula). C. contorta is environmentally bound to ostracod assemblage 4a and
4b: quiet water, moderate depths (?100—200 m, inner-mid shelf).

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 19

Fig. 7. A. Cytherelloidea umzambaensis Dingle, 1969, SAM-—K5573, BH9 106,0 m, LV,
Campanian I. B. Cytherelloidea contorta Dingle, 1980, SAM-K5576, BH9 110,0 m, RV,
Campanian I. C. Cytherelloidea griesbachi Dingle, 1980, SAM-K5575, BH9 88,39 m, RV,
Campanian II. D. Platella africana sp. nov., holotype, SAM—K5669, locality 21-1, Monzi, LV,
Campanian V. E. Platella africana sp. nov., SAM-—K5670, locality 113-0, Nibela Peninsula,
RV, Campanian IV.
Scale bars all 100u.

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

Cytherelloidea griesbachi Dingle, 1980
Fig. 7C
Cytherelloidea griesbachi, Dingle, 1980: 10-11, figs 3B, 4D.

Remarks

This rare species has not been encountered outside the Richards Bay BH-9
borehole.

Age, distribution, palaeoecology

Upper Santonian III to Campanian I (Richards Bay BH-9 borehole). C.
griesbachi occurs in ostracod assemblages 3, 4a and 5a and 5b, but the small
numbers available do not allow a precise assessment of its environmental
preference. However, because it is restricted to the deeper water populations in
assemblage 3, C. griesbachi was probably confined to quiet, moderate to
deep-water environments (~100—500 m, inner-outer shelf).

Cytherelloidea mfoloziensis sp. nov.
Figs 8A-E, 9A-B

Derivation of name

Locality of type.

Holotype
SAM-K5664, LV, locality 20—7/1, Mfolozi River, Maastrichtian II

Paratypes

SAM-K5665, RV, locality 20-7/1, Mfolozi River, Maastrichtian II
SAM-K5666, LV, locality 113-3, Nibela, Campanian IV
SAM-K5667, RV, locality 113-3, Nibela, Campanian IV
SAM-K5668, LV, locality 20-7/1, Mfolozi River, Maastrichtian II

Diagnosis
Species with three longitudinal ribs and a prominent AM rim. The most
conspicuous feature is a concave-upward ventrolateral ridge.

Description

External features. In lateral view rectangular. AM symmetrically rounded,
PM truncated in females, rounded but posterodorsally truncated in males. DM
generally straight but with slight concavity in front of mid point. VM straight to
slightly concave. Surface ornamented with three longitudinal ribs and a wide
AM rim. The latter merges into the VM and DM, but is not connected to the
other longitudinal elevations. Dorsal rib is curved and ventrally deflected at
about quarter length, in some specimens its anterior end merges with the median
rib; it rises posteriorly and joins the valve margin posterodorsally. The median

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 21

Fig. 8. Cytherelloidea mfoloziensis sp. nov. A. Holotype, SAM-K5664, locality 20-7/1,
Mfolozi River, LV, Maastrichtian Il. B. SAM-—K5665, locality 20-7/1, Mfolozi River, RV,
Maastrichtian II. C. SAM-K5666, locality 113-3, Nibela Peninsula, LV, Campanian IV.
D. SAM-K5667, locality 113-3, Nibela Peninsula, LV, Campanian IV. E. SAM-K5668,
locality 20-7/1, Mfolozi River, LV, Maastrichtian II.
Scale bars all 100y.

22 ANNALS OF THE SOUTH AFRICAN MUSEUM

rib runs from the posterodorsal corner to the mid point where it is deflected
horizontally; it frequently swells anteriorly. The ventral rib is short and concave
dorsally; in males it is not attached at its extremities, but in females merges with
a posteroventral swelling.

Internal features. No internal views seen.

Remarks

C. mfoloziensis has a similar rib pattern to C. umzambaensis (Fig. 9) and it
is suggested that the former evolved from the latter in lowermost Campanian
IV. The two can be distinguished by the prominence of the curved ventrolateral
rib and distinct separation of the AM and dorsal ribs in C. mfoloziensis, and the
more bulbous nature of the posterior ends of the dorsal and ventral ribs of C.
umzambaensis. C. mfoloziensis is an important subsidiary species in the Maas-
trichtian faunas where it constitutes up to 17 per cent of the cytherellid element
(5% total ostracod fauna).

Dimensions (mm)

length height
K5664 0,45 @:25
K5665 0,47 0,28
K5666 0,50 0,28
K5667 0,50 0,28
K5668 0,45 0:25

Age, distribution, palaeoecology

Campanian IV to Maastrichtian II (Nibela, Monzi and Mfolozi areas,
Zululand). C. mfoloziensis occurs in ostracod assemblages 4b, 5a, 5b, 6-7,
indicating that it tolerated a variety of medium to deep water environments (Fig.
6, Table 4). However, it preferred the deep (>500 m outer shelf/upper continen-
tal slope) stable conditions represented by ostracod assemblage 7.

Genus Platella Coryell & Fields, 1937
Platella africana sp. nov.
Fig. 7D-E

Derivation of name

Locality of type.

Holotype
SAM-K5669, LV, locality 21-1, Monzi, Campanian V

Paratypes

SAM-K5670, RV, locality 113-0, Nibela, Campanian IV
SAM-K5671, LV, locality 113-0, Nibela, Campanian IV

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 23

umzambaensis

Fig. 9. Sketches of Cytherelloidea species, right valves, with positive features shaded.
A. C. mfoloziensis sp. nov., SAM-—K5665, locality 20-7/1, Mfolozi River, Maastrichtian II.
B. C. mfoloziensis sp. nov., SAM-K5667, locality 113-3, Nibela Peninsula, Campanian IV.
C. C. umzambaensis Dingle, 1969, BH9 120,22 m, Santonian III.
Scale bars all 300y.

Diagnosis
Species with curved ventrolateral and rounded posterodorsal swellings.

Description

External features. In lateral view, asymmetrically ovate. AM broadly and
symmetrically rounded, PM asymmetrically rounded, truncated ventrally. DM
weakly convex, VM straight. There is a narrow AM rim and a broad pos-
teroventral marginal elevation. Central area has short, curved ventrolateral and
rounded, posterodorsal swellings. There is a median sulcus at about mid length
and within this the impressions of the adductor MS are clearly visible: they
consist of a curved double row of six elliptical scars—the typical ‘feather’ shape
associated with the genus Cytherella. Surface covered overall with coarse, widely
spaced ovate fossae, in some areas (e.g. adjacent to the AM rim) forming a
reticulate pattern.

Internal features. No internal views available.

Remarks

This genus has been recorded from the Campanian of Western Australia by
Bate (1972), who illustrated two juveniles in open nomenclature. Our species
differs from these in details of surface relief.

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

Dimensions (mm)

length height
K5669 0,50 0,29
K5670 0,49 0,29
K5671 0,47 0,29

Age, distribution, palaeoecology

Campanian IV to Maastrichtian III (Monzi, Mfolozi, and Nibela areas
Zululand, and Agulhas Bank). P. africana occurs in ostracod assemblages 4b,
6-7, and sample 818 (Fig. 6G). This shows that whilst it always occurs in minor
amounts it was moderately environmentally tolerant. It seems to have preferred
the deep-water (>500 m outer shelf/upper continental slope) unstable environ-
ment represented by ostracod assemblage 6.

Suborder PopocoPIna Sars, 1866
Superfamily BAIRDIACEA Sars, 1888

The Bairdiacea are one of the most important, and locally dominant,
ostracod groups in the Upper Cretaceous of south-east Africa. Present in the
Santonian sequences of Richards Bay BH—9 borehole and Umzamba in subsidi-
ary numbers (up to 10% total ostracod population), they suddenly appear in
large numbers in the lower Campanian I with the onset of deeper-water
sedimentary environments, and rapidly increase through the lower Campanian,
replacing the Cytheracea to become the dominant group (40-45% total popula-
tion) in most of the deeper water habitats. The superfamily is represented by
two genera, Bairdoppilata and Bythocypris, and although both reach their
maximum numerical development in the deeper water environments, Bythocy-
pris demonstrates the greater environmental tolerance by maintaining a more
consistent presence in the harsher (less stable) shallow-water facies represented
by assemblages 1 and 2 and the deep-water facies of assemblage 6 (Fig. 10C).

Family Bairdiidae Sars, 1888
Genus Bairdoppilata Coryell, Sample & Jennings, 1935

One of the most important genera in the Campanian—Maastrichtian rocks of
south-east Africa, it forms 29 per cent of total fauna (average of six assemblage
means) in the Zululand outcrops and BH-9 borehole (>30% in four of the six
assemblages), 25 per cent at Igoda, 90 per cent at Needs Camp, 28 per cent in
borehole JC-1, and 1 per cent in Agulhas Bank sample 818.

Three species (B. andersoni, B. africana, B. sp. A) and one subspecies (B.
andersoni aequalis) have been recognized, but only in the shallow water environ-
ments of the Transkei Swell does notable mixing appear to have taken place
(Table 1). Here, B. andersoni, B. africana, and B. sp. A all occur, whereas in
Zululand, BH-9, and JC-1, the faunas are probably monospecific (B. andersoni
and B. cf. africana, respectively).

N
WN

CAMPANIAN AND MAASTRICHTIAN OSTRACODA

BAIRDIACEA

stage
locality
ostr.
assembl.

ao)
2 BAIRDOPPILATA BYTHOCYPRIS

Bythocypris

>80 %/o
Bairdoppilata

CAMPANIAN MAASTRICHTIAN
andersoni
richardsbayensis

IWISANTONIAN

50 150 250

Fig. 10. Distribution of Bairdiacea in Campanian—Maastrichtian strata of Zululand and

Agulhas Bank. A. Total number of valves of Bairdoppilata andersoni. B. Total number of

valves of Bythocypris richardsbayensis. C. Bairdoppilata andersoni and Bythocypris richards-
bayensis as percentage of Bairdiacea.

Bairdoppilata andersoni Dingle, 1980
Figs 11A—D, 13A-B
Bairdoppilata andersoni Dingle, 1980: 12-14, fig. SA-F.

Remarks

Large numbers of well-preserved valves and carapaces of this robust species
have been recovered from Zululand and BH-9 (Fig. 10A), whilst mainly
carapaces occur at Needs Camp. B. andersoni is numerically the single most
important ostracod species in the Campanian—Maastrichtian of south-east
Africa, where it dominates (>20%) five of the six ostracod assemblages recog-
nized in the Zululand and BH-9 populations.

Age, distribution, palaeoecology

Santonian II to Maastrichtian HII (Richards Bay BH-9 borehole, Zululand
outcrops, Igoda, Needs Camp, and Agulhas Bank). B. andersoni occurs in all

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 11. Bairdoppilata. A-—D. B. andersoni Dingle, 1980. A. SAM-K5577, BH9 88,39 m, RV,
Campanian II. B. SAM—K5578, BH9 88,39 m, LV, Campanian II. C. SAM—-K5672, Lower
Quarry Needs Camp, RV, late Campanian/early Maastrichtian. D. SAM-—K5673, Lower
Quarry Needs Camp, RV, late Campanian/early Maastrichtian. E. B. andersoni aequalis
(Chapman, 1916), holotype, SAM-—2736/20, Lower Quarry Needs Camp, LV, late Cam-
panian/early Maastrichtian. F. B. andersoni aequalis (Chapman, 1916), holotype, SAM-2736/
20, Lower Quarry Needs Camp, internal LV, late Campanian/early Maastrichtian.
Scale bars: C-D, F = 100u, A-B, E = 300p. ;

CAMPANIAN AND MAASTRICHTIAN OSTRACODA ai

Fig. 12. Bairdoppilata. A. B. andersoni aequalis (Chapman, 1916), SAM-2736/17, Lower
Quarry Needs Camp, RV, late Campanian/early Maastrichtian. B. B. andersoni aequalis
(Chapman, 1916), SAM-K5674, Lower Quarry Needs Camp, LV, late Campanian/early
Maastrichtian. C. B. africana (Chapman, 1916), holotype, SAM-—2736/19, Lower Quarry
Needs Camp, RV, late Campanian/early Maastrichtian. D. B. africana (Chapman, 1916),
holotype, SAM-2736/19, Lower Quarry Needs Camp, dorsal view, carapace, late Campanian/
early Maastrichtian. E. B. cf. B. africana (Chapman, 1916), SAM—K5675, JC-1 1625 m, RV,
Maastrichtian. F. B. cf. B. africana (Chapman, 1916), SAM-K5676, JC-1 1884 m, RV,
Campanian.
Scale bars: A, F = 300u, others = 100y.

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

Sa

andersoni

a. 2 Moe

andersoni aequalis

F

SS africana

cf. africana

Fig. 13. Sketches of Bairdoppilata. A. B. andersoni Dingle, 1980, SAM—K5578, BH9 88,39 m,
LV, Campanian II. B. B. andersoni Dingle, 1980, SAM-K5577, BH9 88,39 m, RV, Campa-
nian II. C. B. andersoni aequalis (Chapman, 1916), SAM-2736/20, Lower Quarry Needs
Camp, LV, late Campanian/early Maastrichtian. D. B. andersoni aequalis (Chapman, 1916),
SAM-2736/17, Lower Quarry Needs Camp, RV, late Campanian/early Maastrichtian. E-F. B.
africana (Chapman, 1916), SAM-2736/19, Lower Quarry Needs Camp, LV(E) and RV(F), late
Campanian/early Maastrichtian. G-—H. B. cf. B. africana (Chapman, 1916), SAM-—KS5675, JC-1
1625 m, LV(G) and RV(H), Maastrichtian.
Scale bars all 300y.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 29

the ostracod assemblages recognized in the Richards Bay—Zululand area, but its
environmental preference was for the deeper-water facies, especially 5a and 7
(i.e. water depths >300 m in stable environments) (Table 4).

Bairdoppilata andersoni aequalis (Chapman, 1916)
Figs 11E-F, 12A-B, 13C-D
Bairdia subdeltoidea Minster sp. var. aequalis var. nov. Chapman, 1916: 115, pl. XV, fig.

17a—b. (SAM-2736/20.)
Bairdia subdeltoidea Minster, Chapman, 1916: 114-115 (no ijlustration). (SAM-2736/17.)

Remarks

Two specimens of this subspecies occur in slide SAM-2736. Square 18 is
obviously the holotype illustrated by Chapman (1916, fig. 17a—b) and square 17
is probably the carapace referred to Bairdia subdeltoidea by Chapman, but not
illustrated.

These specimens have been retained here within a subspecific allocation
because they differ subtly but significantly in outline from typical examples of
Bairdoppilata andersoni s.s. These differences are shown in Figure 13 and can be
summarized as: a straighter VM (particularly in LV), and a more rounded DM
(RV) in subspecies aequalis. Consequently, andersoni s.s. has a more almond-
like outline in lateral view. MS in the two varieties are, however, almost
identical.

The ‘deep sinus’ on the VM of specimen SAM-2736/17 recorded by
Chapman (1916) is merely a result of damage to the carapace. Small teeth are
clearly visible on the hinge of the holotype, which is remarkably well preserved
in comparison with other ostracod specimens recovered from the Needs Camp
outcrop.

Age, distribution, palaeoecology

Late Campanian/early Maastrichtian (Needs Camp, Lower Quarry).
Restriction of this subspecies to this locality suggests that it may have been
confined to very shallow (<20 m) normal marine, moderate to high-energy
environments with coarse carbonate sand substrates.

Bairdoppilata africana (Chapman, 1916)
Figs 12C-D, 13E-F
Bairdia africana Chapman, 1916: 115, pl. 15, fig. 19a—c. (SAM-2736/19.)

Remarks

The only specimen of this species available is the carapace of the holotype
described by Chapman (1916). It has distinctive RV and LV lateral outlines with
a strong upward tilt of the RV AM and a tapering outline to the posterior part of
the valve. Greatest height occurs at 43 per cent length (cf. 49% in B. andersoni)
and greatest width (dorsal view) occurs at 50 per cent length (cf. 47% in B.
andersoni).

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

Age, distribution, palaeoecology

Late Campanian/early Maastrichtian (Needs Camp, Lower Quarry).
Restriction of this species to this locality suggests that it may have been confined
to very shallow (<20 m) normal marine, moderate to high energy environments
with coarse carbonate sand substrate. Comparable specimens have been re-
corded from borehole JC-—1 (see below).

Bairdoppilata cf. B. africana (Chapman, 1916)
Figs 12E-F, 13G—-H
Remarks

Specimens of Bairdoppilata from the JC—1 borehole are very close to the
holotype of B. africana from Needs Camp. In particular, they possess the
upturned AM outline, and asymmetric DM (maximum height at 40% length)
that typifies Chapman’s species. The material from borehole JC—1 cannot,
however, be unequivocally assigned to B. africana because it has a slightly more
convex VM outline. Without more topotypic material it is not clear whether this
variation falls within the intraspecific morphological range of B. africana.

Age, distribution, palaeoecology

Campanian—Maastrichtian in borehole JC—1 (upper and lower limits not
known). B. cf. africana is the most abundant and widespread ostracod species in
this part of the borehole, so presumably it was relatively tolerant of the
restricted circulation, high sedimentation rate of the 100-200 m deep delta top
basin that the JC-1 Campanian—Maastrichtian sequence is thought to represent.

Bairdoppilata sp. A.
Fig. 14E

Remarks

A distinctive, elongate species with a strongly arched DM. It possesses only
a weakly developed typical ‘bairdia’ boat-shape and has a finely punctate surface
ornamentation. No internal views were available and, although this is probably a
new species, better preserved material is required to define it.

Age, distribution, palaeoecology

Late Campanian/early Maastrichtian Igoda Formation. B. sp. A constitutes
6 per cent of the Igoda ostracod population and, as such, is subordinate to B.
andersoni which constitutes 19 per cent (Table 11). Its restriction to Igoda
suggests that it might have been confined to low-energy, normal-marine, moder-
ate-water depths (~100 m, inner shelf).

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 31

Genus Bythocypris Brady, 1880
Bythocypris richardsbayensis Dingle, 1980
Fig. 14A—C
Bythocypris richardsbayensis Dingle, 1980: 14-16, fig. 6(A-E).

Remarks |

Large numbers of well-preserved specimens of this species have been
recovered from the outcrops in Zululand north of the Richards Bay borehole,
where it consistently constitutes >10 per cent of the total ostracod fauna. In the
palaeoecological analyses attempted in the following section, variations in the
numbers present of this species are found to be useful indicators in palaeo-
sedimentary environmental discrimination and have led to the concept of the
‘Bythocypris line’ to divide fields on the Cytheracea/Cytherellidae/Bairdiacea +
Cypridaea (CCBC) triangular diagram (Fig. 68).

Age, distribution, palaeoecology

Santonian II to Maastrichtian III (Richards Bay BH-9 borehole, Monzi,
Mfolozi, and Nibela areas in Zululand, Agulhas Bank sample 818, Igoda). B.
richardsbayensis occurs in ostracod assemblages 1-2, 4a, 4b, Sa, 5b, 6-7,
indicating that it had a tolerance of environmental conditions that ranged from
high-energy, restricted, shallow marine to quiet, deep, open ocean upper slope.
However, Table 4 shows that B. richardsbayensis had a definite preference for
the deep-water (>500 m, outer shelf/upper slope) environments represented by
ostracod assemblages 6—7. Its relationship with Bairdoppilata shows some in-
teresting variations (Fig. 10C). In the small number of samples from the
shallow-water environments of the Richards Bay BH-9 borehole (Santonian,
assemblages 1-2), Bythocypris richardsbayensis never constitutes less than 50 per
cent of the bairdiacean component, and is frequently the only representative
present. This indicates that, although it was at its tolerance limits in these
environments, it was far better able to cope with them than Bairdoppilata. As
soon as deeper-water conditions were established in the Zululand—Richards Bay
area in Campanian times, however, the position was immediately reversed, with
Bairdoppilata consituting less than 40 per cent of the bairdiacean element at only
one horizon. In fact, Bythocypris richardsbayensis forms >30 per cent of the
bairdiacean element only in ostracod assemblage 6. Further examination of
Figure 10 indicates that, given the fact that Bairdoppilata is always dominant in
the deep-water habitats, Bythocypris is relatively more tolerant of extremes:
shallow, restricted circulation, high energy; and deep, unstable, outer shelf/
upper slope, environments.

Bythocypris? sp.
Fig. 14D

Remarks

Six poorly preserved carapaces from borehole JC-1 have tentatively been
assigned to Bythocypris. Because no internal views are available allocation is

32 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 14. A-C. Bythocypris richardsbayensis Dingle, 1980. A. SAM-K5678, locality 20-1/2,
Mfolozi River, LV, Maastrichtian I. B. SAM-—K5679, locality 20-1/2, Mfolozi River, RV,
Maastrichtian I. C. SAM-—K5680, Igoda Formation, Igoda estuary, RV, late Campanian/early
Maastrichtian. D. Bythocypris sp., SAM-K5681, JC-1 1756 m, RV, Maastrichtian. E. Bair-
doppilata sp. A, SAM-K5677, Igoda Formation, Igoda estuary, RV, late Campanian/early
Maastrichtian.
Scale bars all 100y.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 33

purely on external outline, and it is possible that the specimens could belong to
Pontocyprella or even Paracypris. The species differs from B. richardsbayensis in
having its highest point of outline in the anterior half and in possessing a more
asymmetric posterior outline with a more strongly sloping posterodorsal region.

Age, distribution, palaeoecology

Maastrichtian (levels 1 780 m to 1 570 m) in JC-1 borehole. No specimens
of this species occur in association with Jnoceramus prisms in the Campanian—
lower Maastrichtian strata, nor at horizons in which charophytes occur. The
latter phenomenon suggests that the species was not tolerant of the environ-
ments that prevailed during the large influxes of fresh-water debris in the
mid-Maastrichtian times.

Superfamily CYPRIDACEA Baird, 1845

The Cypridacea forms a consistent, subsidiary element of the ostracod
assemblages in the Upper Cretaceous of south-east Africa, rarely constituting
more than 10 per cent. It is represented by two genera (Paracypris and
Pontocyprella) and five species, whose distribution shows strong environmental
control (Fig. 1SA—C, Table 4). Paracypris occurs throughout the Santonian II to
Maastrichtian III but is rare above the shallow-water facies of the Santonian,
whilst Pontocyprella occurs solely in the moderate to deep-water facies of the
Upper Campanian and Maastrichtian. The superfamily (Paracypris only) has
been encountered only occasionally and in trace amounts away from the
Zululand—Richards Bay area.

Family Paracyprididae Sars, 1923
Genus Paracypris Sars, 1866

The genus is present throughout the Campanian—Maastrichtian succession
of Zululand, but away from the BH-9 borehole it occurs only as scattered
individual valves and carapaces (Fig. 15A) which are invariably poorly pre-
served. In strata younger than Campanian II it never constitutes more than 2 per
cent of the total fauna (Fig. 15C). The two species which were recognized by
Dingle (1980) in the well-preserved material of BH—-9 borehole (P. umzambaen-
sis and P. zululandensis) can be recognized in outcrops, but because confident
allocation is based on MS pattern, it can rarely be made in these younger strata
where good internal views are not available. Consequently, it has not been
possible meaningfully to plot the distributions of these two species through the
Campanian—Maastrichtian succession in Zululand (for instance in Figs 15,
77-79), and they have been combined under the grouping Paracypris spp.

In the other areas, the genus is very rare and poorly preserved: Igoda (11
specimens, P. umzambaensis, P? sp. A); JC-1 borehole (1 specimen, P? sp.);
and Agulhas Bank (2 specimens P. zululandensis).

34 ANNALS OF THE SOUTH AFRICAN MUSEUM

> r=]
® = E
o F 3 PARACYPRIDIDAE =3
a & PARACYPRISJ] PONTOCYPRELLA PARICERATINA o9

MAASTRICHTIAN
TOOT

Ls}
~
s
”
-
<=

nibelaensis

CAMPANIAN

SANTONIAN 5 24

Fig. 15. Distribution of Paracyprididae and Pariceratina in Campanian—Maastrichtian strata of
Zululand and Agulhas Bank. A. Total number of valves of Paracypris spp. B. Total number
of valves of Pontocyprella nibelaensis. C. Percentage of total ostracods: Paracypris spp
(shaded), Pontocyprella nibelaensis (blank). D. Total number of valves of Pariceratina hirsuta.

Paracypris umzambaensis Dingle, 1969
Fig. 16A—C
Macrocypris simplex Chapman, 1898, Chapman, 1904: 233, pl. 29 (fig. 22).

Paracypris? umzambaensis Dingle, 1969: 354-356, fig. 5.
Paracypris umzambaensis Dingle, 1980: 17, figs 7A-C, 9A.

Remarks

In the absence of good internal views, differentiation between P. umzam-
baensis and P. zululandensis is based on the more acuminate posterior outline of
the former. In the case of the plumper varieties of both species, however,
differentiation is often subjective, and on the range charts it has not been
attempted. Specimens of P. umzambaensis from Igoda compare favourably with
the plumper varieties from BH-9 borehole (compare Fig. 16A-B), but no
internal views were available.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 35

Age, distribution, palaeoecology

Santonian III (Umzamba), Santonian II to Maastrichtian II (BH-9, and
Mfolozi River, Zululand), late Campanian/early Maastrichtian (Igoda). The
sparse distribution of the genus Paracypris in assemblages 5b, 6-7 (Fig. 15C)
indicates that it preferred the shallow-water environments of 4a (100-200 m,
inner-mid shelf). The meagre evidence available indicates, however, that P.
umzambaensis was probably more tolerant of the deeper water conditions than
was its close relative P. zululandensis.

Paracypris zululandensis Dingle, 1980
Fig. 16D
Paracypris zululandensis Dingle, 1980: 17-19, figs 7D-G, 9B.

Remarks

Identifications based solely on valve shape suggest that P. zululandensis
occurs at outcrops on the Nibela Peninsula, as well as on the Agulhas Bank. As
indicated above, however, several identifications in the genus are equivocal, and
on the range charts produced herein, no formal attempt has been made to
differentiate from P. umzambaensis.

Age, distribution, palaeoecology

Santonian II to ?Maastrichtian I (BH-9 borehole and Nibela Peninsula,
Zululand), Maastrichtian III (Agulhas Bank). What little data are available
suggest that P. zululandensis was less able to tolerate the deeper water environ-
ments of the Maastrichtian than was P. umzambaensis.

Paracypris? sp. A
Fig. 16E

Remarks

An acuminate species, tentatively referred to Paracypris, occurs in the
Igoda Formation. Preservation is poor and all the specimens are carapaces. The
elongate outline is very close to P. umzambaensis, but the DM is less strongly
arched.

Age, distribution, palaeoecology

Late Campanian/early Maastrichtian Igoda Formation at Igoda. This en-
vironment is considered to have been low-energy, normal-marine, moderate-
water depth (~100 m, inner shelf).

Paracypris? sp.
Remarks

One poorly preserved, fragmentary valve is tentatively referred to Paracyp-
ris from level 1 835 m (lowermost bed in the Maastrichtian) in the JC-1

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 16. A-C. P. umzambaensis Dingle, 1969. A. SAM-K5586, BH9 88,39 m,
LV, Campanian II. B. SAM-K5682, Igoda Formation, Igoda estuary, LV, late Campanian/
early Maastrichtian. C. SAM-—K5683, locality. 20-7/1, Mfolozi River, RV, Maastrichtian II. D.
P. zululandensis Dingle, 1980, SAM-K5590, BH9 110,0 m, LV, Campanian I. E. P. sp. A,
SAM-K5684, Igoda Formation, Igoda estuary, LV, late Campanian/early Maastrichtian.
Scale bars all 100.

- A eG a a a
Sp SS sss

CAMPANIAN AND MAASTRICHTIAN OSTRACODA es

borehole. Its significance is that it is the only representative of the genus
recovered from the Campanian—Maastrichtian section of the JC—1 borehole.
Dingle (1976) found small numbers of Paracypris (probably a different species)
in the Eocene—Oligocene part of the borehole.
Pontocyprella Lubimova, 1955
Pontocyprella nibelaensis sp. nov.
Figs 17A-C, 18

Derivation of name

Locality of type.

Holotype
SAM-K5685, RV, locality 113-3, Nibela, Campanian IV

Paratypes ere
SAM-K5686, LV, locality 113°3. Nibela, Campanian IV
SAM-K5687, RV, locality 113-3, Nibela, Campanian IV

Diagnosis
Large’ species with prominent anterodorsal margin concavity in RV lateral
view. :

Description + «

External features. In lateral view reniform. Asymmetrically rounded AM,
broadly pointed PM with apex in ventral region. DM strongly arched with a
wide anterodorsal concavity, particularly in RV. VM concave. Greatest height
about mid length although the greatest arching of the DM is posterior of this.
Valves are plump and large with smooth surfaces.

Internal features. Hinge simple, adont with gently arched groove in RV.
MA moderately wide anteriorly, narrow posteriorly and ventrally. Vestibules
wide, MPC very short and narrow, at least 20 anteriorly. MS consist of four
large oblong scars, two of which are crossed by a small sinus.

Remarks

This large species does not fit comfortably into the genus as erected by
~ Lubimova on P. harrisiana (Jones) and has certain characters in common with
genera such as Macrocypris and Argilloecia. However, it lacks the toothed hinge
and MS rosette of the former, and the small size and very wide MA of the latter.
Our species is very close to ?Pontocyprella sp. described by Neale (1974 pl. 5
(fig. 1)) from the Santonian of western Australia, and has an almost identical
MS pattern with P. dorsoconvexa Bate from the Campanian of western Austra-
lia. The latter species and P. nibelaensis differ significantly in lateral outline,
however. Bythocypris simulata Jones was recorded from the Santonian of

38 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 17. A-C. Pontocyprella nibelaensis sp. nov. locality 113-3, Nibela Peninsula, Campanian

IV. A. SAM-K5686, LV. B. Holotype, SAM-K5685, RV. C. SAM-K5687, internal RV.

D-E. Cythere? postcultrata Chapman, 1916, SAM-2736/18, Lower Quarry Needs Camp,
external view (D) and internal view (E).

Scale bars all 100u.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 39

Umzamba by Chapman (1904, 1923) and is similar in lateral outline to P.
nibelaensis, but lacks the anterodorsal concavity.

P. nibelaensis is a characteristic species of the Upper Campanian—Maas-
trichtian faunas of south-east Africa, locally comprising 9 per cent of the
Maastrichtian I ostracod population.

Fig. 18. Muscle scars, Pontocyprella nibelaensis sp. nov., SAM—K5687, locality 113-3, Nibela
Peninsula, RV, Campanian IV.
Scale bar 90.

Dimensions (mm)

length height
K5685 O91 0,40
K5686 0,94 0,44
. K5687 0,87 0,38

Age, distribution, palaeoecology

Campanian IV to Maastrichtian III (Nibela and Monzi areas of Zululand,
and Agulhas Bank). P. nibelaensis occurs in ostracod assemblages 4b, 5a, 5b,
6-7 (Fig. 15B), and seems to have preferred quiet, deep-water environments
(>500 m, outer shelf/upper continental slope).

Superfamily CYTHERACEA Baird, 1850

The numerical importance of the Cytheracea within the Campanian—Maas-
trichtian ostracod populations of south-east Africa (as monitored in Zululand) is
directly contolled by changes in palaeosedimentary environments in time and
space: in the shallower water environments the superfamily constitutes between
40 and 60 per cent, whereas in deeper-water environments this figure drops to
between 20 and 30 per cent. Thus, on Figure 67 there is a sharp decline in the
cytheracean percentage (50% to 20%) just above the Campanian I-II boundary
in the Richards Bay BH-9 borehole, and a similarly dramatic decline (40% to
20%) in the lower part of Campanian IV at outcrops to the north in Zululand.
As will be explained in a later section, these changes are mirrored in other facets

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

of the overall ostracod populations, and occur at different times in different
localities because of the strongly diachronous nature of the facies changes in
south Zululand.

At the other localities in south-east Africa from which either small numbers
of samples or only small ostracod populations are available, the importance of
the superfamily varies considerably: Igoda and Needs Camp (late Campanian/
early Maastrichtian) 57% and 6%, respectively; sample 818 Agulhas Bank
(Maastrichtian III) 83%; and in the Campanian—Maastrichtian section of the
JC-1 borehole 26% (11% Krithe and 15% ornamented species). In these areas it
has not be possible satisfactorily to monitor the temporal distribution of the
superfamily, but in the discussion section the significance of particular elements
of the cytheracean component are commented upon where a comparison with
the Zululand faunas is thought feasible.

In terms of number of cytheracean species as percentage of total number of
extant species, the superfamily is dominant, even in the sedimentary environ-
ments where it is numerically not important (see:Table 16). There is a slight
decline from 67 per cent of total species in the Campanian I-II to 64 per cent in
the Campanian IV, followed by a sharp increase to a high of 72 per cent in the
Maastrichtian I. Thereafter, this diversity declines to 60 per cent in the Maas-
trichtian II. The Igoda populations follow the regional trend at 71 per cent in the
late Campanian/early Maastrichtian.

Family Cytheridae Baird, 1850
Genus Cythere Mueller, 1785
Cythere? postcultrata Chapman, 1916
Fig. 17D-E
Cythere postcultrata Chapman, 1916: 116, pl. 15 (figs 18a—b). (SAM-—2736/18.)

Remarks

The holotype of this species (SAM-—2736/18) is a poorly preserved fragment
of indeterminate taxonomic status, which almost certainly does not belong to
genus Cythere. In his description, Chapman (1916) considered the specimen to
be a left valve with the ‘keel shaped prominence’ lying along the ventral margin,
but if the fragment is from an ostracod, then this orientation seems unlikely and
it is suggested that it may be a left valve from which the dorsal and posterior
areas have removed. Internal views show no evidence of MS, recognizable MA
features, or hingement, though if the suggestion here on orientation is correct
then the hinge has been broken off.

Chapman assigned no other specimens to this species and in our material
nothing, with the possible exception of Pondoina, looks at all like it.

It is possible that the holotype is not even a fragment of an ostracod and it is
certainly not suitable to be designated as a holotype. This presumably invali-
dates its taxonomic assignment by Chapman although we have not formally
applied to have the name suppressed.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 41

Family Bythocytheridae Sars, 1926
Genus Pariceratina Griindel & Kozur, 1971

Pariceratina hirsuta sp. nov.
Fig. 19A—D

¢

Derivation pf name

Latin hirsuta (rough, uneven): reference to rough valve surface.

7 Holotype

SAM-K5688, RV, locality 20—1/3,, Mfolozi River, Maastrichtian I

Paratypes

SAM-K5689, LV, locality 20—7/1, Mfolozi River, Maastrichtian II
SAM-K5690, LV, locality 20-1/3; Mfolozi River, Maastrichtian I

Diagnosis
Species with rough surface ornamentation of small spines, and an upturned
caudal process. '

Description

External features. In lateral view AM rounded and spinose, PM acuminate
with an upturned caudal process. VM slightly convex, DM undulating. Surface
ornamented with three large ventral spines distributed equidistantly along VM.
Surface ornamented all over with fine reticulation and small conjunctive and
disjunctive spines. The large surface spines bear small secondary spines.

Internal features. The hinge is rudimentary, consisting of a smooth LV bar
and corresponding RV groove, but also has very small smooth terminal ridges in
the RV. Inner surface of valve has large sulci leading to the three large lateral
spines. MS consist of a curved row of six oblong adductor scars and one rounded
anterior scar. No clear views of MA available.

Remarks

P. hirsuta differs from the closely related Australian species P. trispinosa
(Neale, 1975) in being less spinose marginally, having an upturned caudal
process, and in lacking ridges on its ventral surface.

Dimensions (mm)

length height
K5688 0,68 0,35
K5689 0,55 0,30
K5690 0,55 0,31

Other material 0,60 ?

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 19. A-D. Pariceratina hirsuta sp. nov. A. Holotype, SAM-—K5688, locality 20-1/3,
Mfolozi River, RV, Maastrichtian I. B. SAM-—K56839, locality 20—7/1, Mfolozi River, internal
LV, Maastrichtian II. C. SAM-K5689, locality 20—7/1, Mfolozi River, MS LV, Maastrichtian
II. D. SAM-K5690, locality 20-1/3, Mfolozi River, detail posteroventral spine LV, Maastrich-
tian I. E. Eucytherura? pyramidatus sp. nov., holotype, SAM-—K5691, locality 21-1, Monzi,
RV, Campanian V. ;
Scale bars: C = 10u, D = 30, others = 100w.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 43

Age, distribution, palaeoecology

Maastrichtian I to Maastrichtian II (Mfolozi River, Zululand). P. hirsuta is
confined to ostracod assemblage 7 of which it is a minor member. This suggests
that it was environmentally bound by deep, oceanographically stable conditions
(>500 m, outer shelf/upper continental slope) (Fig. 15D).

Family Cytheruridae Miiller, 1894

A family that is present only in the deeper water environments of the upper
Campanian and Maastrichtian in Zululand (Fig. 22). It is represented by three
genera and four species, only one of which (Pedicythere fragilis) attains any
numerical importance (up to 13% of the cytheracean population).

Genus Eucytherura Miller, 1894
Eucytherura? pyramidatus sp. nov.
Fig: 197

Derivation of name

Latin pyramidatus (pyramidal): reference to pyramid-shaped posteroventral
process.

Holotype
SAM-K5691, carapace, locality 21-1, Monzi, Campanian V.

Diagnosis
Species with pyramid-shaped posteroventral process that is joined to a
posterodorsal process by a vertical ridge.

Description

External features. In lateral view, quadrate with broadly rounded AM,
acuminate PM with caudal process. DM and VM ‘straight. Flared ventrolateral
ridge runs from anterior quarter to a pyramid-shaped posteroventral process at
three-quarters length. A similar-shaped, less conspicuous process lies in a
posterodorsal location at the end of a short and poorly defined dorsal rib. The
two processes are connected by a vertical rib. There is a prominent AM rim with
a lipped indentation at about mid height. Valve surface reticulate, fossae being
larger adjacent to the main ribs. Eye spot prominent at anterior cardinal angle.

Internal features. No internal features seen.

Remarks

Generic assignment is uncertain because no internal features were seen,
though valve outline and geometry are typical for the genus. The posteroventral
process is reminiscent of the icositetrahedron-shaped process of E. antipodum
from the Santonian of western Australia (Neale 1975), but the African species
differs by lacking an anterodorsal process and curved posterodorsal rib.

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

Age, distribution, palaeoecology

Campanian V (Monzi, Zululand). The one carapace available occurred in
ostracod assemblage 6 which represents a deep-water (>500 m, outer shelf/
upper continental slope), oceanographically unstable environment (Fig. 22A).

Dimensions (mm)
length height
K5691 0,40 0,23

Genus Cytheropteron Sars, 1866
Cytheropteron brenneri sp. nov.
Fig. 20A

Derivation of name
Named after Dr P. Brenner for his contribution to the knowledge of South
African lower Cretaceous ostracods.

Holotype
SAM-K5692, RV, locality 20-5, Mfolozi River, Maastrichtian I.

Diagnosis
Elongate species with pointed, spinose alae.

Description

External features. RV elongate with narrowly rounded AM and upturned
caudal process. DM broadly arched with a broad step at anterior cardinal angle.
VM straight. There is a large, pointed ala which bears spines along its posterior
edge and has a mucrose terminal spine. The ala protrudes almost at right angles
from the valve surface and is smooth distally and finely punctate proximally. The
rest of the lateral surface is smooth.

Internal features. Not clearly seen, except anterior MA is wide, with a small
central vestibule and possibly 10-12 short, faint MPC. Posterior MA is small,
but MPC could not be studied.

Remarks

C. brenneri is similar to C. (Aversovalva) mccomborum Neale, 1975, but
the latter has an overall pitted surfce, posteriorly deflected alae, and a more
symmetrically acuminate PM. C. (A.) mccomborum is from the Santonian of
western Australia.

C. brenneri is a minor species that never exceeds 1 per cent of the
Cytheracean population.
Dimensions (mm) nin
| length (over ala)
K5692 0,40 0,28

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 45

Fig. 20. Cytheropteron. A. C. brenneri sp. nov., holotype, SAM-K5692, locality 20-5, Mfolozi
River, RV, Maastrichtian I. B. C. cf. C. westaustraliense Neale, 1975, SAM-K5693, locality
20-5, Mfolozi River, LV, Maastrichtian I.

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

Age, distribution, palaeoecology

Late Maastrichtian I to Maastrichtian II (Mfolozi River, Zululand). C.
brenneri occurs in ostracod assemblages 5a, 5b, and 7 (Fig. 22B). Because of the
small numbers found, it is not possible confidently to predict which sedimentary
environment it preferred; it is known to have inhabited environments ranging
from quiet-water, moderate depth (?200—300 m, mid-outer shelf) to quiet, deep
water (>500 m outer shelf/upper continental slope).

Cytheropteon cf. C. westaustraliense Neale, 1975
Fig. 20B

Cytheropteron (Aversovalva) westaustraliense Neale, 1975: 27-30, pl. 13 (fig. 5-7), text fig.
4c-d, g-h.

Remarks

One valve which is very similar to Neale’s original description, notably the
thick rimmed, dorsally pitted alae. The African form differs slightly in having a
more arched DM in LV and subtly more posteriorly deflected alae, and these
differences may reflect a new species. Whatever the case, the two types are very
closely related, though they show a significant age difference: Santonian in the
Gingin chalk of Western Australia, and Maastrichtian I in the St Lucia Forma-
tion.

Dimensions (mm)
height
length (over ala)
K5693 0,30 0,20

46 ANNALS OF THE SOUTH AFRICAN MUSEUM

Age, distribution, palaeoecology

Maastrichtian II (Mfolozi River, Zululand). The one valve available was
found in ostracod assemblage 7 (Fig. 22C), which represents a deep-water
(>500 m, outer shelf/upper continental slope) oceanographically stable environ-
ment.

Genus Pedicythere Eagar, 1965
Pedicythere fragilis sp. nov.
Figs 21A-F, 23A-B

Derivation of name

Latin fragilis (fragile); reference to delicate nature of valves.

Holotype
SAM-K5694, RV, locality 20—7/3, Mfolozi River, Maastrichtian II

Paratypes

SAM-K5695, LV, locality 20-1/2, Mfolozi River, Maastrichtian I
SAM-K5696, RV, locality 20-1/2, Mfolozi River, Maastrichtian I
SAM-K5697, RV, locality 20-1/2, Mfolozi River, Maastrichtian I
SAM-K5698, LV, locality 20-1/2, Mfolozi River, Maastrichtian I

Diagnosis
Small, thin-shelled, acutely alate species with frilled AM and long, pos-
teroventral spines.

Description

External features. In lateral view, AM asymmetric, frilled, somewhat ragged
in appearance, PM with posterodorsal caudal process and three ventrally curved,
tusk-like posteroventral spines. DM straight, inflected upwards at posterior
extremity. VM convex, but hidden in lateral view. Surface features dominated
by massive, hollow, pointed alae, the leading edges of which continue as narrow
ridges to the AM. In dorsal view, the distal ends of the alae project almost as far
back as the PM, and in ventral view the alae bear three short longitudinal ribs.
Valve surface smooth, with rare puncta.

Internal features. MA and MS not seen clearly. Hinge modified amphidont:
in RV, ATE is a subdivided peg, PTE is elongate and denticulate; in LV, ME is
a dentate bar, and the ATE socket is open to the body of the valve ventrally (a
feature noted by Neale (1975) in his Australian species P. australis).

Remarks

P. fragilis resembles P. australis Neale, 1975, from the Santonian of
Western Australia, but differs in having more prominent alae and a frilled AM.
It has a prominent posterodorsal caudal process like P. sp. recorded by Bate
(1972) from the Campanian of Western Australia, but differs in ala shape.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA f 47

inl

is sek
Bane

See ane JF * :

eee |

Fig. 21. Pedicythere fragilis sp. nov. A. SAM-K5695, locality 20-1/2, Mfolozi River, LV,
Maastrichtian I. B. Holotype, SAM—K5694, locality 20—7/3, Mfolozi River, RV, Maastrichtian
II. C. SAM-K5696, locality 20-1/2, Mfolozi River, internal RV, Maastrichtian I. D. SAM-—
K5697, locality 20-1/2, Mfolozi River, dorsal view RV, Maastrichtian I. E-F. SAM-K5696,
locality 20-1/2, Mfolozi River, detail hinge ATE(E) and PTE(F), Maastrichtian I.
Scale bars: E-F = 30y, others = 100u.

48 ANNALS OF THE SOUTH AFRICAN MUSEUM

Dimensions (mm)

length height width
(excl. spines) (over ala) (over ala)
K5694 0,46 0527
K5695 0,45 0,22
K5696 0,45 wh
K5697 0,45 0,33
K5698 0,41 ?
Other material 0,40 G25
Other material 0,40 0.25

Age, distribution, palaeoecology

Maastrichtian I to Maastrichtian Il (Mfolozi River, Zululand). P. fragilis is
found in ostracod assemblages 5a, 5b, and 7 (Fig. 22D), and although it
constitutes between | per cent and 13 per cent of the Cytheracea population, its
distribution does not allow a definitive statement on its environmental prefer-
ences. It appears to have preferred 5a: quiet, open-water, moderate-depth
(2200-300 m, mid-outer shelf) conditions, but clearly could tolerate greater
depths.

Family Schizocytheridae Mandelstam, 1960

Bate (1972) grouped the following genera into the subfamily Schizocy-
therinae Mandelstam, 1960: Schizocythere, Cnestocythere, Acrocythere, Amphi-
cytherura, Apateloschizocythere, and Sondagella. Subdivision is based upon
presence and/or absence of eyes, the possession of schizodont or antimerodont
hinges, and differences in valve outline and ornamentation. Present data show
that four of these genera can be recognized in the South African Lower
(Sondagella and Acrocythere) and Upper Cretaceous (Amphicytherura and
Apateloschizocythere).

Together with the Brachycytheridae, the Schizocytheridae rank second in
importance behind the Tachyleberididae in diversity (6-18%) and consistent
distribution within the cytheracean families of the south-east African Cam-
panian—Maastrichtian strata (Table 17). It is represented by two genera and six
species which consistently constitute between 5 and 10 per cent of the
Cytheracea population, but locally makes up more than 10 per cent (Fig. 22K).

Genus Amphicytherura Butler & Jones, 1957

This genus ranges throughout the Santonian—Maastrichtian rocks of south-
east. Africa, and in the Campanian—Maastrichtian section is represented by four
species: A. tumida, A. zululandensis, A. sp. A, and A. armatus. All four species
occur in the Campanian, but only two persist into the Maastrichtian (Fig.
22E-H).

18s
\O

CAMPANIAN AND MAASTRICHTIAN OSTRACODA

CYTHERURIDAE SCHIZOCYTHERIDAE

CYTHEROPTERO APATELOSCHIZOCY THERE
5 2EUCYTHERURA PEDICYTHERE | AMPHICYTHERURA

locality
assembl.

)

a
©
~—

a

ostr.

MAASTRICHTIAN
uo

brenneri
cf westaustraliense
Ss lie

pyramidatus

n
n
<
®
mo)
i
i}
s
= |
N

armatus

CAMPANIAN

IMI|SANTONIAN

Fig. 22. Distribution of Cytheruridae and Schizocytheridae in Campanian—Maastrichtian strata
of Zululand and Agulhas Bank. A. Total number of valves of Eucytherura? pyramidatus.
B. Total number of valves of Cytheropteron brenneri. C. Total number of valves of Cytherop-
teron cf. C. westaustraliense. D. Total number of valves of Pedicythere fragilis. E. Total
number of valves of Amphicytherura sp. A. F. Total number of valves of Amphicytherura
tumida. G. Total number of valves of Amphicytherura zululandensis. H. Total number of
valves of Amphicytherura armatus. I. Total number of valves of Apateloschizocythere laminata.
J. Total number of valves of Apateloschizocythere mclachlani. K. Schizocytheridae as percen-
tage of Cytheracea.

Amphicytherura tumida Dingle, 1969
Figs 23C, 25A

Amphicytherura (A.) tumida Dingle, 1969: 368-370, fig. 13.
Amphicytherura tumida Dingle, 1980: 20-21, fig. 1OA-F.

Remarks

No further specimens of this species have been found above the top levels
from which it was recorded in the Richards Bay borehole (bed number 110.0)
(Dingle 1980) where it is evidently at the top of its range. One carapace has been
found in sample Pil3 (Campanian I) at Umzamba. Locally, therefore, the
extinction of A. tumida can be used as a rough guide to the Santonian-
Campanian boundary.

S50 ANNALS OF THE SOUTH AFRICAN MUSEUM

Age, distribution, palaeoecology

?Santonian II to Campanian I (Umzamba), early Santonian III to earliest
Campanian I (Richards Bay BH-9 borehole). A. tumida has been found in
ostracod assemblages 2, 2/3 transition, and 4a, indicating that it was moderately
environmentally tolerant, but preferred the shallow-water (<100 m, inner shelf)
low-energy, restricted circulation environments of assemblage 2 (Table 4).

Amphicytherura zululandensis Dingle, 1980
Figs 23D, 25B

Amphicytherura zululandensis Dingle, 1980: 21-24, fig. 11A—G

Remarks

This distinctive species has not been found outside the Richards Bay BH-9
borehole.

Age, distribution, palaeoecology

Campanian I (Richards Bay BH-9 borehole). A. zululandensis is confined
to assemblage 4a and was environmentally bound by quiet, moderate depths
(?100—200 m inner-mid shelf), where it seems to have replaced A. tumida and
locally constitutes between 5 per cent and 10 per cent of the cytheracean fauna
(Fig. 22G, K, Table 4).

Amphicytherura sp. A
Fig. 23E

Remarks

Seven specimens of a species that is close to A. tumida have been found at
three levels in upper Campanian—Maastrichtian outcrops in Zululand. They are
poorly preserved but probably represent a new species which differs from A.
tumida on the following points: sp. A has a more symmetric PM outline and
lacks the typical posteroventral cutaway of A. tumida; the DM and VM of sp. A
are almost parallel, giving it a more rectangular outline than A. tumida; in the
LV hinge, the PTE of A. tumida is larger and more complex and the anterior
end of the ME is oblong, the latter is shorter and peg-like in sp. A.

Age, distribution, palaeoecology

Campanian V to Maastrichtian II (Monzi and Mfolozi River, Zululand)
(Fig. 22E). A. sp. A is found in ostracod assemblages 5b, 6-7, though with the
small numbers involved it is not possible to say which it preferred. Presumably it
was restricted to water depths >300 m, i.e. it inhabited outer-shelf and upper-
slope environments (Table 4).

CAMPANIAN AND MAASTRICHTIAN OSTRACODA ed

Fig. 23. A-B. Pedicythere fragilis sp. nov., SAM-K5698, locality 20-1/2, Mfolozi River,

internal LV PTE(A) and ATE(B), Maastrichtian I. C-E. Amphicytherura. C. A. tumida
Dingle, 1969, SAM-K5699, BH9 110,0 m, LV, Campanian I. D. A. zululandensis Dingle,
1980, SAM-K5599, BH9 97,5 m, LV, Campanian I. E. A. sp. A, SAM-K5700, locality 21-1,
Monzi, LV, Campanian V.
Scale bars: A-B = 30y, others = 100w.

an
bo

ANNALS OF THE SOUTH AFRICAN MUSEUM

Amphicytherura armatus sp. nov.
Figs 24A-E, 25C

Derivation of name

Latin armatus (armoured): reference to encased or armoured appearance.

Holotype
SAM-K5701, LV, locality 20-3, Mfolozi River, Maastrichtian I

Paratypes

SAM-K5702, RV, locality 20-7/3, Mfolozi River, Maastrichtian II
SAM-K5703, LV, locality 20—7/3, Mfolozi River, Maastrichtian II
SAM-K5704, RV, locality 20—7/3, Mfolozi River, Maastrichtian II

Diagnosis

Heavily calcified species with a curved posterodorsal flange-like protrusion.

Description

External features. In lateral view, AM asymmetrically rounded with frilled
edge, PM acuminate, slightly asymmetric. DM and VM almost straight and
nearly parallel. Surface ornamented with irregular pits, indistinct nodes, plate-
shaped swellings and fine ribs, giving a fanciful appearance of a suit of armour.
A wide swelling runs to the posterodorsal corner from the indistinct SCT, and a
narrow ridge runs anteriorly to the AM. There is a prominent posterodorsal
shoulder which in well-preserved specimens is delicately ridged and perforate.
Behind this lies a curved flange which projects above the dorsal margin and
hides the posterodorsal outline. There is a keel-like ventrolateral ridge that runs
from the AM to a stubby, wing-like posteroventral corner. The proximal surface
of this keel has coarse quadrate reticulation. Eye spot is a prominent dome.

Internal features. Hinge schizodont, with anterior tooth of LV ME trifid,
ATE RV bifid, and PTE RV coarsely dentate. ME is denticulate. MA mod-
erately wide, possibly with very narrow vestibules anteriorly. Up to ten straight,
widely spaced anterior MPC; up to six straight, widely spaced posterior MPC.
MS: an oval anterior scar and curved row of four large, ovate adductors, ole
second of which is wedge-shaped.

Remarks

A. armatus bears a resemblance to A. tumida, but differs in being less
elongate, in having a subtly different lateral surface morphology, especially in
the shape of the posterodorsal and posteroventral swellings, and in possessing
the curved posterodorsal flange. The hinges of the two species also differ: the
anterior tooth of LV ME in A. tumida is bifid, and the ATE of A. tumida is
more coarsely dentate. The MS also differ slightly (Fig. 25).

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 53

Fig. 24. Amphicytherura armatus sp. nov. A. Holotype, SAM-K5701, locality 20-3, Mfolozi
River, LV, Maastrichtian I. B. SAM-—K5702, locality 20-7/3, Mfolozi River, RV, Maastrichtian
II. C. SAM-K5703, locality 20—7/3, Mfolozi River, internal LV, Maastrichtian II.

D-E. SAM-K5704, locality 20—-7/3, internal RV(D), MS(E), Maastrichtian II.

Scale bars: E = 10y, others = 100u.

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

>
tumida zululandensis armatus

Fig. 25. Muscle scars of Amphicytherura. A. A. tumida Dingle, 1969, Umzamba cliff, RV,
Santonian. B. A. zululandensis Dingle, 1980, SAM-K5601, BH9 110,0 m, LV, Campanian I.
C. A. armatus sp. nov., SAM-K5704, locality 20-7/3, Mfolozi River, RV, Maastrichtian II.
Scale bars 30u.

Dimensions (mm)

length height
KS5701 0,38 0,22
K5702 0,40 ‘0,25
K5703 0,40 0,24
K5704 0,38 0,24
Other material 0,40 0,24
Other material 0,36 0,24
Other material 0,45 0,26

Age, distribution, palaeoecology

Campanian II to Maastrichtian II (Nibela and Mfolozi River, Zululand)—
all records, except one specimen from locality 110 on the Nibela Peninsula, are
from Maastrichtian I-II of the Mfolozi River section (locality 20) (Fig. 22H). A.
armatus occurs in ostracod assemblages 4b, 5b, and 7, but is consistent, and in
relatively large numbers (for the family) only in 5b and 7. It is not possible to say
precisely where its preference lay, but apparently it was most successful in deep
water (?300->500 m outer shelf to upper slope) (Table 4).

Genus Apateloschizocythere Bate, 1972
Apateloschizocythere mclachlani sp. nov.
Fig. 26A-B
Derivation of name
Named after Dr I. McLachlan for his contribution to the knowledge of
South African Mesozoic stratigraphy.

Holotype
SAM-K5705, Carapace, locality 110-21, Nibela, Campanian III.

Diagnosis

Species with distinct caudal process and four lateral ridges emanating from
the AM.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA ao

Description

External features. In lateral view, small quadrate species with rounded AM
and symmetrical, acuminate PM with a caudal process. DM and VM sfraight.
Cardinal angles clearly defined. Surface strongly reticulate with horizontal and
vertical blade-like muri. Four longitudinal ridges emanate from the AM: a
ventral one which defines the ventrolateral outline; a lateroventral ridge which is
deflected ventrally in the posterior half and which ends in a sharp wing-like
process; a median ridge which deflects ventrally just in front of mid length; and a
short ridge above mid height which has a sharp ventral offset in a subcentral
position and extends to about three-quarters length. Dorsal part of valve
dominated by short vertical ridges and weaker longitudinal muri.

Internal features. Not seen.

Remarks

A. mclachlani differs from the genotype (A. geniculata Bate, 1972) by
possessing a caudal process and a fourth lateral ridge. In Australia, A. geniculata
ranges from Santonian to Campanian. A. /aminata (Maastrichtian, Agulhas
Bank) has a very similar rib pattern to A. mclachlani, but has a prominent LV
anterodorsal ear and delicate secondary reticulation.

Dimensions (mm)
length height
K5705 0,39 0,24

Age, distribution, palaeoecology

Campanian III (Nibela Peninsula, Zululand). A. mclachlani has been found
only in ostracod assemblage 4b, indicating that it was bound to moderate-depth
(?200 m inner-mid shelf) environments (Fig. 22J).

Apateloschizocythere laminata (Dingle, 1971)
Fig. 26C—F
Acrocythere? laminata Dingle 1971: 403—404, fig. 6.

Remarks

The uncertainty of generic assignment for this species when it was erected
was resolved by the creation of Apateloschizocythere by Bate (1972). SEM
photographs included herein supplement the original description and indicate
that the ornamentation of A. /aminata consists of a coarse first-order frame with
prominent blade-like longitudinal ridges and vertical muri, and a lace-like
secondary reticulation. Although Dingle (1971) indicated ‘indistinct eyespots’,
the species is now known to be blind.

A. laminata has a similar first-order surface ornamentation to the genotype
A. geniculata Bate (1972), and both species possess sieve plates over the normal
pore canal apertures with hair pore openings (Fig. 26E). The MS of A. laminata

ANNALS OF THE SOUTH AFRICAN MUSEUM

nN
nN

Fig. 26. Apateloschizocythere. A-B. A. mclachlani sp. nov., holotype, SAM-—KS5705, locality
110-21, Nibela Peninsula, LV(A) and RV(B), Campanian III. C-F. A. laminata (Dingle,
1971), TBD 818, Alphard Formation, Agulhas Bank, Maastrichtian II. C. SAM-—K5706, LV.
D. SAM-KS5707, internal LV, E. SAM-—K5706, sieve plates over normal pore canal apertures
LV. F. SAM-K5707, MS LV. -
Scale bars: E-F = 10p, others = 100y.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 57

are curious: there is an oval anterior scar and an irregular adductor formed by
the fusion of a line of four oval scars.

Age, distribution, palaeoecology

Maastrichtian III (Alphard Formation, Agulhas Bank sample 818) (Fig.
221). The ostracod assemblage of sample 818 is thought to represent an unusual
setting: a quiet shallow-water environment at the outer edge of the continental
shelf or on the uppermost continental slope.

Family Cytherettidae? Triebel, 1952

Genus Klingerella gen. nov.

Derivation of name

Named after Dr H. C. Klinger for his invaluable assistance during fieldwork
in Zululand in 1977.

Type species

K. aranearius sp. nov.

Diagnosis

Elongate quadrate genus, ornamented with longitudinal ribs/reticulation,
having a weakly developed amphidont hinge and very wide marginal areas.

Remarks

The style of the marginal areas of Klingerella suggests that it belongs within
the family Cytherettidae, but its hinge, outline, MS and general ornamentation
are reminiscent of many trachyleberids, and in this respect it has affinities with
Paracytheretta Triebel, 1941, and Neocytheretta van Morkoven, 1963. It differs
from these two genera in details of ornamentation and outline, particularly in
lacking hinge ears and an obtuse posteroventral outline.

In general outline and ornamentation, Klingerella is closer to Para-
cytheretta, whereas its inner lamella and hinge are more like those of
Neocytheretta. In addition, Paracytheretta appears to be blind, whereas Kling-
erella and Neocytheretta have eye spots. Paracytheretta has a range of Senonian
to Palaeocene in Europe, and Neocytheretta has a range Miocene to Recent in
the Indian Ocean and East Indies area.

Exclusively southern hemisphere genera with particularly wide marginal
areas are Paramunseyella Bate (Santonian) and Premunseyella Bate (Santonian—
Campanian) from Australia, but both differ from our new genus in possessing a
typical pectocytherid outline, and in having pectodont hinges. Klingerella is so
far known only from the Maastrichtian of south-east Africa.

ANNALS OF THE SOUTH AFRICAN MUSEUM

nN
(o 6)

ao)
®
QO KLINGERELLA KRITHE XESTOLEBERIS

assembl.

locality
ostr.

stage

MAASTRICHTIAN

nibelaensis

CAMPANIAN

- 5
Lal :
E :
Ls}
®
c
©
=
©

INISANTONIAN

Fig. 27. Distribution of Klingerella, Krithe, and Xestoleberis in Campanian—Maastrichtian
strata of Zululand and Agulhas Bank. A. Total number of valves of Klingerella aranearius.
B. Klingerella aranearius as percentage of Cytheracea. C. Total number of valves of Krithe
nibelaensis. D. Krithe nibelaensis as percentage of Cytheracea. E. Total number of valves of
Xestoleberis luciaensis. F. Xestoleberis luciaensis as percentage of Cytheracea.

Klingerella aranearius sp. nov.
Figs 28A-F, 29

Derivation of name
Latin aranea (spider’s web): reference to spider’s web-like reticulation
pattern.

Holotype
SAM-K5708, LV, locality 20-5, Mfolozi River, Maastrichtian I

Paratypes

SAM-KS5709, RV, locality 20-5, Mfolozi River, Maastrichtian I
SAM-K5710, LV, locality 20-5, Mfolozi River, Maastrichtian I
SAM-K5711, RV, locality 20-5, Mfolozi River, Maastrichtian I
SAM-KS5712, carapace, locality 20-5, Mfolozi River, Maastrichtian I

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 59

Fig. 28. Klingerella aranearius gen. et sp. nov. A. Holotype, SAM-K5708, LV. B. SAM-
K5709, RV. C. SAM-K5710, internal LV. D. SAM-K5711, internal RV. E. SAM-KS5710,
detail inner edge of MA post-adjacent to MS pit LV. F. SAM-K5712, dorsal view carapace.
All specimens from locality 20-5, Mfolozi River, Maastrichtian I.
Scale bars: E = 10y, others = 100y.

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

Diagnosis
Small species in which reticulation radiates from a subcentral site in a
web-like mesh, and which possesses dorsal, ventral and incipient median ribs.

Description

External features. In lateral view outline quadrate, broadly rounded AM,
obliquely rounded PM, and straight DM and VM. AM and PM are weakly
spinose. Anterior area wide, compressed, with a prominent anterior marginal
rim which continues dorsally over a small eye spot. Surface bears narrow but
sharp dorsal and ventral ribs which are not joined to the AM. Ventral rib lies
some way above VM, and is deflected upward posteriorly; dorsal rib commences
at one-third length, is slightly convex upwards, and proceeds along the upper
margin to join the PM just above mid height. There is an incipient median rib,
best developed posteriorly, which in its more anterior part is merely a series of
horizontal fossae muri. No SCT, but the surface reticulation forms a denser net
over a low subcentral node (resembling the centre of a spider’s web) away from
which the reticulation muri radiate. One particularly prominent muri line runs a
crooked course to the eye spot and another runs N-S, transverse to the median
rib line. Reticulation is weakest in the anterior part of the valve. In dorsal view
the carapace has a subtly waisted appearance about mid length. RV eye spot is
more prominent than LV eye spot.

Internal features. Hinge weakly holamphidont with small peg-like ATE in
RV and anterior part of LV ME. PTE more elongate. ME narrow and all
elements are smooth. MS trachyleberid.like with a hooked, almost coiled,
anterior scar and curved row of four oblong adductors, the top two of which are
almost subdivided. There may be two further weak spots in the centre of the
group. MA very wide, covering the entire inner surface except for the MS area
and a short distance along the anterior dorsal margin. No vestibules. MPC
hair-like and wavy, at least thirty in anterior area.

.
NG

Fig. 29. Muscle scars, Klingerella aranearius gen. et sp. nov., SAM-KS5710, locality 20-5,
Mfolozi River, LV, Maastrichtian I.
Scale bar 30y.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 61

Remarks

No other described species appears to be close to K. aranearius. Para-
cytheretta reticosa Triebel from the middle Palaeocene of Denmark has longitu-
dinal ribs and reticulation, but has a different LV outline, and Neocytheretta
snellii (Kingman) from the Recent of Indonesia has similar MA and hinge. Both
of these species differ in important aspects of either internal features or external
morphology.

In south-east Africa, Klingerella aranearius locally becomes an important
and typical member of the cytheracean population (>10%) and for this reason
has been selected as a subzonal marker in the upper part of the Dutoitella dutoiti
Zone.

Dimensions (mm)

length height width
K5708 0,52 0,28
K5709 0,48 0,26
K5710 0,52 0,27
K5711 0,53 0,27
7 12 0,58 0,21

Age, distribution, palaeoecology

Maastrichtian I to Maastrichtian II (Mfolozi River, Zululand). K.
aranearius occurs in moderate numbers in ostracod assemblages 5a, 5b, and 7,
indicating that it inhabited quiet environments ranging from ?200 to >500 m
(mid shelf to upper slope) (Fig. 27A-B). It seems to have preferred the
shallower water end of this range (?200—-300 m).

Family Progonocytheridae Sylvester-Bradley, 1948

The poor representation of this family in the Upper Cretaceous rocks of
south-east Africa is something of a puzzle in view of its strong presence in the
Jurassic-Lower Cretaceous of the Western Indian Ocean area (east Africa,
Madagascar, south-east Africa) with Majungaella and Progonocythere. The
Majungaella/Tickalaracythere group is found in the Santonian—Campanian of
Western Australia, but does not occur in south-east Africa (see Kr6mmelbein
1972).

Genus Hutsonia Swain, 1946
Hutsonia? sp. A
Fig. 30A

Remarks

One poorly preserved carapace of this species was found in the Igoda
section. It has the typical pyriform shape and reticulate ornamentation for the

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 30. A. Hutsonia? sp., SAM-K5713, Igoda Formation, Igoda estuary, RV, late Cam-
panian/early Maastrichtian. B-D. Pondoina igodaensis sp. nov. Igoda Formation, Igoda
estuary, late Campanian/early Maastrichtian. B. Holotype, SAM-K5714, RV. C. SAM-
K5715, internal LV. D. SAM-K5716, dorsal view carapace, anterior is to the right. E-F.
Krithe sp. E. SAM-K5717, JC-1 1625 m, dorsal view carapace, Maastrichtian. F. SAM-—
K5718, JC-1 1743 m, LV, Maastrichtian.
Scale bars all 100.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 63
genus, and possesses a median sulcus. Its hinge and MS could not be seen.
Hutsonia is typical of very shallow marine conditions and has so far been
positively recorded only from the upper Jurassic-Aptian of the east coast of the
North American province. If substantiated, its presence in south-east Africa
would greatly extend its known geographical and temporal range.

Age, distribution, palaeoecology

Late Campanian/early Maastrichtian (Igoda). The Igoda palaeosedimentary
environment is considered to have been low-energy, normal marine, moderate
water depth (~100 m inner shelf).

Family Cytherideidae Sars, 1925
Genus Krithe Brady, Crosskey & Robertson, 1874

Krithe is considered to be a marker for relatively deep-water sedimentary
environments: e.g. ‘most common in infra-neritic and bathyal environments’
(Van Morkhoven 1963: 340) and its absence from the upper, deep-water faunas
of the Richards Bay BH-9 borehole was considered by Dingle (1980) somewhat
anomalous. The present work has located the genus in younger sequences than
those available from BH-9, and extends the local range down into the upper
Campanian. It had previously been recorded in moderate numbers in the Lower
Palaeocene—Oligocene of the JC—1 borehole off-shore Natal (Dingle 1976) and
has now been recorded sporadically in the Maastrichtian. No species of Krithe
have been found at Igoda or Needs Camp. Significantly, Neale (1975) did not
record the genus in the Santonian of Western Australia which he reckoned had
been deposited in water depths of 80-100 m and Bate (1972) recorded only a few
fragments from the Campanian of the Carnarvon Basin.

There are no records of it from the Upper Cretaceous of west Africa or
Argentina, but Bate (in Bate & Bayliss 1969) notes its presence in small
numbers in the Campanian and Maastrichtian of Tanzania.

Krithe nibelaensis sp. nov.
Fig. 31A-F
Derivation of name

Locality of type.

Holotype
SAM-K5719, RV, locality 113-3, Nibela, Campanian IV

Paratypes

SAM-K5720, LV, locality 113-0, Nibela, Campanian IV
SAM-K5721, carapace, locality 113-0, Nibela, Campanian IV
SAM-K5722, LV, locality 113-3, Nibela, Campanian IV
SAM-K5723, carapace, locality 113-0, Nibela, Campanian IV

64 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 31. Krithe nibelaensis sp. nov. A. SAM-—K5720, locality 113-0, Nibela Peninsula, LV,
Campanian IV. B. SAM-K5721, locality 113-0, Nibela Peninsula, RV, Campanian IV.
C. SAM-K5722, locality 113-3, Nibela Peninsula, internal LV,-Campanian IV. D. Holotype,
SAM-KS5719, locality 113-3, Nibela Peninsula, internal RV, Campanian IV. E. SAM-K5723,
locality 113-0, Nibela Peninsula, dorsal view carapace,.Campanian IV. F. Holotype, SAM-
K5719, locality 113-3, Nibela Peninsula, MS RV, Campanian IV.
Scale bars: F = 30u, others = 100wu.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 65

Diagnosis
Slim species with star-shaped anterior MS.

Description

External features. In lateral view symmetrically rounded AM, acuminate
PM. DM slightly convex, VM straight. Greatest height behind mid length.
Outline in dorsal view slim, with posterior indentations clearly visible, and
separated by a narrow bar.

Internal features. MA typical of the genus, wide anteriorly with a central,
narrow vestibule that extends almost to AM. MPC not well developed, up to 4
anteriorly, 8 ventrally, and 3 posteriorly. MS consist of a stellate anterior scar
and four oblong adductors. The lower adductor is small and oval, the central
two oblong, and the upper more quadrate, with a dorsal incision which almost
cuts it in half.

Remarks

K. nibelaensis has all the characters typical of the genus and as such is
similar to many other species. Its distinctive features are its stellate anterior MS
and its morphology in dorsal view. The genus has not been widely recorded from
southern hemisphere Cretaceous strata: Bate (1972) mentions three specimens
only (from the Lower Campanian of Western Australia, only one of which has
the characteristic posterior indentation); and from various DSDP sites in the
western South Atlantic (e.g. Benson 1977). Unfortunately, the latter have not
yet been illustrated.

Our species is close to K. rocana Bertels (1973) from the early Danian of
Argentina, but differs in being slimmer in dorsal view (length: width ratio of 2,59
compared to 2,34) and in having more prominent and deeply incised posterior
indentations.

Although the species recorded from the Palaeocene—Oligocene of the JC-1
borehole by Dingle (1976) was not formally described, it is not conspecific with
K. nibelaensis (see Dingle 1976, pl. 12(44)).

Dimensions (mm)

length height width
K5719 0,55 0,27
K5720 0,60 0,31
KS721 0,70 0,30
K5722 0,61 0:31
K5723 0,70 0:27

Age, distribution, palaeoecology

Campanian IV to Maastrichtian III (Nibela and Mfolozi River, Zululand,
and Agulhas Bank). K. nibelaensis has been found in all the deeper water
assemblages in the Zululand outcrops, i.e. 5a, 5b, 6-7, and in one sample in

66 ANNALS OF THE SOUTH AFRICAN MUSEUM

assemblage 4b (Fig. 27C-D). This distribution suggests that it inhabited water
depths of ?200 to >500 m (mid shelf to upper slope) but with a preference for
the deep stable (>500 m), outer shelf/upper continental slope environments
where it locally reaches >20 per cent of the Cytheracea population. In the
somewhat more unstable upper slope environment represented by ostracod
assemblage 6, it is consistently present at >15 per cent. The non-appearance of
species of Krithe in the Campanian II (ostracod assemblage 5a, 5b) ) still cannot
be satisfactorily explained.

Krithe sp. A
Fig. 30E—-F

Remarks

Four specimens of Krithe have been recovered from borehole JC—1 which
are probably not conspecific with either Krithe nibelaensis (Campanian—Maas-
trichtian of Zululand and Agulhas Bank) or Krithe sp. 1 and K. sp. 2
(Palaeocene-lower Oligocene of higher sections of JC—1 borehole (Dingle,
1976) ). However, because no satisfactory internal views were available, the
absence of the stellate anterior scar in K. nibelaensis could not be confirmed. K.
sp. A and K. nibelaensis differ most notably in the shape of the posterior
indentation in dorsal view—in the former it is larger and more acute. K. sp. 1
and K. sp. 2 have different AM outlines to K. sp. A. The status of the species is
uncertain until more material becomes available.

Age, distribution, palaeoecology

Occurs in four samples in the Maastrichtian of JC—1 borehole, between
levels 1 625 m and 1 743 m. Only one of these levels coincides with horizons in
which charophytes is found and the species always occurs in association with one
or two of the following genera: Bairdoppilata, Cytherella or Bythocypris?. Such
a distribution is consistent with the known preference of Krithe for deeper water
environments and its presence may be helpful in identifying periods of deepen-
ing and/or shallowing on the Tugela Delta top (see Table 10 and discussion
section).

Genus Pondoina Dingle, 1969

This genus is represented in the Santonian strata of south-east Africa by P.
sulcata which reaches 10 per cent of the total ostracod population at Umzamba.
The same species is present in small numbers in the Santonian III of the
Richards Bay BH-9 borehole, but does not range into the Campanian. The only
record of Pondoina in the Campanian—Maastrichtian rocks of south-east Africa
is at Igoda. Kr6mmelbein (1972) records species tentatively assigned to Pon-
doina from the Turonian Sibang Formation of Gabon and the ?Coniacian Macau
Formation of north-east Brazil.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 67

Pondoina igodaensis sp. nov.
Fig. 30B—D
Derivation of name

Locality of type.

Holotype
SAM-K5714, carapace, Igoda Formation, Igoda Estuary, late Campanian/
early Maastrichtian

Paratypes
SAM-K5715, LV, Igoda Formation, Igoda Estuary, late Campanian/early

Maastrichtian
SAM-K5716, carapace, Igoda Formation, Igoda Estuary, late Campanian/

early Maastrichtian

Diagnosis
Plump species with strongly asymmetric AM outline in LV.

Description

External features. In lateral view, AM asymmetrically rounded. PM nar-
rower, symmetrically rounded. DM gently arched. VM in RV nearly straight or
slightly concave in anterior third in LV convex. Highest point of valve at about
one-third length. In dorsal view LV distinctly larger than RV, plump appearance
with maximum width at about two-thirds length. There is a weak, vertical
median sulcus which gives a subtly waisted outline in dorsal view. Surface
smooth with shallow rounded pits concentrated in the vicinity of the median
sulcus and the anterior border.

Internal features. Hinge antimerodont. MS and MA not seen.

Remarks
P. igodaensis differs from P. sulcata by its plumper shape and asymmetrical
LV AM outline.

Dimensions (mm)

length height width
K5714 0,69 0,36
K5715 0,70 0,38
K5716 0,80 0,45
Other material 0,81 0,48

Age, distribution, palaeoecology

Late Campanian/early Maastrichtian (Igoda Formation). The Igoda
palaeosedimentary environment is considered to have been low-energy, normal-
marine, moderate-water depth (~100 m, inner shelf).

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Xestoleberididae Sars, 1928
Genus Xestoleberis Sars, 1866
Xestoleberis luciaensis sp. nov.
Fig. 32A—G
Xestoleberis sp. A Dingle, 1980: 19-20, fig. 8A-B.

Derivation of name
Locality of type—Nibela Peninsula, Lake St Lucia.

Holotype
SAM-K5724, RV, locality 20-5, Mfolozi River, Maastrichtian !

Paratypes

SAM-K5725, RV, locality 20-1/3, Mfolozi River, Maastrichtian I
SAM-KS5726, LV, locality 20—-1/3, Mfolozi River, Maastrichtian I
SAM-K5727, RV, locality 20-1/3, Mfolozi River, Maastrichtian I

Diagnosis
Plump species, with all hinge elements crenulate and MS that consist of two
antennal scars and four adductors, the, top one being almost subdivided.

Description

External features. In lateral view, asymmetrically rounded AM with an
angular, ‘beaked’ overhang in the anteroventral area of the RV. VM convex,
DM strongly arched. Greatest height behind mid length. PM broadly rounded.
In some specimens there is a very weakly developed AM rim in the LV. Valve
surface smooth except for occasional NPC openings and faint ridges in the
anteroventral extremity.

Internal features. Hinge strongly antimerodont with a small DM projection
above ME in RV and ATE in LV. MS consist of four elongate adductors, the
top one being almost bisected, and two antennal scars: a small rounded scar
within a large sickle-shaped scar. The ‘xestoleberis spot’ is well developed. MA
moderately wide anteriorly, narrow posteriorly. Wide anterior vestibules with
apparently few (?up to six) anterior MPC. Posterior MPC could not be counted,
and despite the large numbers of specimens available the number quoted for
anterior MPC is not definite. There are numerous NPC on the interior surface
but few appear to open to the exterior.

Remarks

Because X. /uciaensis has all the characters of the genus, it is close to many
other species, but can be distinguished in well-preserved specimens on details of
hingement and MS. X. luciaensis appears to be the first Mesozoic species
recorded from the southern hemisphere, but Neufville (1979) has described
‘Danian’ species from the Sergipe Basin of eastern Brazil. His closest form is X.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 69

Fig. 32. Xestoleberis luciaensis sp. nov. A—-C. Holotype, SAM-K5724, locality 20-5, Mfolozi
River, internal RV(A) MS(B) hinge (C), Maastrichtian I. D. SAM-K5725, locality 20-1/3,
Mfolozi River, RV, Maastrichtian I. E. SAM-—K5726, locality 20-1/3, Mfolozi River, LV,
Maastrichtian I. F—G. SAM-KS5727, locality 20-1/3, Mfolozi River, dorsal view RV(F) detail
PTE(G), Maastrichtian I.
Scale bars: B; G = 10u, C = 30p, others = 100.

70 ANNALS OF THE SOUTH AFRICAN MUSEUM

chamela van den Bold, which is less plump in dorsal view and somewhat less
angular in its anteroventral outline than X. luciaensis.

Dimensions (mm)

length height width
K5724 0,40 0,25
K5725 0,40 27
K5726 0,42 0,30
K5727 0,40 0,13

Age, distribution, palaeoecology

Companian I to Maastrichtian II (Zululand: Richards Bay BH-9 borehole
and Monzi, Mfolozi, and Nibela outcrops). X. /uciaensis is an environmentally
tolerant species that has been found in ostracod assemblages 4a, 4b, 5a, 5b, 6-7
(Fig. 27E-F, Table 4). Its preference, in increasing compatibility within these,
would seem to be (values are mean % cytheracean population): 4a (~2%), 4b
and Sa (4-7%), 6 (~20%), 5b (~25%), and 7 (~30%). The significant values
here are: (i) a sharp increase in populations that are thought to represent deep-
water environments (ie >300 m), (ii) main preference for assemblage 7 which
represents stable environments >500 m (outer shelf/upper continental slope).

Family Buntoniidae Apostolescu, 1961
Genus Buntonia Howe, 1935 (in Howe & Chambers 1935)
Buntonia? sp. A
Fig. 34A

Remarks

A carapace referred to Buntonia on the grounds of its shape, particularly its
upturned posterior outline. Features not conforming to typical species of this
genus are: lack of eye spot, and smooth posterior area (which typically bears
small longitudinal ribs). Buntonia is common in the uppermost Cretaceous of
west Africa, and has been recorded from the upper Eocene of the JC—1 borehole
(Dingle 1976), but no positive identifications have yet been made in the
Cretaceous of south-east Africa.

Age, distribution, palaeoecology

Late Campanian/early Maastrichtian (Igoda Formation, Igoda). The Igoda
palaeosedimentary environment is considered to have been low-energy, normal-
marine, moderate-water depth (~100 m, inner shelf).

Family Brachycytheridae Puri, 1954
In diversity and consistency of distribution, the Brachycytheridae (jointly
with the Schizocytheridae) rank second in importance within the families of the
Cytheracea (Table 17b). Numerically, however, the Brachycytheridae are far
more important than the Schizocytheridae: in Campanian I the former consti-

|
pa

CAMPANIAN AND MAASTRICHTIAN OSTRACODA

BRACHYCYTHERIDAE

a PONTICULOCYTHERE
2 BRACHYCYTHERE PTERYGOCYTHER

assembl.

locality
ostr.

MAASTRICHTIAN

longicaudata
lanceolata
biremis

2
<
z
<
a
=
ot
Q

Fig. 33. Distribution of Brachycytheridae in Campanian—Maastrichtian strata of Zululand and
Agulhas Bank. A. Total number of valves of Brachycythere longicaudata. B. Brachycythere
longicaudata as percentage of Cytheracea. C. Total number of valves of Brachycythere sicarius.
D. Total number of valves of Pterygocythere lanceolata. E. Total number of valves of
Ponticulocythere biremis. F. Total Brachycytheridae as percentage of Cytheracea.

tutes ~30 per cent of the Cytheracea, and in Campanian II to Maastrichtian II
10-20 per cent (compare Figs 22K and 33F).

The family is represented by three genera and four species: Brachycythere
longicaudata, B. sicarius, Pterygocythere lanceolata, and Ponticulocythere bi-
remis. Only Brachycythere occurs in the Campanian and Maastrichtian (both
species are inherited from the Santonian), whilst Pterygocythere and Ponticu-
locythere are restricted to the Maastrichtian.

Genus Brachycythere Alexander, 1933
Brachycythere longicaudata (Chapman, 1904)
Fig. 34B-C

Cytheridea longicaudata Chapman, 1904: 234-235, pl. 29 (fig. 21). Howe & Laurencich, 1958:

279.
Cythere ?drupracea Jones, 1884, Chapman, 1904: 234.
Brachycythere longicaudata (Chapman), Dingle, 1969: 358-361, fig. 7. Dingle, 1980: 25-26, figs

12A-C, 13A-D.

s

72 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

B. longicaudata exhibits a good deal of intraspecific morphological varia-
tion, and the specimens from Igoda are closest to the more elongate forms at
Umzamba (type horizon), though they are slightly smaller. Bate (in Bate &
Bayliss 1969) illustrates a form which he assigns to B. aff. B. sapucariensis
Kroéommelbein, 1964, from the Turonian of Tanzania. Within the limits of his
sketch, this specimen could easily be accommodated within B. longicaudata
(compare Bate 1969, pl. 7 (fig. 1) with Dingle 1980, fig. 12B).

Age, distribution, palaeoecology

Santonian II to Maastrichtian II (Umzamba, and Richards Bay BH-9
borehole, and Mfolozi and Nibela outcrops in Zululand), late Campanian/early
Maastrichtian (Igoda).

Dingle (1980) found this to be the most common species of ostracod in the
Richards Bay BH-9 borehole, particularly in the lower, Santonian part, where it
locally constituted 88 per cent of the cytheracean population (~50% total
population). Although its importance declines up the borehole into the lower
Campanian, it occurs sporadically throughout the whole of the Campanian-—
Maastrichtian in Zululand. It is one of the most environmentally-tolerant
ostracod species in the south-east African upper Cretaceous and has been found
in ostracod assemblages 1-3, 4a, 4b, 5a, 5b, 6-7 (Fig. 33A-B, Table 4). As
shown in Figure 33B, its importance (expressed as a percentage of the cythera-
cean population) varies between 16 and 62 per cent in water reckoned to be
shallower than about 200 m, but drops sharply to between 3 and 10 per cent in
deeper water environments. It preferred the high-energy, restricted circulation,
shallow water of assemblage 1 (Santonian II, Richards Bay BH-9 borehole), but
curiously stages an ‘ecological recovery’ in the deep water (>500 m) of assem-
blages 6 and 7, compared with the low values and sporadic distribution in the
moderate depth (200-400 m) mid-shelf environments. A somewhat similar
phenomenon was noted in the case of Cythereis klingeri. A possible explanation
is of posthumous transportation into deep water by currents.

At Igoda, B. longicaudata constitutes 26 per cent of the cytheracean
population.

Brachycythere sicarius Dingle, 1980
Fig. 35A
Brachycythere sicarius Dingle, 1980: 27-29, figs 13E, 14A-F.

Remarks
This species has been located at only two horizons above the stratigraphic
level of the Richards Bay BH-9 borehole (Fig. 33C).

Age, distribution, palaeoecology

Santonian II to Maastrichtian I (Richards Bay BH-9 borehole and, Mfolozi
and Nibela outcrops). B. sicarius occurs only sporadically in the Upper Cre-

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 73

Fig. 34. A. Buntonia? sp., SAM-K5728, Igoda Formation, Igoda estuary, LV, late Cam-
panian/early Maastrichtian. B-—C. Brachycythere longicaudata (Chapman, 1904). B. SAM-
K5603, BH9 106,2 m, RV, Campanian I. C. SAM-K5729, Igoda Formation, Igoda estuary,
LV, late Campanian/early Maastrichtian.
Scale bars 100y.

taceous of south-east Africa, but has been found in the following assemblages:
1-3, 4a, 4b, 5a, 5b, and 7. Appearances in the deep-water assemblages 5b and 7
are confined to single samples and the species clearly preferred the moderate
depth (?100—200 m inner-mid shelf) environments represented by assemblage
4a, although even here it is generally insubordinate in numbers to its close
relative B. longicaudata.

Genus Pterygocythere Hill, 1954

Pterygocythere lanceolata sp. nov.
Fig. 35B-F

Derivation of name

Reference to lance-like spine on ventrolateral alae.

Holotype
SAM-K5730, LV, locality 20—1/3, Mfolozi River, Maastrichtian I

74 ANNALS OF THE SOUTH AFRICAN MUSEUM

Paratypes
SAM-KS5731, carapace, locality 20—-1/2, Mfolozi River, Maastrichtian I
SAM-KS5732, carapace, locality 20—-1/3, Mfolozi River, Maastrichtian I
SAM-K5733, LV, locality 20—1/2, Mfolozi River, Maastrichtian I
SAM-K5734, RV, locality 20-1/1, Mfolozi River, Maastrichtian I

Diagnosis
Smooth-surfaced species with a posteriorly directed spine on alae termina-
tions.

Description

External features. Triangular in lateral view, with marginal spines in pos-
teroventral and anteroventral positions. AM broadly rounded, PM acuminate.
DM and VM straight, converging strongly posteriorly. The surface is smooth,
except for small punctae in the posterior part. Its most prominent feature is a
large wing-like ala on each valve which has a sharp posteriorly directed terminal
spine. There is a prominent eye spot and a small post-ocular sinus.

Internal features. Hinge amphidont. PTE and ME crenulate, ATE weakly
subdivided with, in RV, a frilled spur on its anterior end. There is a deep
ventrolateral sinus leading into the ala. MS consist of a hooked anterior scar and
four adductor scars. MA moderately wide. MPC numerous, fine and straight: up
to 20 anteriorly, about 9 posteriorly.

Remarks

Placed in Pterygocythere because of its strongly alate shape. However, in all
other respects the species possesses the characters of local representatives of the
genus Brachycythere: the hinge is identical to B. longicaudata and B. sicarius (the
latter even has the frilled anterior rim in RV ATE) and the MS are identical to
B. longicaudata. It differs from these two species in shape and ornamentation,
but quite clearly is very closely related. From statigraphic and morphological
considerations it is probable that P. lanceolata developed from B. sicarius, which
itself is an alate development of B. longicaudata. The new species has been placed
in Pterygocythere rather than Alatacythere because of its prominent accommoda-
tion groove in the LV, a feature also common to the local Brachycythere species.

Dimensions (mm)

length height ~ width
K5730 0,75 0,45
K5731 0,78 0:55
K5732 0,75 0,45
K5733 0,70 0,40

K5734 0,68 0,38

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 9 is)

Fig. 35. A. Brachycythere sicarius Dingle, 1980, SAM—K5606, BH9 89,0 m, RV, Campanian
II. B-—F. Pterygocythere lanceolata sp. nov. B. SAM-K5731, locality 20—-1/2, Mfolozi River,
dorsal view carapace, Maastrichtian I. C. Holotype, SAM-—KS5730, locality 20-1/3, Mfolozi
River, LV, Maastrichtian I. D. SAM-—K5732, locality 20-1/3, Mfolozi River, RV, Maastrich-
tian I. E. SAM-—K5733, locality 20-1/2, Mfolozi River, internal LV, Maastrichtian I.
F. SAM-K5734, locality 20-1/1, Mfolozi River, MS RV, Maastrichtian I.
Scale bars: F = 30y, others = 100y.

76 ANNALS OF THE SOUTH AFRICAN MUSEUM

Age, distribution, palaeoecology

Maastrichtian I to Maastrichtian II (Mfolozi River Zululand) (Fig. 33D). P.
lanceolata occurs in ostracod assemblages 5a, 5b, and 7, but preferred the quiet,
deeper-water conditions of 5b and 7: ?300 m — >500 m (outer shelf to upper
continental slope). |

Genus Ponticulocythere gen. nov.

Derivation of name
Latin ponticulus (a small bridge) + generic name cythere: reference to
ponticulate lateral ribs.

Type species
Ponticulocythere biremis sp. nov.

Diagnosis

Subquadrate in lateral view with: ponticulate dorsal and ventral ribs; no
median rib; eye spot; smooth or postulate intercostal areas; strong AM rim.
Alate in dorsal view. Hinge entomodont or modified entomodont.

Remarks

The family placement of this genus is not certain. Externally it has features
in common with Ponticocythereis McKenzie, 1967, which has a reported range
Tertiary to Recent, with the genotype Recent from south-east Australia.
McKenzie’s species differs from Ponticulocythere in possessing a longitudinal
lateral median ridge and in having a holamphidont hinge. Because Ponticulo-
cythere has no median lateral rib, and does not have an amphidont hinge, it is
not placed in the Trachyleberididae. Its alate appearance and hinge which,
although entomodont, could be considered a modification of a weak amphidont
structure, suggest that it has affinities with genera such as Bosquetina and
Pterygocythereis. In the latter respect, Ponticulocythere is closer to Bosquetina,
with the genotype and typical species of Pterygocythereis having holamphidont
hinges. However, Ponticulocythere differs strongly from Bosquetina in their
respective lateral architecture, in which it has affinities with Pterygocythereis.
The main difference here is the spinose nature of the ribs in Pterygocythereis and
their ponticulate aspect in Ponticulocythere. In the light of these similarities, the
new genus is placed in the Brachycytheridae.

Age
It is known so far only from the Maastrichtian of south-east Africa.

Ponticulocythere biremis sp. nov.
Fig. 36A-E
Derivation of name

Latin biremis (galley with two banks of oars): fanciful reference to its
appearance in lateral view.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA ay

Holotype
SAM-K5735, RV; locality 20—1/2, Mfolozi River, Maastrichtian I

Paratypes
SAM-K5736, LV, locality 20-8, Mfolozi River, Maastrichtian II
SAM-KS5737, RV, locality 20-7/3, Mfolozi River, Maastrichtian II

Diagnosis
Species with small post-ocular sail-like projection on DM, and coarse spines
on posteroventral margin.

Description

External features. In lateral view subquadrate, broadly rounded AM,
slightly acuminate PM with coarse posteroventral spines. DM and VM straight,
converging only slightly posteriorly. DM hidden by curved ponticulate rib that
commences post-adjacent to the large rounded eye spot and ends at the
posterodorsal angle where it bears a small spine. There is a small pointed
sail-like projection on the DM immediately behind the eye spot. Ventrolateral
ponticulate rib abruptly joins the AM rim and posteriorly rises, ending at about
three-quarters length. Central area smooth with occasional small perforate
pustules and an indistinct swelling representing the SCT. Anterior area of the
valve is somewhat compressed with large, ill-formed fossae in the AM rim. In
dorsal view, arrow-shaped with ventrolateral ponticulate rib prominent along an
alate projection. Widest part of valve in posterior third.
~ Internal features. MS not seen. MA apparently narrow, but this may be due
to damage to valves. Hinge entomodont: RV has elongate dentate terminal
elements, the PTE being more prominent than ATE in dorsal view. LV has a
narrow, weakly dentate ME with a small, elongate anterior swelling, ATE
consists of a shallow, anteriorly widening depression. LV PTE not seen.

Remarks

P. biremis is not closely related to any described species. Superficially it has
features in common with several taxa with ponticulate, carinate or undercut
clavae, for example: Carinocythereis antiquata (Baird), Recent; and Ponticocy-
thereis militaris (Brady), Recent, but differs on essential points of generic
morphology.

Dimensions (mm)

length height width
K5735 0,82 0,40
K5736 0,60 0,32
K5737 0,58 0,18

Age, distribution, palaeoecology
Maastrichtian I to Maastrichtian II (Mfolozi River, Zululand) (Fig. 33E). P.
biremis occurs in very small numbers in ostracod assemblages 5a, 5b, and 7. The

78 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 36. Ponticulocythere biremis gen. et sp. nov. A. SAM-—K5736, locality 20-8, Mfolozi
River, LV, Maastrichtian II. B. Holotype, SAM—K5735, locality 20-1/2, Mfolozi River, RV,
Maastrichtian I. C-E. SAM-K5737, locality 20-7/3, Mfolozi River, ATE dorsal view (D) PTE
dorsal view (C) dorsal view RV(E), Maastrichtian II.
Scale bars: C-D = 30u, others = 100w.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 79

sparse data available suggest that the species preferred the quiet-water moderate
depths (?200-500 m mid-outer shelf) represented by assemblage 5.

Family Trachyleberididae Sylvester-Bradley, 1948

In terms of numbers of species, the Trachyleberididae is the most important
cytheracean family in the Campanian—Maastrichtian rocks of south-east Africa.
Only in the deep-water environments, represented by assemblages 5b, 6 and 7,
does it numerically fall below 40 per cent, and have a diversity (number of
species) below 50 per cent of the total cytheracean populations (see Table 17).

Subfamily Pennyellinae Neale, 1975

Neale (1975) separated a group of strongly reticulate, blind genera of
trachyleberid-type in which he placed the following taxa: Pennyella (Santonian-—
Maastrichtian), Agulhasina (Maastrichtian) and Agrenocythere (Eocene—-
Recent). In the Upper Cretaceous, these genera are confined to the South
Africa—Australia—West Pacific area, but from the Oligocene onwards, Agrenocy-
there develops into a deep-water cosmopolitan genus. Genus B. described by
Bate (in Bate & Bayliss 1969) from the Campanian of Tanzania may also belong
here.

Genus Agulhasina Dingle, 1971

Despite the availability of more material from the Upper Cretaceous of
south-east Africa, the genus remains monospecific and confined to the Maas-
trichtian III of the Agulhas Bank, suggesting that it developed late in the
Maastrichtian. However, Genus B of Bate (in Bate & Bayliss 1969) is super-
ficially similar to the genotype, raising the possibility that Agulhasina in fact
originated during the Campanian in the east Africa area. The phylogeny of the
genus is at present, therefore, unknown.

Agulhasina quadrata Dingle, 1971
Figs 37A-B, 44A
Agulhasina quadrata Dingle, 1971: 414-416, fig. 15, Pl. VII(b).

Remarks

Additional work on specimens from sample 818 on the Agulhas Bank allows
a more detailed presentation of valve morphology and MS patterns than was
possible in the original description. The MS pattern of Agulhasina quadrata (Fig.
44A) differs from that of Pennyella pennyi in having a kidney-shaped anterior
scar (the latter has a V-shaped scar), and in having two small rounded scars in
the lower positions of the posterior row, rather than elongate scars as in
Pennyella. Also, under high magnification, the indistinct SCT of Agulhasina
quadrata is seen to have two short median ribs (about 50u long) on its posterior
end.

80 ANNALS OF THE SOUTH AFRICAN MUSEUM

. 8 >

oo 4

ic)

. . * @xg
a tN
3" wo MF yas OG

. ~ .
MED RTA

s

«

wt & — se 5
=< sea .

Fig. 37. A-B. Agulhasina quadrata Dingle, 1971, TBD, 818 Alphard Formation, Agulhas
Bank, Maastrichtian HI. A. SAM-K5738, LV. B. SAM-K5739, RV. C-—E. Unicapella sacsi
Dingle, 1980. C. SAM-K5740, locality 20-7/1, Mfolozi River, LY, Maastrichtian II.
D. SAM-KS5741, locality 21-1, Monzi, RV, Campanian V. E. SAM-K5740, locality 20-7/1,
Mfolozi River, detail anterior area LV, Maastrichtian II. F. Dutoitella mimica gen. et sp. nov.,
SAM-K5748, TBD 818 Alphard Formation Agulhas Bank, detail anterior area RV, Maastrich-
tian III.
Scale bars: E = 30u, F = 10, others = 100y.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 81

Age, distribution, palaeoecology

A. quadrata is known only from the Maastrichtian HI (sample 818) on the
Agulhas Bank, and because of its distinctiveness it has been selected as the zonal
ostracod for the A. quadrata Zone. The assemblage of sample 818 is thought to
represent an unusual setting: quiet, shallow-water environment on the outer
edge of the continental shelf or on the uppermost continental slope.

Subfamily Unicapellinae subfam. nov.
Type genus
Unicapella Dingle, 1980 (Santonian—Maastrichtian).

Other genera

Dutoitella gen. nov. (Campanian—Maastrichtian), Paleoabyssocythere Ben-
son, 1977 (Campanian—Palaeocene), Atlanticythere Benson, 1977 (Campanian-
Miocene), Herrigocythere Griindel, 1973 (Campanian—? Maastrichtian).

Diagnosis

Blind, weakly reticulate to foveolate genera of general trachyleberid aspect
with prominent domed or elongate—oval SCT, strong to moderately developed
hinge ear in LV, and nodose, massively spined, or bullate surface features.

Remarks

In the same way that the Pennyellinae constitute a distinctive blind reticu-
late trachyleberid-like group, so the Unicapellinae brings together an equivalent
grouping that is characterized by its combination of finely reticulate often
delicately foveolate surface texture with coarse spines, massive nodes, and
bullae (Figs 39, 43).

Although the earliest record of the subfamily so far is of Unicapella in the
Santonian of Zululand (personal unpublished data) its main development is in
the Campanian, with the appearance of the genera Dutoitella in east and
south-east Africa, Atlanticythere and Paleoabyssocythere in the South Atlantic
basin (Benson 1977), and Herrigocythere in north Germany (Herrig 1965). In
addition, the species described by Holden (1964) as /diocythere triebeli from the
Upper Campanian—Lower Maastrichtian of California may belong to Herrigocy-
there, as suggested by Gritindel (1974). Atlanticythere and Paleoabyssocythere are
reckoned to have lived in water depths of the order of 1 000 m (Benson 1977)
and, although so far described only in detail from DSDP sites on the Rio
Grande Rise (sites 356, 21 & 22), they are said to occur ‘worldwide’ (Benson
1977). Unicapella and Dutoitella, on the other hand, are thought to occur in
moderate to deep continental shelf or upper slope environments in east Africa.

The northern hemisphere genera appear in the late Campanian—early Maas-
trichtian, whereas the southern hemisphere representatives are first found in
slightly older rocks (Santonian—early Campanian). On this evidence it seems
likely that the subfamily originated in the proto South Atlantic-South-western
Indian Ocean area and rapidly migrated northwards.

82 ANNALS OF THE SOUTH AFRICAN MUSEUM

= r

® = E
eS ae UNICAPELLINAE 3
= 3 4 an
2 & 8 UNICAPELLA DUTOITELLA og

818 Sel
z | 5 15 | 5a |
“ cs F [5b |
» ome
xr £
= E
ad £
= 7
Ke
Ww
<q
<q
=

D i E

ou

15 10

dutoiti

4a

reticulata >
Po ae

CAMPANIAN

IIISANTONIAN

Fig. 38. Distribution of Unicapellinae in Campanian—Maastrichtian strata of Zululand and
Agulhas Bank. A. Total number of valves of Unicapella reticulata. B. Total number of valves

of Unicapella sacsi.

C. Unicapella sacsi as percentage of Cytheracea. D. Total number of

valves of Dutoitella dutoiti. E. Dutoitella dutoiti as percentage of Cytheracea. F. Total number

of valves of Dutoitella mimica.

TABLE 5
Distinguishing features: Unicapella, Paleoabyssocythere, and Herrigocythere.

postero- _ventro- surface holotype
ventral lateral ornamen-_ length,
AM rim SCT bullae nodes tation LV mm
Unicapella ; ' . narrow large, prominent small, foveolate 0,55
dome- small rounded, to coarsely
shaped discon- reticulate
tinuous
Paleoabyssocythere » “Massive. » large." massive large, foveolate 0,80
wide elongate, fused to weakly
with dis- into reticulate
tinct ridge
posterior
taper
Herrigocythere ; . massive small, large, indistinct, smooth to 0,62
wide elongate, rounded fused into foveolate

posteriorly ridge

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 83

Genus Unicapella Dingle, 1980

Although the genus probably first appears in the Santonian of Zululand
(unpublished data), Unicapella is typical of the Campanian—Maastrichtian,
where two species are known, U. reticulata and U. sacsi (Fig. 38A-C). It is
closely related to Paleoabyssocythere Benson, 1977, and Herrigocythere Grin-
del, 1973. All have similar surface morphologies, shell outlines and hingement
(MS are so far known only for Herrigocythere), and generic subdivision is based
primarily upon differences in the following characters: AM rim, SCT, surface
ornamentation, posteroventral bulla, and ventrolateral ridge. These differences
are listed in Table 5 and are sketched in Figure 39.

Unicapella reticulata Dingle, 1980
Figs 39H, 40A
Unicapella reticulata Dingle 1980: 33-34, fig. 17A-B.

Remarks

Despite the large amount of new material studied, the only records of this
distinctively ornamented species remain the three valves from sample 88.0 in the
Campanian I of the Richards Bay BH-9 borehole (Dingle 1980).

Age, distribution, palaeoecology

Campanian I, Richards Bay BH-9 borehole, confined to ostracod assem-
blage 5a which represents a low-energy, open-water, moderate-depth environ-
ment (200-300 m, mid to outer shelf). U. reticulata is the only species restricted
to this ecofacies.

Unicapella sacsi Dingle, 1980
Figs 37C-E, 39A, E
Unicapella sacsi Dingle, 1980: 30-32, fig. 16A-G.

Remarks

There is some intraspecific morphological variation throughout the Zulu-
land succession, with the Campanian II forms tending to be more coarsely
reticulate and having somewhat narrower posterodorsal bullae than their
younger (e.g. Maastrichtian) counterparts. In addition, the prominent LV
anterodorsal ear is not always well developed in the Campanian II populations.
It should be noted that the small dorsal ridge in the LV of one specimen
illustrated by Dingle (1980, fig. 16B, SAM-—K5611) is now considered to be an
artefact caused by slight distortion of the dorsal valve edge during sedimentary
compaction.

Age, distribution, palaeoecology

Campanian II—Maastrichtian II (Mfolozi, Monzi, and Nibela outcrops,
Zululand and Richards Bay BH-9 borehole), Maastrichtian III (Agulhas Bank,

84 ANNALS OF THE SOUTH AFRICAN MUSEUM

sample 818). This species is widely distributed throughout the Campanian—
Maastrichtian rocks of south-east Africa. In Zululand, it is a consistent member
of the fauna, and in the Campanian II to early Campanian IV constitutes
between 10 and 70 per cent of the cytheracean element, and locally >15 per cent
of the total ostracod population (Fig. 38B—C). U. sacsi has been found in
ostracod assemblages 4b, 5a, 5b, 6-7, but is most abundant in 4b and 5,
indicating that it mainly inhabited quiet, low-energy, moderate to deep-water
(200-500 m) environments, probably with a preference for the mid-shelf (~200-
300 m) areas (Table 4). Its presence may be particularly correlated with strong,
open ocean influences.

Dutoitella gen. nov.

Derivation of name

In honour of Dr A. L. du Toit for his contributions to South African
geological knowledge and his fostering of the concept of Gondwanaland.

Type species
D. dutoiti sp. nov.

Diagnosis

Non-reticulate trachyleberid with connected AM and ventrolateral ridges or
clavae. The median area is occupied by a prominent nodose or dome shaped
SCT and an unconnected post-adjacent node or short nodose ridge.

Remarks

The hinge of Dutoitella varies from weakly hemiamphidont (D. dutoiti) to
strongly hemiamphidont (D. mimica), and whilst the MS in the genotype have
not been seen clearly, those of D. mimica show subdivided antennal and second
posterior scars.

Fig. 39. Comparative morphology of various genera and species of the. Unicapellinae subfam.
nov. A-D. Left valves. E-H. Right valves. A. Unicapella sacsi Dingle, 1980, SAM-K5740,
locality 20-7/1, Mfolozi River, Maastrichtian I]. B. Herrigocythere definita (Herrig, 1965),
genotype, GPIG 14/1, borehole Stubnitz 1/60 Rigen, East Germany, Lower Campanian.
C. Paleoabyssocythere cenozoica Benson, 1977, genotype, USNM 190285, DSDP site 21A-—3—4
50-56 cm, Thanetian (Globorotalia velascoensis Zone). D. Paleoabyssocythere cretacea Ben-
son, 1977, USNM 190168, DSDP site 21-6—4 53-59 cm, Campanian (Pseudotextularia elegans
Zone). E. Unicapella sacsi Dingle, 1980, genotype, SAM-K5610, BH9 88,39 m, Campanian II.
F. Herrigocythere definita (Herrig, 1965), genotype, GPIG 14/1, borehole Stubnitz 1/60 Rigen,
East Germany, Lower Campanian. G. Paleoabyssocythere cretacea Benson, 1977, USNM
190169, DSDP site 21-6-4 53-59 cm, Campanian (Pseudotextularia elegans Zone). H. Uni-
capella reticulata Dingle, 1980, SAM-K5614, BH9 88,76 m, Campanian II.
Scale bars all 200w.
Illustrations from following sources: A, this paper; B, Herrig (1965 fig. 1a); C, Benson (1977
pl. 2 fig. 7); D, Benson (1977 pl. 2 fig. 8); E, Dingle (1980 fig. 16A); F, Herrig (1965 fig. 1b);
G, Benson (unpublished data, with permission); H, this paper.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 85

Dutoitella is close to the early (Campanian—Maastrichtian) species of
Atlanticythere Benson, 1977. The two genera differ in the following points:
Atlanticythere does not have a continuous AM-ventrolateral ridge; the SCT and
post-adjacent node or nodes are less prominent in Atlanticythere; species of
Dutoitella are significantly smaller (lengths of holotypes): D. dutoiti (0,57 mm),
D. mimica (0,80 mm); A. maestrichtia (1,03 mm), A. murareticulata (0,87 mm),

U.sacsi

G IN

w,

H mm PC)
ES 20°)
)
Hove %

Pcretacea ———= U.reticulata

86 ANNALS OF THE SOUTH AFRICAN MUSEUM

A. prethalassia (0,87 mm). In addition, Benson (1977: 876-877) emphasizes the
‘almost equally rounded anterior and postericr marginal rims’ of Atlanticythere,
whereas although this condition is almost achieved in the LV of D. mimica, it is
not met with in the genotype or the RV of D. mimica (Fig. 43).

The stratigraphic and morphologic relationships between Unicapella and
Dutoitella suggest that the latter evolved from the former, probably by modifica-
tion of U. sacsi in Campanian III-IV times. The two genera have a common
basic geometry and morphology, but differ in their connection or separation of
the ventrolateral and AM ridges and the presence or absence of swellings
post-adjacent to the SCT.

Bate (in Bate & Bayliss 1969) recorded the genus (as Genus C. sp.) from
the middle to upper Maastrichtian of Tanzania.

Dutoitella, like its close relative Unicapella, is a relatively deep-water genus
(outer continental shelf, possibly as deep as 500 m).

Dutoitella dutoiti sp. nov.
Figs 40B-F, 43A

Derivation of name

In honour of Dr A. L. du Toit for his pioneering contribution to geological
knowledge in South Africa.

Holotype
SAM-K5742, LV, locality 20-1/2, Mfolozi River, Maastrichtian I

Paratypes

SAM-K5743, RV, locality 20-1/2, Mfolozi River, Maastrichtian I
SAM-K5744, LV, locality 20-1/2, Mfolozi River, Maastrichtian I
SAM-K5745, RV, locality 20-1/2, Mfolozi River, Maastrichtian I
SAM-K5746, carapace, locality 20—1/2, Mfolozi River, Maastrichtian I

Diagnosis

Species with two large, smooth, rounded median nodes.

Description

External features. In lateral view rounded AM and PM, the latter carrying
stubby, posteriorly deflected spines. DM and VM straight, converging slightly
posteriorly. Thick anterior marginal rim with small tubercles and spines which
leads posteriorly to a thick ventrolateral ridge which probably represents five
fused clavae. This ridge swings upwards posteriorly and ends at about three-
quarters length. There is a short keel on the ventral surface which just protrudes
beyond the VM. DM is surmounted by a weak ridge on which are situated four
perforate conuli with a small posterodorsal bulla. In the RV the anterodorsal
corner has a step, whereas in LV it is flaired slightly to form a small ear.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 87

Fig. 40. A. Unicapella reticulata Dingle, 1980, SAM-—K5614, BH9 88,76 m, RV, Campanian IT.
B-F. Dutoitella dutoiti gen. et sp. nov., locality 20-1/2, Mfolozi River, Maastrichtian I.
B. SAM-K5746, dorsal view carapace. C. Holotype, SAM-K5742, LV. D. SAM-KS5743,
RV. E. SAM-K5744, internal LV. F. SAM-—K5745, internal RV.
Scale bars all 100.

88 ANNALS OF THE SOUTH AFRICAN MUSEUM

Medianly there is a large smooth domed SCT, and a smaller smooth domed
node at about three-quarters length. In dorsa! view these give a characteristic
asymmetrically mammalate appearance to each valve. The surface is delicately
ornamented with fine fossae that are superimposed on a faint, coarse, first-order
reticulation pattern, particularly in the anterior region. There are numerous
small perforate nodes on the valve surface. Ventral surface has two longitudinal
ridges which frequently develop into small keels.

Internal features. Hinge amphidont, all the elements are smooth except PTE
which is rounded and weakly subdivided. There is a narrow, terminally widening
groove immediately above the hinge in RV. MS not seen. MA moderately wide,
no vestibule. MPC fine, slightly sinuous: fifteen anteriorly, ten posteriorly.

Remarks

D. dutoiti differs from D. mimica in possessing a node rather than a ridge
post-adjacent to SCT, and in having a continuous ridge rather than a nodose
lineament ventrolaterally (Fig. 43).

Dimensions (mm)

length height width
K5742 0,57 0,33
K5743 0,55 0.32
K5744 0,54 0,30
K5745 0,53 0,29
K5746 0,54 0,28
Other material 0,58 0,34
Other material O57 0,30

Age, distribution, palaeoecology

Campanian IV to Maastrichtian I] (Monzi, Mfolozi and Nibela areas
Zululand) (Fig. 38D-E). This species is consistently present in small numbers
(2-3% total population, 8-11% total cytheraceans) throughout its range, and
because it has been found evenly distributed in ostracod assemblages 5a, 5b,
6-7, it has been selected as a zonal marker for the upper part of the succession in
south-east Africa. The even distribution of D. dutoiti through the mid shelf
(?200 m) to deep water (?outer shelf/upper slope, >500 m) sedimentary en-
vironments indicates that it is not as environmentally bound as its relative U.
sacsi, which becomes relatively scarce in the deep-water facies. It is replaced in
the Maastrichtian III by D. mimica. -

Dutoitella mimica sp. nov.
Figs 37F, 41A-F, 42A-B, 43B, F, 44B

Genus C. sp. Bate, 1969 (in Bate & Bayliss 1969): 143, pl. 7 (fig. 15).

Trachyleberis schizospinosa Dingle 1971: 406-408, fig. 10. (Some of the paratypes designated
MG-4-1-+4 belong to D. mimica.)

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 89

Derivation of name
Latin mimicus (counterfeit): reference to similarity with Atlanticythere
maestrichtia Benson, 1977.

Holotype
SAM-K5747, LV, sample 818, Agulhas Bank, Maastrichtian II]

Paratypes

SAM-K5748, RV, sample 818, Agulhas Bank, Maastrichtian III
SAM-K5749, LV, sample 818, Agulhas Bank, Maastrichtian III
SAM-K5750, RV, sample 818, Agulhas Bank, Maastrichtian III
SAM-K5751, LV, sample 818, Agulhas Bank, Maastrichtian III
SAM-K5752, LV, JC-1 borehole, 1 811 m, Maastrichtian

Diagnosis
Species with nodose SCT, and three perforated nodes sub-adjacent to it.

Description

External features. AM broadly rounded with spinose and nodose rim. PM
and LV broadly rounded, in RV asymmetric and slightly acuminate, ventrally
spinose and nodose in both valves. DM straight with numerous short spines, and
a distinct step at the anterior cardinal angle in RV. VM slightly convex, more so
in the RV, the outline is partly obscured by a short keel on the ventral surface.
Lateral surface smooth or foveolate with fine ridges forming a loose network in
the central part of the valve and a first-order reticulation in the anterior area.
The dorsal area has a poorly developed lineament of stout, perforated spines,
several of which in the LV are deflected posteriorly. They form a weak bulla at
the posterior cardinal angle. There is an upward deflected ventrolateral line of
stout, perforate nodes which is continuous with the AM rim. The SCT is
prominent, domed and nodose and there is a short ridge of three perforate
nodes post-adjacent but not connected to it.

Internal features. Hinge hemiamphidont with fist-like PTE in RV. MS
consist of two rounded anterior scars and a vertical row of elongate adductors,
the second of which is subdivided. MA fairly narrow with small anterior and
posterior vestibules. Up to eighteen thin, straight anterior MPC; seven to ten
short, straight posterior MPC.

Remarks

Dingle (1971) included representatives of this species in his population of
Trachyleberis schizospinosa. They can be separated from it on the basis of valve
outline and surface ornamentation. D. mimica differs from D. dutoiti in the
morphology of the posteromedian elevation and ventrolateral ridge (Fig. 43).
Surface nodes on the specimens from JC-1 borehole are slightly more massive
than those from the Agulhas Bank, although this may in part be due to slight
abrasion.

90 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 41. Dutoitella mimica gen. et sp. nov. TBD 818 Alphard Formation, Agulhas Bank,
Maastrichtian III. A. Holotype, SAM-K5747, LV. B. SAM-K5748, RV. C. SAM-K5749,
internal LV. D. SAM-—K5750, internal RV. E. SAM-—K5751, dorsal view LV. F. SAM-—
K5749, MS LV.
Scale bars: F = 30u, others = 100w.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA QI

Fig. 42. Dutoitella mimica gen. et sp. nov., SAM-—K5752, JC-1 1811 m, LV, Maastrichtian.
A. Lateral view. B. Detail anterior area.
Scale bars: A = 100u, B = 30yu.

D. mimica superficially resembles Atlanticythere maestrichtia Benson, 1977,
but differs in the essential genetic characters (see remarks and designation of
Dutoitella).

Bate (in Bate & Bayliss 1969) recorded one valve of D. mimica (his Genus
C. sp.) from the Upper Cretaceous of Tanzania (1969, pl. 7 (fig. 15)) but there
is some confusion in his text as to the exact locality and age of the specimen.
One page 123 it is stated to have been found in the upper Campanian, but on
page 143, and according to the sample number (BM 108) it is said to be
mid-upper Maastrichtian. The latter will be assumed here because on fig. 7
sample 168 is clearly shown as Maastrichtian.

Dimensions (mm)

length height width
K5747 0,81 0,41
K5748 0,80 0,42
K5749 0,78 0,44
K5750 0,80 0,43
K5751 0,78 0,20
K5752 0,88 0,46

Age, distribution, palaeoecology

Maastrichtian III (sample 818, Agulhas Bank), late Campanian to lower
Maastrichtian (1 871-1 811 m), JC—1 borehole, and Maastrichtian (Tanzania).
The Agulhas Bank sedimentary environment is thought to represent an unusual
setting: a quiet, shallow-water environment on the outer edge of the continental
shelf or the uppermost continental slope. It is an important member of this
population, constituting 10 per cent of the total fauna. In the JC—1 borehole, D.
mimica occurs at level 1811 (Maastrichtian) with Bythocypris? and Cytherella
sp., and at level 1871 (Campanian) on its own.

9? ANNALS OF THE SOUTH AFRICAN MUSEUM

A
D.dutoiti
€
D
”) Ars
CeO e.
joe cs
re) > ea
Genus C A.maestrichtia

wee Pe OO

re} =)

‘J —

Fig. 43. Comparative morphology of various genera and species of the Unicapellinae subfam.
nov. All left valves. A. Dutoitella dutoiti gen. et sp. nov., genotype, SAM-—K5742, locality
20-1/2, Mfolozi River, Maastrichtian I. B. Dutoitella mimica gen. et sp. nov., SAM-—K5747,
TBD 818 Alphard Formation Agulhas Bank, Maastrichtian HII. C. Genus C Bate,
1969 = Dutoitella mimica, BM Io. 1304, sample BM168 Kilwa area Tanzania, Maastrichtian.
D. Atlanticythere maestrichtia Benson, 1977, genotype, USNM .190166, DSDP site 21-44
60-66 cm, Maastrichtian (top Rugotruncana-subcircumnoidifera Zone). E. Atlanticythere sp.
nov. USNM 190755, DSDP site 21—-7—4 53-59 cm, Campanian. F. Dutoitella mimica gen. et sp.
nov., SAM-K5752, JC-1 1811 m Maastrichtian.
Scale bars all 200y.
Illustrations from following sources: A, this paper; B, this paper; C, Bate & Bayliss (1969 pl. 7
fig. 15); D, Benson (1977 pl. 2 fig. 4); E, Benson (unpublished data, with permission); F, this
paper.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 93

Sry,
ew
=

Fig. 44. Muscle scars. A. Agulhasina quadrata Dingle, 1971, TBD 818 Alphard Formation.
Agulhas Bank, LV, Maastrichtian III. B. Dutoitella mimica gen. et sp. nov. SAM-K5S749,
TBD 818 Alphard Formation, Agulhas Bank, LV, Maastrichtian III.

Scale bars 60.

a B

Subfamily Trachyleberidinae Sylvester-Bradley, 1948

This subfamily is well represented by twenty-two species in the Campanian—
Maastrichtian rocks of south-east Africa, with the genera Haughtonileberis,
Oertliella, Trachyleberis, and Hermanites particularly important.

Genus Haughtonileberis Dingle, 1969

This genus is one of the most important of the Cytheracea taxa in the Upper
Cretaceous in south-east Africa. Its earliest records in this area are from the
Santonian, and Dingle (1976) has reported one species in the Eocene, but
recently Grosdidier (1979) has tentatively assigned several species to it from the
Cenomanian—Turonian of Gabon.

For most of the Campanian—Maastrichtian period in south-east Africa, the
genus had numerically passed its peak, although during the Campanian [ it is
represented by four species (Fig. 45): H. haughtoni, H. fissilis, H. vanhoepeni,
and H. nibelaensis. Only H. nibelaensis is confined to strata of Campanian—
Maastrichtian age, and only H. fissilis ranges up into the Maastrichtian. Numeri-
cally, the genus is most important in the Santonian (Dingle 1980) where it locally
reaches over 50 per cent of the total ostracod populations in two species (H.
haughtoni and H. fissilis). There is a steady, if erratic, decline in numbers
through the upper Santonian, so that, although a further species appears in the
uppermost Santonian (H. vanhoepeni), the Campanian opens with the genus
constituting about 20 per cent of the total ostracod population (Fig. 45E). This
abundance is maintained throughout Campanian I, where H. vanhoepeni
reaches its local acme (5-8%), but a rapid decline occurs across the Campanian
I-II boundary (to about 6%) and despite a temporary resurgence at the top of
Campanian II (~20%) where H. nibelaensis temporarily becomes important, the
genus is represented in small numbers by only two species in rocks younger than
Campanian II. Although ecological factors must have played a role in this
decline, the fact that both H. haughtoni and H. fissilis show considerable
environmental tolerance in the Santonian (Dingle 1980) suggests that their

94 ANNALS OF THE SOUTH AFRICAN MUSEUM

decline at the end of Campanian I was to a large extent a phylogenetic
phenomenon.

The genus is represented in the Maastrichtian by one record only: two
specimens of H. fissilis at locality 20 (Mfolozi River), but the closeness of H.
radiatus (Eocene, borehole JC—1, Dingle 1976) to H. fissilis indicates that the
latter species, or a development of it, survived locally through the rest of
Maastrichtian and Palaeocene times.

stage
locality
ostr.
assembl.

HAUGHTONILEBERIS

MAASTRICHTIAN

nibelaensis

haughtoni
fissilis
vanhoepeni

E

CAMPANIAN

SANTONIAN

Fig. 45. Distribution of Haughtonileberis in Campanian—Maastrichtian strata of Zululand and

Agulhas Bank. A. Total number of valves of H. haughtoni. B. Total number of valves of H.

fissilis. C. Total number of valves of H. vanhoepeni. D. Total number of valves of H.
nibelaensis. E. Haughtonileberis as percentage of total ostracod population.

Haughtonileberis haughtoni Dingle, 1969
Fig. 48E
Haughtonileberis haughtoni Dingle, 1969: 372-373, fig. 15; 1980: 39, fig. 21A-E.

Remarks

Of the two morphotypes recognized by Dingle (1980), only the elongate
form occurs in the Campanian I section of the Richards Bay BH-9 borehole.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 95

Here the species is at the top of its range, and occurs in only small numbers
compared to its importance in the Santonian strata.

Age, distribution, palaeoecology

Santonian II to Campanian I (Umzamba and Richards Bay BH-9 bore-
hole). H. haughtoni is an environmentally tolerant species, having been re-
corded from ostracod assemblages 1-3, and 4a (Dingle 1980). It is most
abundant in assemblage 2 and preferred shallow-water (<100 m), low-energy
environments with a restricted access to the open ocean (Table 4).

Haughtonileberis fissilis Dingle, 1969

Fig. 48F
Haughtonileberis fissilis Dingle, 1969: 374-375, fig. 16; 1976: 59, fig. 3(48); 1980: 39, fig.
22A-B.
Remarks

Of the two morphotypes recognized by Dingle (1980), only the more
coarsely reticulate variety occurs in the Campanian—Maastrichtian strata.

Age, distribution, palaeoecology

Santonian II to Maastrichtian IJ (Umzamba, and Richards Bay BH-9
borehole and Mfolozi and Nibela outcrops, Zululand). Although H. fissilis has
been found in ostracod assemblages 1-3, 4a, 4b, 5a, 5b, and 7, and is obviously
an environmentally tolerant species, it is found in relatively large numbers only
in assemblage 4a. On this evidence the species preferred low-energy, moderate-
depth (?100-200 m inner-mid shelf), open-water environments, suggesting that it
inhabited deeper and less-restricted areas than its close relative H. haughtoni
(Table 4).

Haughtonileberis vanhoepeni Dingle, 1980
Fig. 46A
Haughtonileberis vanhoepeni Dingle, 1980: 42-44, figs 22H, 23A-F.

Remarks

No important morphological variations have been recognized in this species
throughout its short range.

Age, distribution, palaeoecology

Uppermost Santonian HI to Campanian IV (Richards Bay BH-9 borehole
and Nibela Peninsula). Although two specimens have been found in assemblage
3, H. vanhoepeni is to all intents and purposes environmentally bound to
assemblage 4, with a clear preference for the conditions represented by sub-
assemblage 4a: low-energy, moderate-depth (?100-200 m inner-mid shelf),
open-water environments (Table 4).

96 ANNALS OF THE SOUTH AFRICAN MUSEUM

Haughtonileberis aibelaensis sp. nov.
Fig. 46B-F

Derivation of name

Locality of type.

Holotype
SAM-K5754, LV, locality 110-14, Nibela, Campanian II

Paratypes

SAM-K5755, RV, locality 110-14, Nibela, Campanian II
SAM-—K5756, LV, locality 110-14, Nibela, Campanian II
SAM-—KS5757, RV, locality 110-14, Nibela, Campanian II
SAM-—K5758, carapace, locality 110-14, Nibela, Campanian II

Diagnosis
Heavily calcified species with intercostal ornamentation that ranges from

coarsely reticulate to almost smooth. Anterior cardinal angle is prominently
rounded and the ventrolateral ridge is looped.

Description

Moderately large, heavily calcified species.

External features. In lateral view, AM is broadly rounded, with a somewhat
flaired appearance, PM is triangular in outline, DM and VM straight, posteriorly
tapering. Highest point over anterior cardinal angle, widest point in dorsal view
over SCT. Ornamentation is dominated by three narrow, sharp, longitudinal
ridges. The dorsal ridge is the shortest and is slightly convex, obscuring the
dorsal margin: the median ridge crosses a subdued SCT and typically splits
anteriorly: the ventral ridge forms a loop that widens posteriorly. A large
rounded eye spot lies on a short anterodorsal ridge that extends most of the way
round the anterior margin. Surface ornamentation within the species varies from
coarsely reticulate to almost smooth with ghost reticulation and small prominent
pustules.

Internal features. Hinge amphidont with weakly subdivided terminal ele-
ments in the RV. The RV ATE has a prominent anterior shoulder which fits
into a small socket in the LV. ME is smooth. MA moderately wide, up to
twenty-five long, fine anterior MPC, up to fifteen posterior MPC. MS consist of
a hooked (sometimes almost subdivided) anterior scar ‘and four elongate adduc-
tors, all lying in a shallow pit.

Remarks

This species is closest to H. fissilis with its split median rib, but has a more
symmetrical anterior outline, a more triangular posterior outline and a looped
ventrolateral ridge. H. nibelaensis is similar in lateral outline to H. vanhoepeni
but the latter has a different rib disposition and is a smaller species.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 97

Fig. 46. A. Haughtonileberis vanhoepeni, SAM-K5632, BH9 100,0 m, RV, Campanian I.
B-F. Haughtonileberis nibelaensis sp. nov., locality 110-14, Nibela Peninsula, Campanian II.
B. SAM-K5758, dorsal view carapace. C. Holotype, SAM-K5754, LV. D. SAM-K5755, RV.
E. SAM-K5756, internal LV. F. SAM—K5757, internal RV.
Scale bars all 100.

98 ANNALS OF THE SOUTH AFRICAN MUSEUM

Dimensions (mm)

lengih height width
K5754 0,65 0,31
RS 755 0,58 0,30
K5756 0,62 0,34
K5757 0,67 0,32
K5758 0,62 0:25
Other material 0,65 WS |
Other material O72 0,30
Other material 0,61 032
Other material O37 O25

Age, distribution, palaeoecology

Campanian I to Campanian II (Richards Bay BH-9 borehole and Nibela
Peninsula). Although H. nibelaensis occurs in assemblages 4a, 4b, and Sa (Fig.
45D) it has a strong preference for 4b, indicating that it is to all intents and
purposes environmentally bound to conditions of low energy, moderate depth
(?200 m inner-mid shelf) with open-water connections. H. nibelaensis has the
deepest water preference of all the known species of the genus (Table 4).

Genus Oertliella Pokorny, 1964

This genus occurs sporadically in the late Cretaceous of south-east Africa,
and is numerically important (10-20% of total cytheraceans) in Campanian I and
II and Maastrichtian I and II strata (Fig. 47E). It is represented by four species:
O. pennata, O. sp. A, O. africana, and O. maastrichtia of which the last two are
confined to Campanian—Maastrichtian, and the first two to Santonian—Campa-
nian strata.

Oertliella pennata Dingle, 1980
Fig. 48A

Acanthocythereis? aff. A. horridula (Bosquet, 1854), Dingle, 1969: 378-380, fig. 19.
Oertliella pennata Dingle 1980: 46-49, fig. 26A—-E.

Remarks
No additional material is available for this species beyond that already
recorded by Dingle (1980).

Age, distribution, palaeoecology

Santonian III to Campanian II (Richards Bay BH-9 borehole), Santonian
III (bed Pi 3) (Umzamba). Two valves of O. pennata were found in ostracod
assemblage 2, but the species is consistently present only in assemblages 3, 4a,
and 5a, with a maximum in 4a indicating a preference for low-energy, moderate-
depth (?100—200 m, inner-mid shelf) open-water conditions (Fig. 47A, Table 4).
Because of its apparent tolerance over various environments in the inner to mid
shelf locations, the disappearance of O. pennata during Campanian II times is
probably phylogenetically significant.

\O
\o

CAMPANIAN AND MAASTRICHTIAN OSTRACODA

stage
assembl.

<
= .
co | locality

ostr.

OERTLIELLA

MAASTRICHTIAN

ed
uo
wo
uo

TT 2 ©
S ss
e < = =
® joe - =)
a 77) cs ies
i a
Ls]
Ly]
£
z
=
P4
<q
| Yy 4a
=
be B Cc
= 5
IIISANTONIAN 5 3 5 15

Fig. 47. Distribution of Oertliella in Campanian—Maastrichtian strata of Zululand and Agulhas

Bank. A. Total number of valves of O. pennata. B. Total number of valves of O. sp. A.

C. Total number of valves of O. africana. D. Total number of valves of O. maastrichtia.
E. Oertliella as percentage of Cytheracea.

Oertliella sp. A
Fig. 48B

Oertliella sp. A Dingle, 1980: 50-52, fig. 26F.

Remarks

Three valves of this species were recorded by Dingle (1980) in the Richards
Bay BH-9 borehole. No additional specimens have come to light, so their
taxonomic position remains uncertain.

Age, distribution, palaeoecology

Santonian III to Campanian II (Richards Bay BH-9 borehole). Too few
valves are available for an assessment of the species’ palaeoenvironmental

preferences, though it appears to be confined to moderate-water depth (Fig.
47B, Table 4).

L100 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 48. A-D. Oertliella. A. O. pennata Dingle, 1980, SAM-—-K5759, BH9 102,6m, LV,
Campanian I. B. O. sp. A, SAM-K5760, BH9 106,0 m, LV, Campanian I. C. O. africana
Dingle, 1980, SAM-K5648, BH9 82,03 m, LV, Campanian II. D. O. africana, SAM-K5761,
locality 20-1/1, Mfolozi River, RV, Maastrichtian I. E. Haughtonileberis haughtoni Dingle,
1969, SAM-K5627, BH9 92,27 m, LV, Campanian I. F. Haughtonileberis fissilis Dingle, 1969,
SAM-KS5762, locality 20-1/3, Mfolozi River, LV, Maastrichtian I.
Scale bars all 100y.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 101

Oertliella africana Dingle, 1980
Fig. 48C-—D
Oertliella africana Dingle, 1980: 49-50, figs 26G, 27A-E.

Remarks

Two specimens closely comparable to the lower Campanian material from
the Richards Bay BH-9 borehole (Dingle 1980) have been found at locality 20
(Mfolozi River) in Maastrichtian I sediments. They differ only in possessing a
double-bladed spine post-adjacent to the weak eye spot, a somewhat more
inflated ventral outline and an additional spine immediately below the postero-
dorsal margin spine (compare C and D in Fig. 48). In addition they are slightly
smaller than the types described by Dingle (1980).

Dimensions (mm)

length height
K5761 0,61 0,31
Other material 0,59 0333

Mean of four specimens quoted by Dingle (1980) from the BH-9 borehole:
length 0,73 mm and height 0,34 mm.

Age, distribution, palaeoecology

_ Accepting that the two specimens mentioned above fall within the definition
of the species, O. africana ranges Campanian I to Maastrichtian I (Richards Bay
BH-9 borehole, and Mfolozi and Nibela outcrops) (Fig. 47C). It has been found
in ostracod assemblages 4a, and Sa, 5b, and it is significant that the Maastrich-
tian occurrence is in assemblages assigned to 5b. It seems likely that this species’
preference is for the environments represented by assemblage Sa, viz: low-
energy, Open-water, moderate-depth (?200—300 mid-outer shelf) environments.
On this evidence it occupied slightly deeper-water conditions than O. pennata
(Table 4).

Oertliella maastrichtia sp. nov.
Fig. 49A-E
Derivation of name

Age of largest populations.

Holotype
SAM-K5763, LV, locality 20-7/1, Mfolozi River, Maastrichtian II

Paratypes

SAM-K5764, RV, locality 20-7/1, Mfolozi River, Maastrichtian II
SAM-K5765, LV, locality 20-7/1, Mfolozi River, Maastrichtian II
SAM-K5766, RV, locality 20—-7/1, Mfolozi River, Maastrichtian II

102 ANNALS OF THE SOUTH AFRICAN MUSEUM

of

Fig. 49. Oertliella maastrichtia sp. nov., locality 20-7/1, Mfolozi River, Maastrichtian II.
A. Holotype, SAM-K5763, LV, B. SAM-K5764, RV. C. SAM-K5765, internal LV.
D. SAM-K5766, internal RV. E. SAM—K5765, MS LV.

Scale bars: E = 10y, others = 100y.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 103

Diagnosis
Species with large turreted eye spot and pennate spine at posterior end of
dorsal ridge.

Description

External features. In lateral view quadrate. Symmetrically rounded, spinose
AM. Slightly acuminate, spinose PM, with pronounced posterodorsal concavity.
DM straight, VM gently convex. Surface strongly reticulate with numerous
conjunctive spines, and occasional normal pore openings within fossae. Narrow,
well-developed anterior marginal rim with numerous stubby spines. There is a
well-developed ventrolateral ridge which is upcurved at its anterior and posterior
ends, and is surmounted by numerous large spines. The dorsal ridge is short, set
away from the margin and carries three spines which lie anterior of a large
pennate bulla at the posterior termination. There is a prominent spinose SCT,
and the eye spot is large, spherical, and perched prominently at the anterior
cardinal angle on a spinose turret.

Internal features. Hinge amphidont with the RV PTE distinctly subdivided:
other elements are smooth. MA narrow, no vestibule. MPC straight: seventeen
anteriorly, seven posteriorly. MS lie in a deep sub-central pit and consist of four
ovate adductors, the middle two being particularly elongate, and a V-shaped
anterior scar. The top adductor is dorsally indented.

Remarks

~ QO. maastrichtia is very close to O. exquisita Bate from the Campanian of
Western Australia. The new species differs on the following points: it is
somewhat more elongate, and has a distinct posterodorsal concavity which O.
exquisita lacks; the adductor MS of the two species are different in shape; the
RV PTE of O. maastrichtia is clearly subdivided whereas it is only weakly so in
the Australian form; O. exquisita lacks the pennate spine at the posteror end of
the dorsolateral ridge. Although O. exquisita has not been proven from Maas-
trichtian strata, it is possible that the two species have very similar time ranges.
Within the South African context, it appears that O. maastrichtia evolved from
O. pennata (Santonian—Campanian) by a process of ornament modification and
slight changes in outline. It is an important element of the cytheracean popula-
tion (>10%, up to 20%) in Maastrichtian times, and locally constitutes 10 per
cent of the total ostracod fauna.

Dimensions (mm)

length height
K5763 0,58 033
K5764 0,65 0,34
K5765 0,61 0,34
K5766 0,66 0,35
Other material 0,64 0,36
Other material 0,65 0,36

Other material 0,65 0,34

104 ANNALS OF THE SOUTH AFRICAN MUSEUM

Age, distribution, palaeoecology

Campanian IV to Maastrichtian [I (Monzi, Mfolozi and Nibela outcrops,
Zululand) (Fig. 47D). O. maastrichtia occurs in ostracod assemblages 4b, 5a, 5b,
6-7, with a clear preference for the conditions represented by 7, viz.: deep
(?2>500 m, outer shelf/upper continental slope), oceanographically stable en-
vironments. It therefore inhabited distinctly deeper water than O. africana
(Table 4).

Genus Hermanites Puri, 1955

This genus is represented by two species (H. kennedyi, H? arcus), in the
Campanian—Maastrichtian strata of south-east Africa, one of which (H. ken-
nedyi), for much of its range, constitutes an important element of the ostracod
fauna: 8-10 per cent total population, and 30—40 per cent cytheracean popula-
tion (Fig. 50). Both species appear to have favoured moderate to deep water
environments.

>
~
e =
a s
o (s) GS
oe
a &

ostr.
assembl.

®
2 HERMANITES C’'THEREIS GIBBERLEBERIS

kennedyi
klingeri

La
)
s
= 7)
|
°
a

CAMPANIAN

SANTONIAN

Fig. 50. Distribution of Hermanites, Cythereis, and Gibberleberis in Campanian—Maastrichtian

strata of Zululand and Agulhas Bank. A. Total number of valves of Hermanites kennedyi.

B. Total number of valves of Hermanites? arcus. C. Hermanites kennedyi as percentage of

Cytheracea. D. Total number of valves of Cythereis klingeri. TE. Cythereis klingeri as

percentage of Cytheracea. F. Total number of valves of Gibberleberis elongata. G. Total
number of valves of Gibberleberis sp. A.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 105

Hermanites kennedyi Dingle, 1980
Fig. 51A-B
Hermanites kennedyi Dingle, 1980: 44—46, figs 22C—G, 24A-F.

Remarks

H. kennedyi is a distinctive, relatively environmentally tolerant, and locally
abundant species. For this reason it has been selected as a zonal fossil for the
ostracod zonation scheme proposed herein. The specimens that occur in the
Richards Bay BH-9 borehole and at outcrops in Zululand show no significant
morphological variations throughout the species’ range, but the representatives
from Igoda tend to be slightly less coarsely reticulate (particularly the SCT) and
have less flared dorsal and ventral ridges. These specimens are, however, worn
and have considerable secondary crystalline calcite overgrowth which might have
reduced the angularity of the positive surface features.

Age, distribution, palaeoecology

Campanian II to Campanian V (Richards Bay BH-9 borehole, and Monzi
and Nibela Peninsula, Zululand), late Campanian/early Maastrichtian (Igoda).
H. kennedyi is a member of ostracod assemblages 4b, 5a, 5b, and 6 with a strong
preference for assemblage 5 (both a and b) where it locally forms between 30
and 40 per cent of the cytheracean element (Fig. SOA, C, Table 4). On this
evidence it evidently preferred low-energy, moderate-depth (?200—-300 m, mid-
outer shelf) environments.

At Igoda, H. kennedyi is a minor element of the ostracod population (4%)
and is relatively unimportant (7%) within the cytheraceans, suggesting that it
was not in a preferred environment during the deposition of the Igoda assem-
blage.

Hermanites? arcus sp. nov.
Fig. 51C
Derivation of name

Latin arcus (bow): reference to bow-shaped combination of dorsal and
median lateral ridges.

Holotype
SAM-K5768, RV, locality 20—1/3, Mfolozi River, Maastrichtian I.

Diagnosis
Species with bow-shaped combination of dorsal and median lateral ridges.

Description

External features. In lateral view, elongate with broadly rounded and
weakly spinose AM, asymmetrically acuminate PM that is deflected ventrally
and bears spines posteroventrally. DM and VM straight, converging slightly

106 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 51. A. Hermanites kennedyi Dingle, 1980, SAM-K5637, BH9 82,03 m, LV, Campanian
II. B. Hermanites kennedyi Dingle, 1980, SAM-K5767, Igoda Formation, Igoda estuary, LV,
late Campanian/early Maastrichtian. C. Hermanites? arcus sp. nov., holotype, SAM—K5768,
locality 20-1/3, Mfolozi River, RV, Maastrichtian I. D. Hermanites? cf. H? arcus sp. nov.,
SAM-K5769, Igoda Formation, Igoda estuary, RV, late Campanian/early Maastrichtian.
E. Hermanites? cf. H? arcus sp. nov., SAM-K5770, Lower Quarry Needs Camp, RV, late
Campanian/early Maastrichtian. F. Rayneria nealei Dingle, 1980, SAM-—K5653, BH9 106,0 m,

LV, Campanian I.

Scale bars all 100w.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 107

posteriorly. Dorsal outline partially hidden by curved, ponticulate dorsolateral
ridge that runs from posterodorsal angle to just behind the eye spot. Median
ridge is connected to dorsal ridge at their respective posterior ends, and runs
diagonally across the valve surface to anterior of the SCT. There is a curved
ventral ridge which parallels the median ridge and connects to the AM ridge.
The latter extends to the prominent eye spot, behind which it forms a small ear.
Valve surface is coarsely reticulate all over. Fossae are polygonal and irregularly
shaped, often partially rounded.
No internal features seen, hence provisional generic placement.

Remarks

The surface features of H? arcus bear a striking resemblance to those of H.
kennedyi. The two species differ in the nature and disposition of the median-
lateral ridge: in arcus it is diagonal and reaches the dorsal margin, whereas in
kennedyi it is vestigial and roughly parallel to the line of greatest length.

Dimensions (mm)

length height
K5768 0,56 0,29
Other material >0,50 0,30

Age, distribution, palaeoecology

Maastrichtian I to Maastrichtian II (Mfolozi River, Zululand). H? arcus is a
rare species that in Zululand is restricted to assemblage 7, suggesting that it was
environmentally bound by deep-water (?>500, outer shelf/upper slope),
oceanographically stable conditions (Fig. 50B, Table 4). The temporal and
environmental range of this species may be greater than the Zululand popula-
tions suggest (see below).

Hermanites? cf. H?. arcus sp. nov.
Fig. 51D-E

Remarks

Two specimens showing great similarity to H?. arcus were recovered from
Igoda (SAM-KS769) and Needs Camp (SAM-KS5770). Shell abrasion and secon-
dary calcite crystallization prevent a detailed comparison of surface features, but
the only noticeable difference with the Zululand specimens is the slightly weaker
upward curvature of the ventrolateral ridge, and a more pronounced VM
concavity immediately behind the SCT.

Dimensions (mm)
length height

K5769 0,70 0,34
K5770 0,68 0,35

108 ANNALS OF THE SOUTH AFRICAN MUSEUM

Age, distribution, palaeoecology

If these specimens are conspecific with H?. arcus then they probably extend
the known range of this species from late Campanian/early Maastrichtian (Igoda
and Needs Camp) to Maastrichtian II (Zululand). They also indicate that it was
capable of surviving in very shallow (20 m) moderate to high-energy environ-
ments (Needs Camp), although even in the slightly deeper (100-200 m) water of
the Igoda Formation it occurs only in trace numbers.

Genus Rayneria Neale, 1975
Rayneria nealei Dingle, 1980
Fig. 51F
Rayneria nealei Dingle, 1980: 55-57, figs 28E-F, 29A-F, 30G.

Remarks

Only two specimens of this species have been recovered from strata younger
than Santonian (Richards Bay RB-9 borehole) where it is at the top of its local
range.

Age, distribution, palaeoecology

Santonian II to Campanian I. R. nealei occurs in ostracod assemblages 1-3
and 4a in which it is equitably, though sparsely distributed. In this respect its
disappearance just above the base of Campanian I may be a useful biostrati-
graphic marker horizon in south-east Africa since it does not appear to be
environmentally bound, except that all its occurrences seem to be in water no
deeper than about 100 m.

Genus Cythereis Jones, 1849

This genus is represented by two species in the Campanian—Maastrichtian of
south-east Africa: C. transkeiensis and C. klingeri. Both range from Santonian to
Campanian/Maastrichtian, but whereas C. transkeiensis is known from Igoda,
Umzamba, and the base of the Richards Bay BH-9 borehole, C. klingeri
appears to be restricted to the northern area: BH—9 borehole and Mfolozi and
Nibela outcrops in Zululand. In their respective geographical areas, each species
is an important member of the cytheracean faunas, C. klingeri especially so.

Cythereis klingeri Dingle, 1980
Fig. 52A
Cythereis klingeri Dingle, 1980: 34-38, figs 18B-F, 19 A-F.

Remarks

The relatively wide range of intraspecific morphological variation noted by
Dingle (1980) in the Santonian—Campanian I rocks of the Richards Bay BH-9
borehole also occurs in the younger outcrops in Zululand, although no environ-
mentally related trends can be identified.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 109

Age, distribution, palaeoecology

Santonian II to Maastrichtian II. C. klingeri appears to be restricted to the
northern area: Richards Bay BH-9 borehole and Mfolozi and Nibela outcrops,
Zululand. Along with Brachycythere longicaudata, it is the most environmentally
tolerant of the cytheracean taxa, occurring in all the ostracod assemblages so far
identified (Table 4). However, it is most abundant in assemblages 3 and 6,
particularly the former, where in the upper Santonian III of the Richards Bay
BH-9 borehole it locally constitutes over 30 per cent of the total ostracod
population. Within the Campanian—Maastrichtian strata it is most abundant in
Campanian III (assemblage 6) where it locally constitutes >20 per cent cythera-
cean element (Fig. 50D-E, Table 4). From our other evidence, the two
sedimentary environments represented by assemblages 3 and 6 do not appear to
have much in common (shallow, <100 m, and deep ?>500 m, respectively) and
the reason for the observed distribution is not known.

Cythereis transkeiensis Dingle, 1969
Fig. 52B-C
? Cythereis ornatissima Reuss, 1846, var. reticulata Jones & Hinde, 1890, Chapman, 1904: 234.
Cythereis transkeiensis Dingle, 1969: 377-378, fig. 18; 1980: 34, fig. 18A.

Remarks

This species has been found in Campanian—Maastrichtian strata only at
Igeda (7% of total fauna) and Umzamba (one fragment). The Igoda population
shows some small morphological differences to topotypic material and to the
Santonian II specimens in the Richards Bay BH-9 borehole: its median and
ventrolateral longitudinal ridges have somewhat sharper crests, the post-anterior
cardinal angle depression on the DM is less pronounced, and the median
depression post-adjacent to SCT is not well developed. In addition, the Igoda
valves are somewhat more heavily calcified, but none of these differences is as
great as the variations found within individual populations of C. transkeiensis.

Age, distribution, palaeoecology

?Santonian II to Campanian I (Umzamba), Santonian II (Richards Bay
BH-9 borehole), late Campanian/early Maastrichtian (Igoda). C. transkeiensis
appears to be restricted to Igoda, Umzamba and Richards Bay, with the latter
locality marking the northern limit of its range. Within the Santonian, this
species is associated with shallow, relatively high-energy sedimentary environ-
ments, but its presence at Igoda indicates that it also inhabited low to moderate-
energy, moderate-depth (?100 m) sedimentary environments, suggesting that,
like its northern counterpart C. klingeri, C. transkeiensis was environmentally
tolerant.

Genus Gibberleberis Dingle, 1969

Gibberleberis is one of the minor genera in the south-east African Upper
Cretaceous, and has been recorded at only three horizons above the Santonian—

110 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 52. A. Cythereis klingeri Dingle, 1980, SAM-K5617, BH9 82,03 m, LV, Campanian II.
B-C. Cythereis transkeiensis Dingle, 1969, Igoda Formation Igoda estuary, late Campanian/
early Maastrichtian. B. SAM-K5771, LV. C. SAM-K5772, RV. D. Gibberleberis elongata
Dingle, 1980, SAM-K5773, BH9 88,76m, RV, Campanian II. E. Gibberleberis sp. A,
SAM-K5774, locality 110-14, Nibela Peninsula, RV, Campanian II.
Scale bars all 100.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA had

Campanian boundary, where it is represented by two species, G. elongata and
G. sp. A (Fig. 50F-G). Although never abundant, Gibberleberis consistently
occurs in the Santonian strata at Umzamba and in the Richards Bay BH-9
borehole (Dingle 1969, 1980), and appears to be at the upper limit of its range in
the lower Campanian.

Gibberleberis elongata Dingle, 1980
Figs 52D, 53A
Gibberleberis elongata Dingle 1980: 57-59, figs 30E-F, 31A.

Remarks

A rare species first recorded from the Richards Bay BH-9 borehole and
since located at one horizon (at locality 110, Nibela Peninsula) at outcrop.

Age, distribution, palaeoecology

Santonian III to Campanian II (Richards Bay BH-9 borehole, Nibela
Peninsula, Zululand). G. elongata occurs in trace numbers in ostracod assem-
blages 2, 2-3 transition, 5a and 4b (Fig. 50F). It is not possible to determine a
preference because of the small numbers of specimens found at each horizon,
although the species seems to be restricted to water shallower than 300 m.

G.elongata

G.sp.A

Fig. 53. Sketches of Gibberleberis species showing main rib patterns. A. G. elongata Dingle,
1980, SAM-K5660, BH9 124,0m, RV, Santonian III. B. G. sp. A, SAM-K5774, locality
110-14, Nibela Peninsula, RV, Campanian II.

Scale bars 200.

Gibberleberis sp. A
Figs 52E, 53B

Remarks

One carapace of this species was found at locality 110 on the Nibela
Peninsula. It resembles G. africanus Dingle, 1969, but differs in lacking a rib
along the dorsal edge of the dorsolateral hump, in having a slightly different
disposition of lateral ribs, in having a less coarsely reticulate surface ornamenta-
tion, and in lacking a well-defined posteroventral caudal process.

112 ANNALS OF THE SOUTH AFRICAN MUSEUM

Dimensions (mm)
length height
K5774 0,50 0,28
Age, distribution, palaeoecology

Uppermost Campanian II (Nibela Peninsula, Zululand). It occurs in ostra-
cod assemblage 4b, indicating that it was deposited in quiet, moderate-water
depths (?200 m inner-mid shelf). This is the youngest record of the genus
Gibberleberis so far.

Genus Australileberis Dingle, 1976

Records of this genus have previously been confined to the Eocene: JC-1
borehole off Natal; and outcrops on the Agulhas. Bank (Dingle 1976).

Australileberis stangerensis sp. nov.
Fig. 54A-B

Derivation of name

Locality of JC—1 borehole, east of Stanger, Natal.

Holotype
SAM-K5775, RV, JC—1 borehole, 1 676 m, Maastrichtian.

Fig. 54. Australileberis stangerensis sp. nov., SAM-K5775, JC-1 1676 m, Maastrichtian.
A. Lateral view RV. B. Dorsal view RV.
Scale bars 100w.

Diagnosis
Species with thick AM rim, and sharp median longitudinal ridge.
Description

External features. In lateral view elongate, broadly rounded, weakly spinose
AM, acuminate PM. Anterior area strongly depressed. DM and VM straight,
converging posteriorly. Surface ornamented with three prominent longitudinal

CAMPANIAN AND MAASTRICHTIAN OSTRACODA v8) Ee,

ridges. Dorsal ridge is shortest and is medianly deflected behind large eye spot.
Median ridge has a sharp crest and continues across the SCT as a narrow feature
to a prominent pustule. Ventral ridge is also sharp crested, deflected medianly at
its anterior end. There is a thick AM rim carrying small pustules, which passes,
via a small flared anteroventral flange, to a narrow VM ridge. Intercostal areas
smooth, except for numerous perforate pustules.

Internal features. Hinge amphidont, all elements appear smooth, but may be
slightly worn. MA wide with at least twenty-five long, thin anterior MPC, the
anterodorsal ones of which curve upwards. Posterior MPC and MS not seen.

Remarks

A. stangerensis is very close, and presumably ancestoral to A. hieroglyphica.
The two species can be distinguished by the presence of a thick AM rim in A.
stangerensis, which contrasts with the unrimmed compressed anterior area of A.
hieroglyphica. Their surface ornamentation is similar, although the new species
is closer to the Agulhas Bank populations of A. hieroglyphica than it is to the
typically more heavily calcified varieties in JC—1 borehole. The holotype of A.
stangerensis is significantly smaller than typical examples of the Tertiary species.

Dimensions (mm)

length height width
K5775 0,80 0,40 0,19

Age, distribution, palaeoecology

Maastrichtian (level 1 676m), JC—1 borehole. The horizon at which A.
stangerensis occurs is within the mid-Maastrichtian charophytes-rich section of
the borehole (Table 10). Other ostracod species are rare in this facies, which is
thought to represent a level at which there were particularly large influxes of
fluvial debris on to the Tugela delta top during local lower sea-level stands. This
contrasts with the occurrence of A. hieroglyphica in the overlying Palaeogene.
Here, this species is absent from the Palaeocene charophytes-bearing strata, and
appears only in the normal marine Eocene sediments.

Genus Trachyleberis Brady, 1898

This genus is represented by three species in the Campanian—Maastrichtian
strata of south-east Africa (Fig. 55): one long ranging (7. zululandensis), and
two short ranging (T. minima and T. schizospinosa), and as far as is known the
genus does not occur in rocks older than Campanian in this region. In the
Campanian I to III it occurs sporadically, though locally constituting up to 10
per cent of the cytheracean population, but in the younger rocks it is consistently
present, albeit in small numbers (2-10% of the cytheracean population) (Fig.
55). In the one sample that was available from the Maastrichtian III (on the
Agulhas Bank), the genus (7. schizospinosa) constituted a remarkable 40 per
cent of the cytheracean population (33% of the total ostracod population).

114 ANNALS OF THE SOUTH AFRICAN MUSEUM

assembl.

-
~
n
ie]

locality
d

a
oO
©
~
“

2 TRACHYLEBERIS PARVACYTHEREIS

d 3 70
schizospinosa

MAASTRICHTIAN

z
<q
- Ls}
< E
a (=
s =
= £
oO

Fig. 55. Distribution of Trachyleberis and Parvacythereis in Campanian—Maastrichtian strata
of Zululand and Agulhas Bank. A. Total number of valves of Trachyleberis zululandensis.
B. Total number of valves of Trachyleberis minima. C. Total number of valves of Trachyleberis
schizospinosa. D. Trachyleberis as percentage of Cytheracea. E. Total number of valves of
Parvacythereis monziensis. F. Total number of valves of Parvacythereis spinosa.

Trachyleberis zululandensis Dingle, 1980
Figs 56A, 57B;, D
Trachyleberis zululandensis Dingle, 1980: 52-54, fig. 28A—C.

Remarks

In his original description, Dingle (1980) recorded the species from two
horizons in the Richards Bay BH-9 borehole. Considerably more material has
now been obtained from outcrops in Zululand and one point of uncertainty with

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 115

regard to a possible assignment to the genus Matronella can be cleared up. The
MS of the species, seen for the first time, definitely preclude such an assignment
as they show a hooked anterior and four complete posterior adductors (Fig.
57D). The second adductor is, however, medianly constricted. There is no
similarity to the ‘splintered’ (élatées) arrangement described by Damotte (1974)
in her diagnosis of Matronella. A further point that should be emphasized is the
slight difference in shape between LV and RV. The former are more rectangular
than the somewhat triangular-shaped RV (compare Figs 56A and 57B).

Age, distribution, palaeoecology

Campanian I to Maastrichtian II (Richards Bay BH-9 borehole, Mfolozi
and Nibela outcrops, Zululand). T. zululandensis is an environmentally tolerant
species, having been found in ostracod assemblages 4a, 4b, 5a, 5b, 6-7 (Fig.
55A, Table 4), but, as suspected by Dingle (1980), it has a distinct preference
for deep-water environments, particularly that represented by assemblage 6:
deep water (>500 m outer shelf/upper slope), with oceanographically unstable
conditions.

Fig. 56. Trachyleberis. A. T. zululandensis Dingle, 1980, SAM-K5776, locality 113-1, Nibela
Peninsula, LV, Campanian IV. B. T. minima Dingle, 1980, SAM-K5654, BH9 110,0 m, RV,
Campanian I. C-D. T. schizospinosa Dingle, 1971, TBD 818 Alphard Formation, Agulhas
Bank, Maastrichtian III. C. SAM-—K5777, LV. D. SAM-K5778, RV.
Scale bars all 100y.

116 ANNALS OF THE SOUTH AFRICAN MUSEUM

Trachyleberis minima Dingle, 1980
Fig. 56B
Trachyleberis minima Dingle, 1980: 54, fig. 28D.

Remarks
No further specimens of this rare species have been recorded since the
original description for the Richards Bay BH-9 borehole.

Age, distribution, palaeoecology

Lowermost Campanian I (Richards Bay BH-9 borehole). The only record
has been from ostracod assemblage 5a, which is reckoned to represent quiet
moderate depths (?200-300 m mid-outer shelf).

Trachyleberis schizospinosa Dingle, 1971
Figs 56C-D, 57A
Trachyleberis schizospinosa Dingle 1971: 406-408, figs. 9-10, pl. la.

Remarks
Although we have no additional data on this species, SEM photographs are
reproduced herein to supplement the original descriptions. In comparison with

T.schizospinosa

schizospinosa | al

. \
0,5 is o
e pr
o : f o..~
2} zululandensis C23"

—@ ee e; os

0,7 length 0,9

Fig. 57. Trachyleberis. A-B. Sketches showing valve outline and arrangement of spines and
nodes. A. T. schizospinosa Dingle, 1971, SAM-—K5778, TBD 818 Alphard Formation, Agulhas
Bank, RV, Maastrichtian III. B. 7. zululandensis Dingle, 1980, SAM-K5779, locality 20-5,
Mfolozi River, RV, Maastrichtian I. C. Length v. height scattergram of adult T. zululandensis
(dots) and T. schizospinosa (circles). D. T. zululandensis Dingle, 1980, SAM-—K5780, locality
20-5, Mfolozi River, MS RV, Maastrichtian I.
Scale bars: A-B = 200u, D = 30wn.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA Miz

T. zululandensis, T. schizospinosa is seen to be plumper, and consistently
possesses a ring or cluster of small spines midway between, and slightly posterior
to, the large posterodorsal and posteroventral spinose processes (Fig. 57A, B).
It is also a considerably larger species (Fig. 57C).

Age, distribution, palaeoecology

Maastrichtian III (sample 818 Agulhas Bank) which is thought to represent
an unusual setting: a quiet, shallow-water environment on the outer edge of the
continental shelf or on the uppermost continental slope. In this fauna, T.
schizospinosa constitutes 33 per cent of the total ostracod population.

Genus Parvacythereis Grundel, 1973

Griindel (1973) quoted a range of ?Cenomanian to upper Tertiary for this
genus, and suggested that it evolved from Cornicythereis. To our knowledge, the
two species found in the Campanian—Maastrichtian of south-east Africa are the
first record of Parvacythereis from the southern hemisphere, although two forms
described by Bate (in Bate & Bayliss 1969) and Ducasse & Grekoff (1976) from
the east Africa area may belong to it.

In south-east Africa, the genus constitutes a minor (up to 7%) but charac-
teristic element in the Maastrichtian cytheracean populations.

Parvacythereis monziensis sp. nov.
Figs 58A-E, 60A
Derivation of name

Locality of type, vicinity of Monzi village.

Holotype
SAM-K5781, RV, locality 20-7/1, Mfolozi River, Maastrichtian II

Paratypes

SAM-K5782, carapace, locality 20-7/1, Mfolozi River, Maastrichtian II
SAM-K5783, LV, locality 20-5, Mfolozi River, Maastrichtian I
SAM-K5784, RV, locality 20-3, Mfolozi River, Maastrichtian I

Diagnosis

Species with recurved ventrolateral rib and a round SCT.

Description

External features. In lateral view AM broadly rounded, PM bluntly triangu-
lar. DM and VM almost straight, converging posteriorly. Strong, narrow AM
rim which commences over prominent eye spot, and is continuous with a VM rib
that recurves at a posteroventral process and runs forward to below the
prominent, rounded SCT. The SCT possesses a small, curved anteroventrally

118 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 58. Parvacythereis monziensis sp. nov. A. SAM-K5782, locality 20-7/1, Mfolozi River,
LV, Maastrichtian II. B. Holotype, SAM-K5781, locality 20-7/1, Mfolozi River, RV, Maas-
trichtian II. C. SAM-KS5783, locality 20-5, Mfolozi River, internal LV, Maastrichtian I.
D. SAM-K5784, locality 20-3, Mfolozi River, internal RV, Maastrichtian I. E. Holotype,
SAM-KS5781, locality 20—7/1, Mfolozi River, RV, Maastrichtian II.
Scale bars: E = 30y, others = 100w.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 119

directed projection. There is a small rounded isolated median process post-
adjacent to the SCT. Dorsolateral rib is weakly developed, is not connected to
the eye spot, and ends in at an angular posterodorsal node. Valve surface is
coarsely punctate with smaller fossae in the anterior area. Fossae are generally
rounded and well separated, but immediately adjacent to the marginal ribs they
are larger and quadrate.

Internal features. MA moderately wide, but no good views of MPC avail-
able. Hinge is apparently holamphidont. MS consist of a hooked anterior scar
and four adductors. The second scar is the largest and is a ‘dog’s bone’ shape,
whilst the first scar has a dorsal extension.

Remarks

P. monziensis differs from the genotype P. subparva (Pokorny 1967) (as
redescribed by Grtindel 1973), by lacking a distinctly concave VM and in
possessing a recurved ventral ridge. It is very close to P. spinosa from the
Maastrichtian III of the Agulhas Bank, but differs in having a less pronounced
triangular valve outline (particularly in the RV) and in having a rounded SCT,
rather than the ovate feature of P. spinosa. Their close relationship suggests that
P. spinosa evolved from P. monziensis during the middle—upper Maastrichtian.

P. monziensis also bears a resemblance to a lower Eocene form referred to
Hazelina sp. 4 from core number 9 at DSDP site 246 off east Africa (Ducasse &
Grekoff 1976), and to Curfsina turonica Bate, 1969 (in Bate & Bayliss 1969)
from the Turonian of Tanzania. It is possible that both the latter species should
be assigned to the genus Parvacythereis.

P. monziensis has been selected as a subzonal species for part of the
Dutoitella dutoiti Zone proposed herein.

Dimensions (mm)

length height
K5781 0,43 eae
K5782 0,45 0,21
K5783 0,47 O25
K5784 0,46 023
Other material 0,44 0,24
Other material 0.42 O22
Other material 0,44 O22

Age, distribution, palaeoecology

Campanian V to Maastrichtian II (Monzi and Mfolozi areas, Zululand). P.
monziensis has been recorded from assemblages 5b, 6, and 7 (Fig. 55E, Table
4), but its patchy distribution in the first two indicates that although it tolerated
conditions that ranged shoreward to mid shelf (?300-500 m), it preferred deep
water (> 500 m outer shelf/upper slope) with oceanographically stable environ-
ments.

120 ANNALS OF THE SOUTH AFRICAN MUSEUM

Parvacythereis spinosa (Dingle, 1971)
Figs 59A-C, 60B

Phacorhabdotus spinosa Dingle, 1971: 408-410, fig. 11.

Remarks

As pointed out by Dingle (1971), although the lateral outline is typical of
the genus Phacorhabdotus, to which it was originally assigned, its ornamentation
differs from other species within the genus. Its translation to Parvacythereis
removes this anomaly.

P. spinosa is very close to P. monziensis sp. nov., but the two can be
differentiated in the distinctly more triangular lateral outline of the former
(especially in RV), on the shape of the SCT which is rounded in P. monziensis
and anteriorly elongated in P. spinosa, and on the somewhat more prominent
dorsal ridge of P. spinosa. Although not clearly visible, under high magnification
the MS of P. spinosa are very similar to those of P. monziensis with the second
adductor being the largest (Fig. 60A—B).

Age, distribution, palaeoecology

Maastrichtian III (sample 818 Agulhas Bank). The ostracod assemblage of
sample 818 is thought to represent an unusual setting: a quiet, shallow-water
environment on the outer edge of the continental shelf or on the uppermost
continental slope.

Genus Phacorhabdotus Howe & Laurencich, 1958
Phacorhabdotus? anomala Dingle, 1971
Fig. 59D-E

Phacorhabdotus? anomala Dingle, 1971: 410-411, fig. 12.

Remarks

An SEM view of the holotype (Agulhas Bank) is included herein to
supplement the original description. Although the specimen from JC—1 borehole
is crushed, the essential morphological elements are visible and allow a confident
assignment. Bate (in Bate & Bayliss 1969, pl. 8 figs 8-9, 12) has recorded a very
similar species from the Maastrichtian of Tanzania: Phacorhabdotus sp.

Age, distribution, palaeoecology

Maastrichtian III (sample 818 Agulhas Bank), Maastrichtian (level 1 625 m)
JC-1 borehole. The ostracod assemblage of sample 818 is thought to represent
an unusual setting: a quiet, shallow-water environment on the outer edge of the
continental shelf or on the uppermost continental slope, whilst level 1 625 m in
the upper part of the JC-1 Maastrichtian was probably deposited during a period
of relatively deep-water (100-200 m) conditions on the Tugela delta top.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 12}

Fig. 59. A-C. Parvacythereis spinosa (Dingle, 1971), TBD 818 Alphard Formation, Agulhas
Bank, Maastrichtian III. A. SAM-K5785, LV. B. SAM-K5786, RV. C. SAM-K5787,
internal LV. D-F. Phacorhabdotus? D. P? anomala Dingle, 1971, holotype, MG—4-1-18,
TBD 818 Alphard Formation, Agulhas Bank, RV, Maastrichtian III. E. P? anomala Dingle,
1971, SAM-K5788, JC-1 1625 m, LV, Maastrichtian. F. P? sp. A, SAM-K5789, JC-1 1756 m,
LV, Maastrichtian.
Scale bars all 100u.

122 ANNALS OF THE SOUTH AFRICAN MUSEUM

<i oe
IZ oF

monziensis spinosa

B

Fig. 60. Muscle scars of Parvacythereis. A. P. monziensis sp. nov., SAM-—K5784, locality 20-3,
Mfolozi River, RV, Maastrichtian I. B. P. spinosa (Dingle, 1971), TBD 818 Alphard
Formation, Agulhas Bank, RV, Maastrichtian III.

Scale bars 30y.

Phacorhabdotus? sp. A
Fig, 59F

Remarks

One valve with the characteristic external morphology of the genus was
recovered from the JC-1 borehole. No internal views were available so generic
assignment is provisional. Preservation is not good, but well-developed intercos-
tal reticulation is present, as well as occasional conjunctive perforate pustules.

Age, distribution, palaeoecology

Maastrichtian (level 1 756m) JC-1 borehole. Phacorhabdotus? sp. A
occurs, together with Bythocypris? sp. and Cytherella sp., at levels that are
thought to have been intermediate between the deeper (?200 m) and ?shallower
(with large fresh-water influxes) water environments (Table 10) on the Tugela
delta top.

Genus Curfsina Deroo, 1966

Within the Indian Ocean area this genus has previously been reported from
the Turonian of Tanzania (Bate & Bayliss 1969) and the Campanian of Western
Australia (Bate 1972). In south-east Africa it has been found at one locality
only, where it constitutes 18 per cent of the cytheracean population (4% total
ostracods).

Curfsina monziensis sp. nov.
Fig. 61A—D
Derivation of name

Locality of type.

Holotype
SAM-K5790, LV, locality 21-1, Monzi, Campanian V

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 123

Paratypes

SAM-K5791, RV, locality 21-1, Monzi, Campanian V
SAM-K5792, RV, locality 21-1, Monzi, Campanian V

Fig. 61. — monziensis sp. nov., ee ea 21-1, Monzi, Campanian V. A. Holotype,
SAM-KS5790, LV. B. SAM-K5791, RV. C. SAM-K5792, MS RV. _D. SAM-K5792, internal
RV.

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

Diagnosis
Smooth species with a stepped, peg-like ATE in RV hinge.

Description

External features. In lateral view, rectangular with rounded AM, slightly
acuminate PM, and straight, slightly converging DM and VM. There is a broad,
rounded AM rim, which is continuous with wide, rounded dorsal and ventral
ribs that both end in posterior swellings. A median elevation, commencing over
a SC swelling, projects and narrows posteriorly, where it curves towards the
posterior end of the dorsal rib. These two elevations are separated by a narrow
depression. Valve surfaces are completely smooth except for small punctae
(about Su across). There is a large, but weak, eye spot in both valves: in RV it
lies on the anterior cardinal angle, in LV below a smooth anterior cardinal
swelling.

124 ANNALS OF THE SOUTH AFRICAN MUSEUM

Internal features. Hinge holamphidont, but PTE is weakly subdivided and
RV ATE is a stepped smooth peg. MS consist of a kidney-shaped anterior scar,
with a rounded scar above it, and a vertical row of 4 adductors: the top two are
oblong, the lower two rounded. MA generally narrow, no vestibule. MPC fine
and straight, up to twenty anteriorly.

Remarks

C. monziensis is close to C. levigata Bate from the Campanian of Western
Australia, but differs in not possessing a subdivided ATE, in having its median
lateral rib separated from its dorsal rib, and in having continuous ventral and
anterior ribs. The two species also have a slightly different MS pattern.

In his generic diagnosis, Deroo (1966: 139) states that the anterior MS is
capped by a small oval scar. In C. monziensis this small scar is distinctly separate
from the reniform anterior scar.

Dimensions (mm)

length height
K5790 0,62 0,36
K5791 0,59 G31
KS792 0,60 0,33

Age, distribution, palaeoecology

Campanian V (locality 21, Monzi, Zululand). C. monziensis has been found
only at locality 21, in a population that belongs to assemblage 6: deep water
(?>500 m, upper continental slope) with unstable oceanographic conditions.

Genus Cativella Coryell & Fields, 1937
Cativella? dubia sp. nov.
Fig. 62A-B
Derivation of name

Latin dubius (uncertain): reference to uncertain taxonomic status.

Holotype
SAM-K5793, LV, locality 20—7/2, Mfolozi River, Maastrichtian II

Paratype
SAM-K5794, RV, locality 20-1/2, Mfolozi River, Maastrichtian I

Diagnosis
Species in lateral view having curved dorsal and ventral ridges that termi-
nate in median deflections, intercostal areas are reticulate.

Description

External features. A small species, with, in lateral view, AM broadly
rounded, PM strongly acuminate, spinose ventrally. VM almost straight, DM

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 125

bo * =
ae M, $ i
i Sed
x ES ae

oP ONS
TL gee
Pa a oe ee 3 eh oe

ce

<
eg af

t-3

Fig. 62. Cativella? dubia sp. nov. A. Holotype, SAM-K5793, locality 20-7/2, Mfolozi River,
LV, Maastrichtian II. B. SAM-K5794, locality 20-1/2, Mfolozi River, RV, Maastrichtian I.
Scale bars 100.

hidden by lateral ridge, but straight, strongly sloping posteriorly. Highest point
of valve over anterior cardinal angle, greatest length below median line. Surface
Ornamentation dominated by ribs. There is a narrow AM rib that runs from the
large eye spot, and is continuous with a VM rib that continues to the posterior
end of the valve. DM is hidden behind a rib that curves medianly at both ends.
A ventrolateral rib, curved medianly at both ends, runs parallel to a median rib
which runs slightly diagonally across the central part of the valve. This median
rib is hooked at its posterior end, and anterior of a point over the MS area is
accompanied by a further, short, parallel rib. Intercostal areas are reticulate,
with quadrate fossae and disjunctive and conjunctive pustules. Reticulation is
coarser in the compressed anterior area, with small perforate pustules within
fossae. LV and RV differ slightly in shape, there being a curved dorsal valve
extension over the eye spot in LV.

Internal features. No internal features seen.

0,4 Rayneria e

= r Costa
= #

= Carinocythereis

0,3 x
®Cativella

0,6 length 0,8

Fig. 63. Length v. height scattergram of holotypes of Cativella, Rayneria, Carinocythereis and
Costa (dots) compared to specimens of Cativella? dubia (crosses) from Maastrichtian of
Zululand.

Remarks

Generic assignment is uncertain because internal views are not available,
and because the external features cannot be fitted unreservedly into any de-
scribed genus. Four genera have various features in common with the new

126 ANNALS OF THE SOUTH AFRICAN MUSEUM

species: Costa, Rayneria, Carinocythereis and Cativella. The first three, whilst
possessing similar rib patterns, lack its posterior outline and are considerably
larger (Fig. 63). Cativella is closest, but the genotype differs in possessing a
slightly different rib pattern, although other species (e.g. Cativella semitrans-
lucens (Crouch) quoted by Van Morkhoven 1963) are more similar. Cativella is
considered a typical Tertiary taxon and most species that have been unequivo-
cally allocated to it have been recorded from Central and South America. The
only South Atlantic record is of C. moriahensis van den Bold, by Neufville
(1979) from the Lower Eocene of eastern Brazil.

Dimensions (mm)

length height
K5793 0553 0,30
K5794 0,50 0,26

Age, distribution, palaeoecology

Maastrichtian I to Maastrichtian I] (Mfolozi River, Zululand). C? dubia is
environmentally bound to conditions represented by ostracod assemblage 7:
deep water (?>500 m outer shelf/upper continental slope), oceanographically
stable.

Genus Paraplatycosta Dingle, 1971
Paraplatycosta reticulata Dingle, 1971
Fig. 64A-E
Paraplatycosta reticulata Dingle, 1971: 416, fig. 16, pl. 8C.

Remarks

No further specimens of this species have been recovered from the Upper
Cretaceous of south-east Africa. SEM photographs of topotypes are included to
supplement the original descriptions.

Age, distribution, palaeoecology

Maastrichtian III (sample 818 Agulhas Bank). The ostracod assemblage of
sample 818 is thought to represent an unusual setting: a quiet, shallow-water
environment in the outer edge of the continental shelf or on the uppermost
slope.

Indeterminate taxa

Indet. sp. 1
Fig. 655A

Remarks

Single battered valve of reticulate, trachylerid-like species. May belong to
Oertliella.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 127

Fig. 64. Paraplatycosta reticulata Dingle, 1971, TBD 818 Alphard Formation, Agulhas Bank,
Maastrichtian III. A. SAM-K5795, RV. B. SAM-K5796, internal RV. C. SAM-—KS5797,
dorsal view RV, anterior to right. D. SAM-K5797, dorsal view PTE RV. E. SAM-KS5796,
MS RV.
Scale bars: D = 30u, E = 10y, others = 100u.

128 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 65. Indeterminate species. A. Indet. sp. 1, SAM—K5798, locality 20-7/1, Mfolozi River,
RV, Maastrichtian II. B. Indet. sp. 2, SAM-K5799, locality 20-1/2, Mfolozi River, RV,
Maastrichtian I. C. Indet. sp. 3, SAM-—K5800, locality 20-7/3, Mfolozi River, LV, Maastrich-
tian II. D. Indet. sp. 4, SAM-—K5801, locality 20-7/3, Mfolozi River, RV, Maastrichtian II. E.
Indet. sp. 5, SAM-K5802, Igoda Formation, Igoda estuary, LV, late Campanian/early Maas-
trichtian. F. Indet. sp. 6, SAM-—K5803, Lower Quarry Needs Camp, LV, late Campanian/early
Maastrichtian.
Scale bars all 100u.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 129

Age, distribution, palaeoecology

Maastrichtian II (Mfolozi River, Zululand). Ostracod assemblage 7.

Indét. sp..2
Fig. 65B

Remarks

Carapace, small, wholly reticulate species with a strong ventrolateral over-
hang. No eye spot or costation.

Age, distribution, palaeoecology
Maastrichtian I (Mfolozi River, Zululand). Ostracod assemblage 7.

Indet. sp. 3
Fig 65C

Remarks

Single valve of Eucytherura-like species with prominent eye spot. There is a
small posterodorsal process, and a more prominent posteroventral process at the
end of a blade-like and anteriorly-upturned ventrolateral ridge. Surface reticu-
late overall. No true caudal process.

Age, distribution, palaeoecology
Maastrichtian II (Mfolozi River, Zululand). Ostracod assemblage 5b.

Indet. sp. 4
Fig. 65D

Remarks

Two valves of a small, delicately ornamented species with fine, inclined
longitudinal ribs in the central area of lateral view, and marginal, parallel ribs in
anterior and ventral areas. Intercostal areas with scattered, small rounded
fossae. There is a subdued SCT and a small eye spot. Grosdidier (1979) has
illustrated a superficially similar specimen which he called Dumontina? GAD12
from the Turonian of Gabon. Our species is not a Dumontina, nor is it
Triginglymus which is a further superficially similar specimen from lower Eocene
from DSDP site 246 (Ducasse & Grekoff 1976).

Age, distribution, palaeoecology

Maastrichtian I to Maastrichtian II (Mfolozi River, Zululand). Ostracod
assemblages 7 and Sa.

130 ANNALS OF THE SOUTH AFRICAN MUSEUM

Indet.isp,5
Fig. 65E
Remarks

Carapace of elongate-ovate, reticulate species with prominent ovate eye
spot. Reminiscent of the more elongate varieties of Haughtonileberis fissilis from
Umzamba, but poor preservation precludes detailed comparisons.

Age, distribution, palaeoecology
Late Campanian/early Maastrichtian [goda Formation.

Indet. sp. 6
Fig. 65F
Remarks

Poorly preserved carapace of a Hermanites-like species. The only evidence
of original ornamentation is coarse reticulation in the anteroventral area.

Age, distribution, palaeoecology
Late Campanian/early Maastrichtian, Lower Quarry Needs Camp.

Indet. sp. 7
Fig. 66A-B
Remarks
Carapace of an inflated spinose and pustulate blind species. The author

suspects that it may be related to the genus quoted as Acanthocythereis? by
Benson (1977).

Age, distribution, palaeoecology

Maastrichtian, borehole JC—-1 (level 1 676m). The only record of this
species is from within the mid Maastrichtian section with abundant charophytes

Fig. 66. Indeterminate species 7, SAM-—K5804, JC-1 1676 m, Maastrichtian. A. Lateral view
LV. B. Detail anterior area LV.
Scale bars 100y.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 131

(same horizon as Australileberis stangerensis), where large influxes of fresh-water
debris on to the Tugela delta top are suspected during a local low sea-level
stand.

DISCUSSION

All the ostracods encountered were benthic types, and a total of 78 species
belonging to 46 genera were identified in the Campanian—Maastrichtian strata of
south-east Africa. Their spatial and temporal distributions are shown in Tables 1
and 2. It is convenient to commence a survey of the ostracod faunas by
discussing the populations on a geographical basis before attempting a regional
stratigraphic synthesis. In this connection, the composite succession from the
various outcrops and the BH-9 borehole in Zululand is by far the most
complete, and provides to a first approximation an unbroken section through the
Campanian—Maastrichtian. The ostracod faunas from this area will, therefore,
be used as a standard for comparison, and will allow correlation with the
ammonite stratigraphy established by Kennedy & Klinger (1975 et seq.).

In his study of the Santonian—Campanian ostracods of the Richards Bay
BH-9 borehole, Dingle (1980) used two principal techniques to discriminate
significant faunal associations. The first highlighted the specific composition of
the ostracod populations, with particular emphasis on dominant (>20%) and
rare or marker types, coupled with population trends such as variations in
species turnover and population similarity. The second technique demarcated
fields of population similarity on a Cytheracea/Cytherellidae/Bairdiacea + Cy-
pridacea triangular diagram (CCBC plot). Five different ostracod assemblages
were recognized, which with further consideration of the local geology and
faunal characters, could be correlated with particular environments of deposition
(Dingle 1980).

In the present paper these two techniques are again applied and further
modified and extended, with the CCBC plot in particular appearing to hold
considerable potential especially when used in conjunction with the distribu-
tion of selected species or groupings. Because the oldest sections in the
Zululand outcrops are time equivalents of the top part of the BH-9 borehole
(Campanian II), it has been possible to extend the well-controlled faunal and
ecological classifications of the borehole study to the various outcrops. This gives
an excellent ‘datum’ for palaeoenvironmental predictions in the late Campa-
nian—Maastrichtian rocks in areas of Zululand to the north of the BH-9
borehole.

The ostracod populations from the more isolated outcrops and boreholes in
Figure 1 can be compared with the relatively well-known Zululand assemblages,
and predictions on their palaeosedimentary environments can then be made.
Once the ecological factors have been assessed, some regional and extra
southern African biostratigraphic comparisons are attempted.

132 ANNALS OF THE SOUTH AFRICAN MUSEUM

PALAEOECOLOGY

NORTHERN AREA
BH-9 borehole and Zululand outcrops

Figure 2 shows the sampling localities in this area, and Figure 3 shows the
stratigraphic relationships of the various outcrops using Kennedy & Klinger’s
(1975) ammonite subdivision of the three uppermost stages of the Cretaceous.
From these it can be seen that there is likely to be only a slight overlap or
possibly a small gap in the succession between the BH-9 borehole and the
composite section from outcrop in the Campanian II. This gives a complete
section from lower Santonian II to upper Maastrichtian II, including the Santo-
nian—Campanian boundary which occurs between samples at 115 m and 110 m in
the BH-9 borehole (Dingle 1980).

In his study of the BH-9 borehole, Dingle (1980) described the Campanian
I and II ostracod faunas and allotted them to two ostracod assemblages (45)
with assemblage 4 appearing 1,5 m beneath the Santonian—Campanian bound-
ary. Applying the same classification, a further two assemblages can be recog-
nized in the Campanian III to Maastrichtian II strata of Zululand, whilst
assemblages 4 and 5 can be subdivided.

Ostracod assemblages 4 to 7

54 species assignable to 35 genera have been recorded from the Cam-
panian—Maastrichtian rocks of this area. Their vertical distributions are shown in
Table 2, whilst various statistical data on species and population trends through
the succession are shown in Figure 67. Two ostracod types numerically dominate
the faunas (>20% total population), and in combinations with five other
secondary species (each constituting 10-20% total population) and variations in
other parameters of the ostracod population (such as similarity, proportion of
higher taxa, distribution of minor and rare taxa) allow six distinct groupings
within the faunas to be recognized. These assemblages have been annotated 4 to
7, with 4 and 5, as originally defined by Dingle (1980) each subdivided into a and
b. Assemblages, with the dominant ostracod types (with maximum %, and mean
%, based on 3-point means in parenthesis), are:

Assemblage 4a Bairdoppilata andersoni (36%, 27%)

Assemblage 4b Bairdoppilata andersoni (36%, 32%)

Assemblage 5a Bairdoppilata andersoni (37%, 32%) with Cytherella sp.

(31%, 28%)
Assemblage 5b Bairdoppilata andersoni (41%, 32%) with Cytherella sp.
(32%, 30%)

Assemblage 6 Cytherella sp. (47%, 40%)

Assemblage 7 Bairdoppilata andersoni (34%, 32%)

Seven of the rare species are restricted to one or other of these assemblages
(see Fig. 67 and Tables 6-7), and certain secondary types, notably Bythocypris
richardsbayensis, have distributions that allow use to be made of them as
assemblage markers. A summary of these associations is given in Table 6.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA (eG:

Maastrichtian Campanian oka
a ar a

au ae

assemblage

similarity a

population

trends
N° species oa
diversity %
60 Bairdiacea f
os ee higher
So 40 on oe = i BE a se
q-. ane art sa = ~ taxa
20 | Sea ate lace y
B.richardsbayensis
40- B.andersoni ele ie ate ; Brachycythere spp. dominant
% af Np U.sacsi oo | Haaghtonileberis species
20-4. -——eN , iia 28 te S ; ME ee spp.
sd rs — ect r] a et a as
| "NY SY a wud Tre ccunnut iO
borehole &
20 ha es locality
8 7-37-2,7-15 3 1-31-21-11 1311 3 1 0 211914 2 110 bed number
R.nealei
i
ie Sciqnesbachi | griesbachi
| | ~~ H.vanhoepeni_
| | | Ccumzambaensis
T. minima __
§ A.zululandensis
C.contorta
a ae O. africana
St a a a
___ SS ae 0 ee ee X.luciaensis
= H.nibelaensis
U.sacsi other
ee Ss | (D A-avovatus
O.maastrichtia
ae) C-mifoloziensis
Peo SS sf |. Pnibelaensis
eee et Kh. nibelaensis
eee Pateicana
[| iD. dutoiti_|

C.monziensis

E? pyramidatus

P.monziensis

P.lanceolata

[Sipe

C? dubia

P. fragilis

K.aranearius

P.biremis

P.hirsuta

H? arcus

Lsp4

_C. cf westaustraliense

C.brenneri
ealspa

l.sp3

Fig. 67. Variations in the Campanian—Maastrichtian ostracod faunas of Zululand (Richards
Bay BH-9 borehole, and outcrops at Nibela Peninsula (locality 110 & 113), Monzi (locality 21),
and Mfolozi River (locality 20)). Discontinuities in the various trends have been used to delimit
ostracod assemblages 4a, b-7, which are described in detail in the text and summarized in
Tables 6 and 9. Population turnover is a measure of the number of new appearances and
extinctions in each sample, and population diversity is calculated as number of species per
hundred specimens and expressed as a percentage. Population similarity is calculated between
adjacent samples in the sequence as number of species common to the two samples/total
number of species in both samples x 100. Fluctuations in population similarity will give an
assessment of population stability—a sequence of highly similar samples will denote a stable
population, and vice versa. Note that in all cases, percentage of species and higher taxa quoted
in this figure and in the text are based on three-point running mean values.

ANNALS OF THE SOUTH AFRICAN MUSEUM

134

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135

CAMPANIAN AND MAASTRICHTIAN OSTRACODA

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136 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 7

Species with restricted distribution in the Campanian-Maastrichtian ostracod assemblages
of Zululand (BH-9 borehole, Monzi, Mfolozi and Nibela outcrops).

common
assemblage 4a 4b Sa Sb 6 Ws to

Trachyleberis minima : ; rf one
Amphicytherura suldandens’. : “
Apateloschizocythere mclachlani _ . #
Unicapella reticulata

Curfsina monziensis 4 , , *
Cativella ? dubia 2 ‘ : : %
Pariceratina hirsuta

Hermanites? arcus . : : 5 *

Cytherelloidea contorta . : % wk two
Haughtonileberis vanhoepeni . ; :
Prerygocythere lanceolata : ; + %

Cytherelloidea griesbachi ‘ - Ee x three
Haughtonileberis nibelaensis . : * Ee a
Amphicytherura armatus : : # Ee
Klingerella aranearius . : : # *

Cytheropteron brenneri_ . ; : = 2
Ponticulocythere biremis : ' Be nk

Pedicythere fragilis . : : ; Ba si
Parvacythereis monziensis : : x *

Ye ee eK KR KH Y

Dutoitella dutoiti . , ; ‘ Bs sf *
Platella africana : : ‘ : # si x

four

Pe aks

Expressing the make-up of the ostracod populations in terms of a CCBC
diagram allows more refined use to be made of the presence and/or absence of
certain secondary taxa, as well as bringing out clearly the association of certain
minor taxa with particular assemblages (Fig. 69). This is particularly important
in the present study where, because of the generally low abundance of the
cytheracean component, all the assemblages are dominated by either Bairdoppi-
lata andersoni or Cytherella sp., a situation that contrasts strongly with the
Santonian part of the BH—9 borehole (assemblages 1-3) where the Cytheracea
provided the dominant assemblage markers (Fig. 68).

Ostracod assemblages 4 to 7, as defined in Figure 67 and Table 6, plotted o on
a CCBC diagram (Fig. 68) cluster into well-defined fields in a similar fashion to
assemblages 1 to 5 from the Santonian II to Campanian II of the BH—9 borehole
(Dingle 1980, fig. 33). Because of the alternating nature of the ostracod
assemblages throughout the Campanian II to Maastrichtian II at outcrop (Fig.
67), the CCBC diagram (Fig. 68), in fact, gives a clearer picture of the
interrelationships of the various faunas than does a conventional distribution
chart (Fig. 67). In the BH—9 borehole, the lowermost Campanian I assemblage
(4) was seen to pass rapidly, in the lower part of Campanian II, into assemblage
5, whose field extended progressively towards the CBC baseline. In the present
study, populations from the lowermost sections at outcrop (Campanian II)

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 137

CYTHERACEA

( AGULHAS BANK

is

“\~=—Cytheracea

Se ee 7. tine
Ab estate ;
Sa" Ay i : 5a
Ree \
Byinocy pis” ee ey,
line Pe a 4 5
Wide poe ps Has
i / | ae.

/Bairdoppilata line A Scytherelta line
XNEEDS CAMP Sb

BAIRDIACEA & CYTHERELLIDAE
CYPRIDACEA

.Fig. 68. Triangular (CCBC) plot of Campanian—Maastrichtian ostracod populations from
Zululand (Richards Bay BH-9 borehole and outcrops at Nibela Peninsula, Monzi, and Mfolozi
River). Individual samples (crosses) from the Agulhas Bank, Igoda, and Needs Camp are also
plotted. Assemblage fields 4a, b-7 are delimited by major discontinuities (‘Bythocypris’,
‘Bairdoppilata’, and ‘Cytherella’ lines) which are discussed in the text. The ‘Cytheracea’ line
demarcates areas in which cytheraceans dominate (above) and do not dominate (below).
Assemblage fields 1-3 identified in Santonian strata in the BH—9 borehole (Dingle 1980) are

shaded.

cluster below the BH—9 assemblage 4 field, yet have very close affinities with it.
Similarly, several faunas cluster within the BH-9 assemblage 5 field, yet lie
between two other assemblage fields (6-7) that were not encountered in the
borehole. Clearly, a redefinition of assemblages 4 and 5 is necessary, but in such
a way as to emphasize their internal similarities. Useful in this regard has been
the recognition of three important boundaries shown in Figure 68: the 10 per
cent (total ostracod) contour of Bythocypris richardsbayensis (the ‘Bythocypris
line’); a line to the left of which Bairdoppilata is the dominant ostracod (the
‘Bairdoppilata line’); and a line to the right of which Cytherella sp. is the
dominant ostracod (the ‘Cytherella line’). Reference to Figures 68 and 69 shows
that the assemblage fields can now be defined in terms of these lines, in addition
to other parameters on Figure 67, and Table 6.

A summary of the more important characters of the various ostracod
assemblages graphically shown in Figures 67-69 and listed in Table 6 now
follows:

138 ANNALS OF THE SOUTH AFRICAN MUSEUM

Assemblage 4: Bairdoppilata/Brachycythere (4a), Bairdoppilata/Unicapella
(4b). The whole field of assemblage 4 lies above the ‘Bythocypris line’ and to the
LHS of the ‘Bairdoppilata line’, with individual cytheracean types making up
>10% of the total population. It consists of two clearly defined populations:
those of the Campanian I strata of the BH—9 borehole (4a), and those of the
Campanian II, III, and IV rocks at outcrop (4b). These can be separated by
reference to two parameters:

(i) the relationship of the Cytheracea and Bairdiacea percentages to the 40
per cent line in Figure 67: in 4a these curves lie above and below the 40
per cent line, respectively; whereas in 4b both fluctuate approximately
along the line;

(ii) the composition of the secondary (10-20%) ostracods types: assem-
blage 4a (in the BH-9 borehole) has Brachycythere longicaudata and
Haughtonileberis haughtoni, whereas 4b (at outcrop) has Unicapella
sacsi and Haughtonileberis nibelaensis.

As discussed below, we believe that these population differences reflect
subtle contrasts in the environmental preferences of the two populations, and
are not merel; a biostratigraphic phenomenon. It is possible, however, that the
differences in character of the minor and rare species between assemblages 4a
and 4b (Tables 6-7) reflect the age difference between the two. Five species are
restricted to assemblage 4: Trachyleberis minima and Amphicytherura zululand-
ensis to 4a; Apateloschizocythere mclachlani to 4b; and Cytherelloidea contorta
and Haughtonileberis vanhoepeni to 4 (both a and b).

Assemblage 5: Bairdoppilata/Cytherella (Sa), Bairdoppilata/Cytherella/
Bythocypris (5b). The narrow field occupied by assemblage 5 lies between the
‘Bairdoppilata’ and ‘Cytherella’ lines, and as such represents a transition be-
tween Bairdoppilata- and Cytherella-dominated fields to the LHS and RHS,
respectively. The essential unity of this transitional field can be demonstrated by
the distribution of Hermanites kennedyi (>20% total cytheraceans), which lies
precisely within it, whereas subdivision across the ‘Bythocypris line’ (5a above,
5b below) identifies populations which have subtle differences at the tertiary and
minor taxa levels (Figs 69-70, Table 6)—for examples, the distributions of
Xestoleberis luciaensis, Dutoitella dutoiti, disappearance:appearance ratio in
turnover rate, occurrence of blind species, and population similarities.

Only one species is restricted to assemblage 5 (Unicapella reticulata to Sa),
though six of the other rarer and specialized types occur in either 5a or 5b
(Tables 6-7). |

Assemblage 6: Cytherella/Bythocypris. The field of this assemblage lies
beneath the ‘Bythocypris line’ and to the right of the ‘Cytherella line’. As such,
it is the only assemblage in the Campanian—Maastrichtian succession of Zululand
in which the genus Cytherella completely dominates over Bairdoppilata (Fig.
67). Only one species (Curfsina monziensis) is restricted to this assemblage
which shares other rare and minor forms with assemblages 5 and 7 (Table 7),
although their distribution often extends only to the left of field 6 (e.g.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 139

Parvacythereis monziensis, Oertliella maastrichtia, Hermanites kennedyi). The
only cytheracean ostracod of any numerical significance in assemblage 6 is
Cythereis klingeri, which is rather a curious entrant below the ‘Bythocypris line’,
having earlier in the BH-9 borehole been dominant in the shallow-water,
cytheracean-dominated (80-90%) assemblage 3, and only appearing in a tertiary
role (5-10%) elsewhere in the Campanian I of assemblage 4a.

Assemblage 7: Bairdoppilata/Bythocypris. This is the most distinctive ostra-
cod assemblage in the Campanian—Maastrichtian succession of Zululand. On the
CCBC diagram (Fig. 68) it lies below the ‘Bythocypris line’ and to the left of the
‘Bairdoppilata line’, and is characterized by the presence of a relatively large
number of tertiary and minor species (Table 6), most of which reach their
maximum development within it (e.g. Xestoleberis luciaensis, Krithe nibelaensis,
and Oertliella maastrichtia). In addition, there are several rare cytheracean
types, three of which are restricted to this assemblage (Cativella? dubia,
Pariceratina hirsuta, and Hermanites? arcus), whilst six others are found in only
three other assemblages (Tables 6-7).

Sedimentary environments

In his study of the Richards Bay BH-9 borehole, Dingle (1980) concluded
that the various ostracod assemblages reflect specific sedimentary environments
~ of deposition, and for assemblages 4 and 5 predicted the following parameters:
moderate-depth (?100—200 m), low-energy, open water; and deep (?200-500 m,
outer continental shelf), low-energy, open water, respectively. The present study
has confirmed these observations, but additional data on other ostracod assemb-
lages allow further palaeoenvironments to be identified and the definitions of
environments represented by 4 and 5 to be modified.

The shallow-water environments of BH—9 borehole (assemblages 1-3) are
characterized by a cytheracean component of between 60 and 90 per cent, which
drops sharply in the moderate-depth environments of assemblage 4 to between 50
and 60 per cent. A further decrease to less than 30 per cent is evident in crossing the
‘Bythocypris line’ into the fields occupied by assemblages 5b, 6, and 7 on the CCBC
diagram (Figs 67-68). Various authors (e.g. Van Morkhoven 1962; Rosenfeld &
Bein 1978) have remarked that a preponderance of smooth-shelled blind genera
(e.g. Bythocypris, Bairdoppilata, Cytherella, and Krithe) are characteristic of
deep-water environments (mid-outer shelf, upper slope), whilst recent work
prompted by the Deep-Sea Drilling Programme (e.g. Benson 1977) has corrobo-
rated earlier observations (e.g. Brady 1880) that certain of the architecturally
complex, typically blind, trachyleberid forms are characteristic components of
deep-abyssal populations (mid slope, deep-ocean basin).

The establishment of populations with deep-water (mid-outer shelf, upper
slope) aspects accompanies the subordination of the total cytheracean element
below the ‘Bythocypris line’ on the CCBC diagram (Fig. 68). For example,
Krithe nibelaensis, Bairdoppilata andersoni, Cytherella sp., Bythocypris richards-

140 ANNALS OF THE SOUTH AFRICAN MUSEUM

Cc

Pnibelaensis X.luciaensis

B&C 90 = 6C B&C 50 C

B&C 50. 6 C B&C 50 C

e - H? arcus

H.kennedyi

Pek

tia D. dutoiti turnover

Omaastrich

B&C 30 6C B&C 50 C

Fig. 69. Various population parameters and species distributions plotted on Cytheracea/
Cytherellidae/Bairdiacea + Cypridacea triangular diagrams. Insert shows assemblage field
designations used on Figure 68 and in the text. A. Percentage of total ostracod populations:
>5% Xestoleberis luciaensis (diagonal lines); >4% Krithe nibelaensis (horizontal lines); >5%
Pontocyprella nibelaensis (vertical lines). B. >13% total ostracod population Bythocypris
richardsbayensis (shaded). C. Similarity between adjacent samples in vertical sequence (con-
tour interval 10%). D. Population diversity (contour lines at <5%, 8% and 20%).
E. Percentage of total Cytheracea population: >20% Hermanites kennedyi (black); >16%
Oertliella maastrichtia (blank); >10% Dutoitella dutoiti (horizontal lines); >9% Unicapella sacsi
(vertical lines). Two occurrences of Hermanites? arcus shown by dots. F. Areas where
extinctions dominate over appearances in species turnover.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 141

bayensis, Pontocyprella nibelaensis (all considered typically infra-neritic to bath-
yal genera by Van Morkhoven (1963) ) reach their maximum individual develop-
ment in the fields of assemblages 5b, 6 and 7 (Fig. 69A-B). Xestoleberis
luciaensis does the same, although there is no consensus that the genus is a
typical deep-water form. Further evidence for deep-water affinities for these
assemblages (5b, 6-7) is given by the change in numbers of blind cytheracean
species recorded across the ‘Bythocypris line’ (Fig. 70A, Table 8). Above the
line, the number of blind cytheracean species (six) and density of occurrence (an
average of one species per sample) is relatively low, whereas below the line,
nine species are recorded at an average density of three species per sample.
Similarly, architecturally complex cytheraceans such as Oertliella maastrichtia
and Hermanites? arcus cluster below the ‘Bythocypris line’, whilst Hermanites
kennedyi occurs above and below the line, but to the right of the ‘Bairdoppilata
line’ (Fig. 69E).
TABLE 8

Distribution of blind cytheraceans in the Campanian-Maastrichtian of Zululand (BH—-9 bore-
hole, and Monzi, Mfolozi and Nibela outcrops), shown as number of samples containing each
species per ostracod assemblage.

assemblages

4a 4b Sa 5b 6 7
Oertliella africana. : . ste 1 3
Trachyleberis EP adensis : , : : 1 1 2 2 6

Apateloschizocythere mclachlani

el oe

Unicapella sacsi ‘ : b : ; 2 3 4 6
Dutoitella dutoiti : : : ‘ : : 1 2 3 6
Pedicythere fragilis 1 1 2
Cytheropteron brenneri 1 1 1
C. cf. westaustraliense 1
Pariceratina hirsuta 3
Indet. sp.. 2 1
No. of recordings per assemblage . 3 qt Z 12 9 26
No. of species per assemblage 2 4 6 6 3 8
No. samples per assemblage 7 5) 3 4 4 a
No. records per sample per assemblage 0,4 1,4 v.95 3,0 253 3,7

The weight of this evidence indicates that the assemblage fields below the
‘Bythocypris line’ represent moderate to deep-water (mid shelf to upper slope)
sedimentary environments. Of these, assemblage 7 probably represents the
deepest and most ‘specialized’, as reference to Table 7 and Figures 67 and 69
shows that it has the highest number of. restricted species (three), highest
incidence of blind cytheracean types (eight), and is the preferred environment
for Oertliella maastrichtia, Krithe nibelaensis, Xestoleberis luciaensis, Pontocy-
prella nibelaensis, and Bythocypris richardsbayensis. In addition, assemblage 7
shows the highest values (typically >60%) in population stability as expressed in
terms of percentage faunal similarity (Figs 67, 69C, Table 9), suggesting quiet,
physically stable environments, and yet has a very low faunal diversity (<5%)

142 ANNALS OF THE SOUTH AFRICAN MUSEUM

suggesting somewhat ‘hostile’ conditions (e.g. cold water, high pressure). On the
other hand, abyssal populations, such as those described by Benson (1971), are
not evident. This all points to a deep (>500 m) outer continental shelf/upper
slope environment.

TABLE 9

Suggested palaeoenvironments and summary of population characters for ostracod assemblages 4—7
in the Campanian-Maastrichtian of Zululand (BH-9 borehole, and Monzi, Mfolozi and Nibela outcrops).

assemblage 4a 4b Sa Sb 6 7

all environments are low—energy, open—water, normal—marine

palaeoenvironment: moderate depth moderate depth moderate depth deep water deep water deep water
(?inner-mid (?inner-mid (mid-outer (?outer shelf, (?upper slope, (?upper slope
shelf shelf ?200 m) _ shelf, _ 2300-500 m) >500m), >500 m)
2100-200 m) possibly 2200-300 m) unstable stable
quieter or (fluctuating
colder than 4a currents,
temp. etc)

diagnostic ostracoda:

=>200,7- ‘ ; ‘ . B. andersoni B. andersoni B. andersoni B. andersoni Cytherella sp. B. andersoni
Cytherella sp. Cytherella sp. Cytherella sp.
specialized types . ; é H. nibelaensis
T. minima A. laminata U. reticulata none C. monziensis C? dubia
A. zululandensis P. hirsuta
H. arcus

other characteristics:

Cytheracea 2 : . 50-60% 35-45% 30-50% 20-25% ~20% 20-30%
Bairdiacea . : : . 15-40% 30-40 % <40% >40% 30-40 % 40-50%
Cytherellidae ‘ ; . 10-20% 10-25% 25-30% 25-35% 40-50% 20-30%
Bythocypris richardsbayensis —<MNOW/A <110\7, <10\% usually =10%, =13, Sis y,
Faunal diversity . : : <20% (low) >20% (high) ~20% (med.) <20% (low) <{0'4% <5%
(number spp/100 spec.) (very low) (very low)
Population stability . . 30-60% (low) 26-63% (low) 37-58% (low) 45-67% (med.) 47-57% (med.) 50-66 % (high)

(av. similarity index)

Assemblage 6 is more difficult to define. Its lower incidence of blind
cytheracean species (three), architecturally complex forms, and specialized
species (one), as well as its somewhat less stable populations (47-57%), and very
low faunal diversity (<10%) suggests a ‘hostile’, but physically less stable
environment than assemblage 7. This evidence can be interpreted as indicating
that assemblages 6 and 7 represent geographically similar environments (i.e.
both >500 m, outer continental shelf/upper slope), but that assemblage 6
differed in being oceanographically unstable (i.e. subject to more fluctuations in
energy of the sedimentary environment, or changes in water temperatures).

Assemblage 5b is intermediate in the sense that it is a pathway for
population trends between fields 4b and 6, or 7 and 6. However, characteristics
that it shares with 6 and 7, such as the fact that it has the second highest
incidence of blind cytheracean species (Fig. 70, Table 8), place it firmly in the
‘deep-water’ category. No species are restricted to it, and within it the number
of specimens of Bythocypris richardsbayensis shows a distinct low (>10%,
<13%). These features suggest that assemblage 5b represents a somewhat
shallower environment than either 6 or 7, an hypothesis strengthened by the

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 143

wider range of similarity indices (45-63%) and higher faunal diversity (<20%)
than either of the two adjacent fields (Fig. 69C-D, Tables 6, 9). The latter
indicates greater ranges of physical parameters than 6 and 7, but less hostile
conditions. It is suggested that assemblage 5b represents a quiet, outer-shelf
(2300-500 m) environment.

Above the ‘Bythocypris line’ shallower water conditions are represented by
assemblages 4a, 4b, and 5a. Dingle (1980) has discussed the ostracod faunas of
assemblage 4a (his 4) and reasoned for moderate+depth, low-energy, inner-mid
shelf (7100-200 m) environments. The new data do not contradict this assess-
ment, but necessitate the recognition of a further, closely comparable environ-
ment to accommodate assemblage 4b. The two (4a, 4b) have many features in
common, for example: the dominance of Bairdoppilata andersoni; Bythocypris
richardsbayensis at <10%; and the presence of individual cytheracean species in
secondary abundances (10-20%). They differ, however, in the number of blind
species recorded (1,4 species per sample in 4b, and 0,4 species per sample in 4a)
and the presence in 4b of the moderate to deep-water species Unicapella sacsi
and Hermanites kennedyi. It is possible that both these types had not evolved in
Campanian I times, but coupled with the presence of Pontocyprella nibelaensis,
Oertliella maastrichtia, Platella africana, and Amphicytherura armatus in 4b,
several of which only appear in large numbers in the deeper water assemblages,
the evidence suggests that there is a slightly deeper water element in the 4b
faunas compared to 4a. These differences may however, have been controlled by
factors only loosely related to depth differences such as quieter water, lower
temperatures, lower water turbidity etc., but we have no way at the moment of
quantifying them. For the lack of specific data, 4b is tentatively equated with
marginally deeper water conditions (?200 m) than 4a (?100—200 m), whilst both
represent inner-mid shelf environments.

Assemblage 5a was not distinguished by Dingle (1980) as a separate
grouping, but recognition in the present study of the importance of the
‘Bythocypris line’ necessitates subdivision of assemblage 5 as originally defined.
The area above the ‘Bythocypris line’ (5a) differs in several respects from 5b. In
addition to its smaller populations of Bythocypris richardsbayensis and Bairdop-
pilata andersoni, there are significantly lower numbers of Krithe nibelaensis,
Pontocyprella nibelaensis, and Xestoleberis luciaensis, and particularly Dutoitella
dutoiti (which occurs in one sample only). This gives assemblage 5a a distinctly
shallower-water aspect compared to 5b. On the other hand, it has certain
characteristics in common with 5b which indicate that the two were influenced
by similar physical conditions. Notable is the presence of Unicapella sacsi and
Hermanites kennedyi. The latter is an architecturally complex cytheracean whose
20 per cent (total ostracod population) distribution coincides precisely with the
5a and 5b fields. Blind cytheracean types are less important in assemblage 5a
than in Sb (2,3 species per sample compared to 3,0 per sample), although both
have the same species present. Finally, the population stability within assemb-
lage 5a is lower than within 5b (37-58% compared to 45-67%),

144 ANNALS OF THE SOUTH AFRICAN MUSEUM

suggesting somewhat less stable physical conditions. 5a is, therefore, considered
to be a transitional environment between the moderate depths of field 4 and
areas below the ‘Bythocypris’ line with which it is linked via assemblage 5b. A
mid-outer shelf (?200—-300 m) environment is suggested.

The concept of field 5 (a and b) as transitional between 4 and 6 and 7 is
strengthened by plotting the incidence of species turnover in the CCBC diagram
(Fig. 69F). Levels of high turnover, with disappearances predominating, cluster
close to the corridor between the ‘Bairdoppilata’ and ‘Cytherella’ lines (and
also, incidentally, in field 4a). This suggests that certain sedimentary environ-
ments (such as 4a and 5) were not conducive to stimulating phylogenetic
development (appearances), whereas others (notably 4b) might have been more
sO.

€ Cc
ii
50 ——— 50
Sat : Al srl] ra Y Ae C11
un A: an
f ae RY imit-%
Mp:
Bal C
oe
BLIND SPECIES BH-9 AGES ZULULAND OUTCROP AGES
eR ee Se ee
Bac 2 C Bac 50 C Bec 50 e

Fig. 70. Population and stratigraphical data plotted on Cytheracea/Cytherellidae/Bairdiacea +
Cypridacea triangular diagrams. Assemblage field boundaries shown by dotted lines—designa-
tion as in Figures 68 and 69. A. Distribution of blind species of cytheraceans: small figures
show numbers of blind species per sample, areas enclosing 3 or more species/sample are
shaded; large figures show numbers of different species per ostracod assemblage field. B-C.
Ages of samples within ostracod assemblage fields; shading identifies areas enclosing samples of
same age; CI = Campanian I, etc. B. Richards Bay BH-9 borehole. C. Zululand outcrops
(Monzi/Mfolozi/Nibela).

Having discussed the nature of the ostracod assemblages and the
sedimentary environments that they represent, it remains to see how these are
distributed in time and space in Zululand. From Figure 67, it is clear that the
development of assemblages 4a and 5a, 5b in the Richards Bay BH-9 borehole,
and 4b and Sa, 5b at outcrops farther north in Zululand were not synchronous:
the moderate to deep-water environments were established earlier in the south
(Richards Bay area). These relationships can be graphically presented on the
CCBC diagram (Fig. 70B—C), which shows the temporal and spatial distribution
of the various sedimentary environments. For example, the occupation of field
4b during Campanian II times in.the Zululand outcrops (Fig. 70C) was synchro-

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 145

nous with the occupation of field 5 (a and b) farther south at Richards Bay (Fig.
70B). These diagrams also vividly show that for the northerly outcrops there was
a change in the sedimentary environments (i.e. migration across the assemblage
fields) during the Campanian II to Maastrichtian II interval (approximately 9,4
m.y.) from inner-mid shelf through deep water back to mid shelf (4b to 5a via 6
and 7). Similarly, the change from inner-mid shelf to outer shelf (4a to 5b)
during the period Santonian III to Campanian II (approximately 3 m.y.) can be
traced in BH-9 (Fig. 70B).

ostracod
assemblages

cwaleie 8 © 6 ele sa) Oye) sw ies). «

|
q
.
7
|

MAASTRICHTIAN

9°)
<
~
>
°
is)
<
a2)
=
Yn
=
@
b
a
E
z
&

\

\

\

\

\
)
iL

TEE
CAMPANIAN

coenate tee tc elects ste es me © ee ee eee se ee eee

nese e a ow © 6 e o's

eS eee

aT elle Ebi tee
SS |, Sa

a... lil SANTONIAN
G00 SUDO OOOO OOO
BH-9 MONZI/NIBELA
south north

Fig. 71. Temporal distribution of ostracod assemblages in Zululand. Sequence for the northern

area is a composite section based on outcrops between the Monzi area (including Mfolozi

River) and the Nibela Peninsula. The diagram shows that deep-water assemblages were

established earlier in the south, and that the youngest strata in the north indicate a re-

establishment of shallower water faunas. These trends are well illustrated by plotting the
‘Bythocypris’ line.

146 ANNALS OF THE SOUTH AFRICAN MUSEUM

Lateral correlation of lithostratigraphic units (Fig. 71) between the BH-9
borehole and Zululand outcrops shows that the shallow-water (<100 m), inner
and mid-shelf environments (assemblage 4a, 4b) persisted longer in the north,
but that the deepening of the water column that followed (in Campanian IV)
was very rapid. Because we have no data from the Campanian III to Maastrich-
tian II section in BH-9, we do not know if a comparable deepening took place in
the Richards Bay area, nor if environments such as those represented by
ostracod assemblages 6 and 7 were ever established there. The recent review of
oil company borehole data from Zululand by McLachlan & McMillan (1979) is
too generalized to provide the necessary information.

JC-1 borehole

The JC—1 borehole lies on the continental shelf north-east of Durban on the
proximal end of the Tugela delta (cone) (Figs 1; 3, 72). 375 m of Campanian—
Maastrichtian sediments were recorded by Du Toit & Leith (1974, fig. 3), who
described the sedimentary succession as consisting of monotonous light-grey
claystones with thin (<60 cm), hard limestones and subordinate siltstones, and
minor very fine-grained sandstones (their lithofacies 5). Stratigraphic control was
provided by ‘an abundance of planktonic foraminifera’ (Du Toit & Leith 1974:
249), and in their summary McLachlan & McMillan (1979, fig. 3) suggest that
there is a stratigraphic break at the top of the Maastrichtian in the zones
characterized by Globotruncana contusa and G. stuarti. Du Toit & Leith (figs 4,
6) indicate a coarsening of grain-size and an increase in sedimentation rates
(from 18 m/m.y. to 46 m/m.y.) across this hiatus, whereas McLachlan &
McMillan (1979) report a decrease in sedimentation rates (47 m/m.y. to 25
m/m.y.) across the Maastrichtian—Palaeocene boundary.

Ostracod faunas

Prior to the present study, no details had been published of the Cretaceous
ostracod faunas of the JC—1 borehole, although Dingle (1976) has given a
preliminary account of the Tertiary faunas. 28 samples were available for study
from the Campanian—Maastrichtian sequence in the borehole, but only 18
contained ostracods: 3 (43%) in the Campanian, and 15 (71%) in the Maastrich-
tian. The faunas are sparse: a total of 11 species encountered (3 in the
Campanian and 11 in the Maastrichtian); no sample contained more than 10
valves, and half contained 2 or less. This distribution contradicts figure 3 in
McLachlan & McMillan (1979) which shows the Campanian populations as more
abundant than those of the Maastrichtian. |

The ostracods and their distribution are shown in Table 10, and the overall
impression of the combined Campanian—Maastrichtian populations is of the
importance of smooth-shelled, moderately deep-water indicators. In terms of
genera, five of the eight identified fall into this category, but in terms of number
of valves, their dominance is even more impressive (70%): Bairdoppilata 28%,
Bythocypris? 17%, Cytherella 14%, and Krithe 11%. Ornamented cytheracean
forms (five species) account for only 15 per cent.

PALAEO-
CENE

MAASTRICHTIAN

CAMPANIAN

SANT-
ONIAN

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 147

Australileberis stangerensis

=
IS}
Ww
Sond
>
a. 5
oS
Ba &
ae
Sey, SS CS
= = Lees
esos SS
Me OY
x x
x
Ot VEX
x
> ae a <
ew Ox
x x
x
x
x OX
Pate
x
x

Dutoitella mimica

Phacorhabdotus ? anomala

P? sp. &

TABLE 10

Distribution of Ostracods and suggested palaeoenvironments in Campanian-
Maastrichtian section of JC—1 borehole.

Paracypris ? sp.

Indet sp. 7

Indet sp.

charophytes

Inoceramus prisms

NS) No. ostracod valves

NY

NN Ww

NNOLLRO

NN

palaeoenvironments

suggested

1
?shallow-water,
?restricted environment,
essentially barren

2
rapid return to moderately
deep water
(?200 m)

3
shallow, major, fresh-
water influxes

4
moderately deep water
(?200 m) evidence of rapid
shallowing at top

5
depth gradually increasing
upwards to moderate-
water depths (?100—200 m).
Establishment of sparse
ostracod fauna

6
?shallow water

barren of ostracoda

148 ANNALS OF THE SOUTH AFRICAN MUSEUM

Because of the small numbers of specimens encountered, and the very small
populations available at any one horizon, it is not possible to recognize assem-
blages in the same way as in the Zululand area. Nevertheless, some significant
associations can be discerned, and these, in descending order down hole, can be
summarized as:

1 570-1 597 m (uppermost Maastrichtian)—barren, except one carapace of
Bythocypris?

1 625-1 652 m (Maastrichtian)—a Bythocypris?/Krithe fauna with secondary
Cytherella and Bairdoppilata

1 664-1 701 m (Maastrichtian)—a charophytes association with minor ostra-
cod fauna (Bairdoppilata, Australileberis, indet. sp.)

1 719-1 780 m (Maastrichtian)—a mixed Bythocypris?/Krithe/Cytherella/
Bairdoppilata fauna

1 811-1 884m (early Maastrichtian/late Campanian)—a Bairdoppilata/
Cytherella fauna associated with abundant Jnoceramus prisms

1 895-1 934 m (early Campanian)—barren of ostracods, abundant Jnocer-
amus prisms

The rare ornamented cytheracean ostracods occur scattered throughout this
sequence and it is difficult to discern a meaningful distribution. It is suggested
that the following may be significant: the two records of Dutoitella mimica are
confined to the 1811-1 884m association; the relative abundance of
ornamented types in the charophytes-rich horizon 1 664-1 701 m; and the res-
triction of Phacorhabdotus? to associations in which both Bythocypris? and
Krithe are important (1 625-1 652 m and 1 719-1 780 m).

Finally, although ornamented cytheraceans are rare in JC~—1, the three
identified genera have significant geographical distributions. Most interesting is
the presence of two Agulhas Bank Maastrichtian species in JC-1, Phacorhabdo-
tus? anomala, and Dutoitella mimica, which have not been recorded in the
near-by Zululand sequences. Similarly, the genus Australileberis, which is com-
mon in the Eocene of JC—1 and the Agulhas Bank, has been recorded in the
Maastrichtian of JC—1, but not from Zululand. .

Probably this affinity between the Tugela Cone and the Agulhas Bank
(about 1 000 km apart), and the dissimilarity between the ornamented cythera-
cean elements of JC-1 and south Zululand (130 km between JC-1 and Monzi)
reflects contrasting sedimentary environments, although it might have been
influenced by palaeooceanographic factors such as currents (i.e. water tempera-
ture). Although neither Du Toit & Leith (1974) nor McLachlan & McMillan
(1979) give details of the foraminiferal zonation of the Campanian—Maastrich-
tian, the levels at which D. mimica was found (1 911 m Maastrichtian, and
1 871 m Campanian) probably lie stratigraphically below that of sample 818 on
the Agulhas Bank (Maastrichtian III), thereby extending its known range to late
Campanian to Maastrichtian III.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 149

Sedimentary environments

In addition to the ostracod distribution outlined above, any prediction of
the palaeosedimentary environment of the Campanian—Maastrichtian section of
the JC-1 borehole must accommodate the following facts: the ostracod fauna as
a whole is impoverished; throughout the sequence there is an abundance of
authigenic pyrite and agglutinated benthic foraminifera; there is a moderately
rich planktonic foraminifera component; ostracod carapaces dominate over
single valves. Finally, the general geological setting of the area indicates a
location on the top of the Tugela Cone (i.e. a delta top situation) (Du Toit &
Leith 1974; Dingle 1976, 1978).

Generally, the sea-floor conditions were not conducive to the establishment
of a diverse and abundant ostracod community. This was probably because of
the relatively high. terrigenous sediment accumulation rate (McLachlan &
McMillan 1979), which could account for the abundance of agglutinated forami-
nifera and the high carapace:single valve ratio, and mildly anoxic bottom waters
(giving rise to the authigenic pyrite). On the other hand, the area was in
communication, at surface water levels, with the open ocean, thus ensuring a
steady influx of planktonic foraminifera. If the association of Bythocypris?/
Krithe is used as an indicator of moderately deep water (?200 m), and the
abundance of charophytes is taken to indicate a major influx of fresh water
(possibly accompanied by significant shallowing), then the succession of
sedimentary environments, as shown in Table 10, can be postulated. This
suggests that the period under consideration opened and closed with shallow-
water conditions and that there was a further shallow-water episode in ‘mid’
Maastrichtian times. Periods of deepening and moderately deep water inter-
vened in late Campanian to early Maastrichtian, and in late Maastrichtian times.

This suggests that JC—1 was located in a small delta-top depression, open to
the Indian Ocean, but with poor bottom-water circulation. Such an environment
was vulnerable to sea-level fluctuations, being delicately balanced between the
Open Ocean on one side and the influence of a major fluvial sediment source on
the other. Further implications of the sea-level fluctuations implied by Table 10
will be discussed below.

Campanian—Maastrichtian palaeogeography of southern Natal—Zululand Basin

Because of the upward limit of data in BH-9, we have sufficient information
only to construct a Campanian II palaeogeography, although it is unlikely that
the overall geography of the southern part of the Natal—-Zululand Basin would
have altered significantly throughout most of Campanian—Maastrichtian times.

Figure 72 shows a Campanian II reconstruction. To the north of the Tugela
delta, the coast was drowned and rugged, with a mountainous hinterland. The
sites of BH-9 and the Monzi and Nibela outcrops lay in mid-outer shelf
locations, with the shelf edge to the west of the present coast. Although the
Campanian II continental shelf was narrow (up to 25 km), it was probably about
double the present width and was considerably deeper, ranging from 300-500 m

150 ANNALS OF THE SOUTH AFRICAN MUSEUM

2>

rugged hinterland

Tugela Delta

coast
---> Shelf edge
—-- present coast

100 km
——EE eh ee

2 2 Pr oS

"bo mts
SS S>>5>S55> >>> >>

>>> Ss Sas

Se

ae O= ° oe
e : m~

CaaS ee
(ga

Fig. 72. Campanian II palaeogeography of the Tugela—Zululand area of south east Africa.

in the BH-9 area to ?200 m around Nibela (compared with 50 m today). BH-9
lay about 30 km north of the northern edge of the Tugela delta, which extended
as a deep-water cone at least 150 km into the Natal Valley. Immediately south of
BH-9, the continental shelf bulged southwards over the delta top, where the site
of JC—1 lay in a mid-outer shelf location. We do not have a zonation for the
Campanian in JC-1, but from Table 10, Campanian II times probably coincided
with shallow-water conditions which were not conducive to an ostracod fauna.
The only abundant benthos at this time were benthic foraminifera and [nocer-
amus.

Whilst there are insufficient data to reconstruct further palaeogeographies,
it is possible to summarize regional sea-level changes. The most complete data,
from BH-9 and Monzi and Nibela, are plotted on Figure 73 as sedimentary
environment (related to water dépth) versus time. This allows an assessment of

CAMPANIAN AND MAASTRICHTIAN OSTRACODA iS]

rapid minor rapid rapid
regr. regr. /transagr. transgr. transgr.
| | slow transgr.
—
water
depth r.
(00m) .assembl.

6&7

_—- —

Sb

?-- Igoda 4a

?_ —_ Needs Camp

Hl I Vv IV Hl I
MAASTRICHTIAN CAMPANIAN SANTONIAN

65 70 75 80 m.y.B.P.
JC-1 borehole
1600 1700 1800 metres

deep
intermediate
shallow

Fig. 73. Sea-level fluctuations in south-east Africa during Santonian II to Maastrichtian II
times. Suggested water depths relate to predicted sedimentary environments for ostracod
assemblages 1-7 (see Tables 6, 9, and Dingle 1980). Time scale is that of Van Hinte (1976), and
ages of Kennedy & Klinger’s (1975) ammonite stage subdivisions are nominal. Upper part of
diagram relates to Zululand (BH-9 and outcrops between Monzi and Nibela). Single ostracod
populations from Igoda, Needs Camp, and Agulhas Bank have been used to predict water
depths at these localities for specific times. Lower part of diagram shows suggested temporal
distribution of sections 1-6 (shaded) in JC—1 borehole. Table 10 gives details of predicted
sedimentary environments for these sections of the borehole.

the spacing and rapidity of the various sea-level fluctuations that affected the
area in late Cretaceous times. As noted by several authors (e.g. Kennedy &
Klinger 1975; Dingle 1978), the Upper Cretaceous transgression in Zululand is a
southward younging phenomenon and the BH-9 succession documents its
appearance in the Richards Bay area in Santonian II times. Water depth
increase was relatively slow (perhaps 200 m in 3,5 m.y., 57 m/m.y.) through
Santonian II to Campanian I times, with a more rapid rise (~133 m/m.y.) in late
Campanian J/early Campanian II. Data from the area farther north (Monzi and
Nibela) indicate a stand (in this area at a mid shelf, say 200 m depth) for about
3 m.y. during Campanian II to lower Campanian IV times (assemblage 4b),
followed by a major, rapid transgression (involving a water depth increase
probably in excess of 300 m) in early Campanian IV. A further stand occurred
during Campanian IV to lowermost Maastrichtian I (~3 m.y.), with outer
shelf/upper continental slope sedimentary environments (assemblages 6 and 7)
prevailing in the Mfolozi and Nibela area. A short-lived, minor regression is
suspected in Lower Maastrichtian I, which was followed by a minor transgres-

152 ANNALS OF THE SOUTH AFRICAN MUSEUM

sion that restored deep water (assemblage 7) for a further 1,5 m.y. or so. A
major regression started in early Maastrichtian II times, continued evidence for
which is found in our youngest samples ffom south Zululand (mid-upper
Maastrichtian II). The data suggest a sea-level fall of at least 200 m over a
period of 1 m.y. and this may represent the worldwide end-Mesozoic event, that
is locally represented by an uppermost Maastrichtian/early Danian hiatus
(Dingle 1978).

No detailed comparison between the sea-level fluctuation curves for Zulu-
land and JC-1 can be made because there is no refined zonation for the latter.
However, a rough correlation can be attempted by assuming constant sedi-
mentation rates for the Maastrichtian (56 m/m.y.) and Campanian (12 m/m.y.)
sections and plotting the sample levels for JC—1 against the age scale on Figure
73. Several possibly significant points emerge. The basal, barren, ?shallow-water
unit (6) correlates with the late’Santonian/early Campanian slow transgression/
rapid transgression/still stand sequence. The relatively long period represented
by unit 5, which shows evidence of a gradual water deepening and the establish-
ment of a sparse ostracod fauna, correlates with the Campanian IV rapid
transgression and high sea-level stand in the late Campanian/early Maastrich-
tian, whilst the short, deep-water episode (4) correlates with the early Maas-
trichtian high stand. Influxes of charophytes and the general impoverishment of
the ostracod fauna in unit 3, which has been tentatively linked with shallow
water on the delta top, coincides with the initiation of the rapid mid Maastrich-
tian regression, but a further short return to moderately deep water in JC—1
(unit 2) has no equivalent in the Zululand succession because no data are
available over this time sector. On the Agulhas Bank and in JC-1, the late
Maastrichtian was a period of lower sea-levels. Two cautionary notes need to be
sounded about these superficially attractive correlations between the Tugela
delta top and the continental shelf sites farther north. Firstly, the time scales can
be matched only very approximately and, secondly, the two areas were tectonic-
ally different—the Tugela Cone subsidence history was long and relatively
constant, whereas the Zululand shelf was relatively more stable. Non-eustatic
fluctuations may, therefore, have been out of phase and Figure 73 may be
comparing different events. On the other hand, this effect could account for the
apparent lack of evidence in JC—1 for the rapid early Campanian II transgres-
sion, and the smoothing out of the major, rapid Campanian IV transgression.
Taken at face value, the sea-level fluctuation histories of the JC—1 borehole, and
the Zululand sites show a remarkably good correlation.

EASTERN AREA

Two small outcrops of Upper Cretaceous strata occur south-west of East
London. Klinger & Lock (1978) considered the Lower or East Quarry at Needs
Camp to represent a shallower water, restricted, lateral facies equivalent of the
Igoda Formation, both of which they dated as late Campanian/early Maastrich-
tian.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 153

Igoda

With its basal conglomerate and rapidly upward fining sequence into sandy
limestones and calcareous sandstones, the Igoda Formation represents a trans-
gressive series on to the Palaeozoic-Lower Mesozoic basement of the Transkei
Swell. The basal conglomerate contains shell fragments, but all the fossils
identified by Klinger & Lock (1978), and the ostracods described herein, came
from the calcareous member which lies above the brown conglomerate member.
The macrofauna is dominated by ostreid lamellibranchs, whilst other taxa, in
descending order of importance are: brachiopods, baculitid ammonites, echi-
noids, corals, and rare, normally coiled ammonites. The only identifiable macro-
faunal element in common between the Igoda Formation and the Umzamba and
Zululand areas is Saghalinites sp. cf. S. cala (Forbes), whilst Baculites subanceps
Haughton occurs along the west coast of Africa in Angola (Klinger & Lock
1978). The general aspect of the macrofauna suggests moderate to shallow-water
depths.

TABLE 11

Ostracods from Igoda Formation (Igoda estuary), late Campanian/early Maastrichtian

no.specimens % total

Bairdiacea ; . Bairdoppilata andersoni 18 19
B. sp. A 6 6
Bythocypris ee: pete 2 Z 39%
Cypridacea ‘ . Paracypris umzambaensis . 9 12
PP. spcA ; 2
Cytherellidae . . Cytherella sp. 4 4 4%
Cytheracea : . Cythereis Peer 7 %
Hermanites kennedyi . 4 4
HM? cf. arcas 1 1
Pondoina igodaensis 14 ts
Hutsonia ? sp. Z 2 “ee
Buntonia ? sp. : Z 2
Brachycythere longicaudeta 14 15
Xestoleberis luciaensis 2 2
Indet sp. 5 p: 2
Indet spp 6 6
Dominant (>20%) . ; ; . Bairdoppilata spp.
Secondary (10-20%) ; é . Paracypris spp.
Pondoina igodaensis
Brachycythere longicaudata
Tertiary (S-10%) . ; ; . Cpythereis transkeiensis
Cytheracea: total of 54 specimens = 57% fauna
Trachyleberididae MEP! Se 4
Brachycytheridae . : >. 27,
Cytherideidae A é ie HOT
Progonocytheridae : ae,
Xestoleberididae_ . : e's 4
Buntoniidae . ; : A ass ie

Wee wee 2) aS Agee

154 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fifteen species of ostracod belonging to twelve genera have been identified
from the calcareous member of the Igoda Formation (Table 11). Cytheraceans
are the dominant group (57%), and the assemblage ‘plots on the CCBC diagram
close to the BCC line, in the vicinity of assemblage field 4a (Fig. 68). Bairdoppi-
lata spp. is the dominant taxa (29%) followed by Paracypris spp. (12%),
Pondoina igodaensis (15%), and Brachycythere longicaudata (15%). Two impor-
tant points of difference between the Igoda population and those of assemblage
4a from the Lower Campanian of the BH-9 borehole are the low level of
Cytherellidae (4% at Igoda compared to 10-20%) and the low level of the
trachyleberid types (26%). Such low levels of Cytherellidae and Trachyleberidi-
dae were recorded only in the restricted, shallow-water, high-energy of assem-
blage 1 of BH—9 (Dingle 1980). As discussed above, field 4a populations are
thought to represent quiet, normal marine, moderate-depth (?100—200 m, inner-
mid shelf) environments and we consider the Igoda Formation calcareous
member was deposited at the shallow (or higher energy) end of this range, i.e.
say 100-150 m, inner shelf.

The presence of certain taxa within the Igoda Formation call for further
comment. Pondoina igodaensis and Cythereis transkeiensis indicate a close
palaeoenvironmental link with the Santonian—Campanian ostracod populations
at Umzamba: although present in south Zululand (BH-9 borehole), the genus
Pondoina is never common (2-5%), whereas at Igoda and Umzamba it is one of
the more important taxa (15% and 10-15%, respectively); similarly, Cythereis
transkeiensis, which occurs only as a few broken specimens in one sample in the
Santonian of BH-9, is relatively important at Igoda and Umzamba (7% and 5%
respectively). It is possible that the northern limit of the usual geographical
range of Pondoina and Cythereis transkeiensis lay between Umzamba and
Zululand.

Hutsonia? has not been previously recorded from southern Africa, and its
environmental preference is usually considered to be brackish to very shallow
marine conditions. The presence of a single specimen and the uncertainty of its
taxonomic status should caution against too much weight being placed on this
record.

Needs Camp, Lower or East Quarry

The sample available for study consisted of a friable to moderately lithified
creamish coloured calcarenite. Disaggregation could be only partially achieved
and the residues consisted mostly of polyzoa fragments and echinoid spines with
abundant red-stained quartz grains. Glauconite grains and benthic foraminifera
were common and ostracods rare. No planktonic forams were seen, but they
have been reported from the Needs Camp Lower Quarry by McGowran &
Moore (1971).

The ostracod fauna was sparse (54 specimens) with a low diversity (<10%)
(Table 12). The population is dominated by Bairdoppilata (91%), but it is not
possible to say whether this is real or a result of the robustness of the
recrystallized carapaces allowing this genus to withstand any subsequent decal-

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 155

TABLE 12

Ostracods from Needs Camp beds (Lower or East Quarry, Needs Camp),
late Campanian/early Maastrichtian.

a. Present study
no.specimens % total

Bairdiacea : . Bairdoppilata andersoni ; 39
B. andersoni aequalis . : : 10 90 )
94
Cypridacea ; . Pontocyprella sp. ; ; 2 4 f
Cytheracea ef? ef areus : : l 2
indet sp. 6. . : 2 4 :

Polyzoan calcarenite. Residue consists of polyzoan fragments and echinoid spines, benthic
calcareous foraminifera, red quartz and golauconite

b. Ostracoda recorded by Chapman (1916) from Needs Camp Lower Quarry

present assignment S.A. Museum no.
Cythere postcultrata sp. nov. . : Cythere ? postcultrata Chapman 2736/18
(probably invalid)

Bairdia subdeltoidea Minster. . Bairdoppilata andersoni aequalis 2736/20

sp. var. aequalis var. nov. (Chapman)
Bairdia subdeltoidea Minster. . Bairdoppilata andersoni aequalis 2736/17
- (Chapman)
Bairdia africana sp. nov. . : . Bairdoppilata africana (Chapman) 2736/19

cification. Chapman (1916) recorded three species of ostracod from this forma-
tion, two of which were Bairdoppilata, although he did not quote the size of his
population. Our population plots on a CCBC diagram well outside any of the
other assemblage fields encountered in our studies, which itself is to be expected
because the sedimentary environment of this lithofacies (coarse, bioclastic sand)
is unlike any other from which we have examined ostracods in the Campanian—
Maastrichtian of south-east Africa. Its lithology and invertebrate fauna indicate
a very shallow-water (<20 m), normal-marine, moderate to high-energy en-
vironment, with coarse carbonate sand substrate.

Comparing the ostracods from the Lower Needs Camp beds with those
from the Igoda Formation, the two populations contrast strongly, with the latter
having a higher diversity (18% compared to 7%), and a moderately well-
represented cytheracean element (12 spp (57%) compared to 2 spp. (6%)).
They have in common the dominance of the genus Bairdoppilata, and the
presence of two species: Bairdoppilata andersoni, and Hermanites? ct. H? arcus.
The latter is evidence in favour of, but does not corroborate, Klinger & Lock’s
(1978) suggestion that the Lower Needs Camp beds and Igoda Formation are
the same age, although the macro- and microfaunal evidence indicate that they
are both of late Campanian/early Maastrichtian age. Our ostracod evidence does
corroborate Klinger & Lock’s (1978) suggestion that the Igoda Formation was
deposited in deeper water than the Lower Needs Camp beds (see Fig. 73).

156 ANNALS OF THE SOUTH AFRICAN MUSEUM

Umzamba

The Umzamba Formation at its type section consists of alternating sands,
calcareous sands and sandy limestones. Klinger & Kennedy (1977) notated these
beds Pil to Pil9, and on their ammonite faunas assigned them a Santonian II to
Campanian II age (Klinger & Kennedy 1980). These datings have been corrobo-
rated by Makrides (1979) using planktonic foraminifera. In the present study
two samples were collected from the Campanian I section (beds Pi8 to ?Pi14),
but, as found by Makrides in her foraminiferal studies, microfossils in this part
of the sequence are sparse and poorly preserved. Table 13 lists the species

TABLE 13

Campanian ostracod faunas from two samples from the Umzamba Cliff section (locality 4 on
Figs 1, 3). Bed numbers after Kennedy & Klinger (1977).

Sample from bed Pi 13 no. specimens
Brachycythere longicaudata . : : ; d 5
Bairdoppilata cf. andersoni. : ! ‘ ; 2
Amphicytherura tumida : , ; : : 2
Cythereis transkeiensis . : . . fragment

Sample from bed Pi 9
Brachycythere longicaudata . i : 2
Haughtonileberis fissilis 2

extracted, all of which have previously been recorded from the underlying
Santonian III beds by Dingle (1969) and the Santonian III to Campanian I strata
in the BH-9 at Richards Bay (Dingle 1980). The main absentees compared with
the Santonian III at Umzamba are Pondoina sulcata and Haughtonileberis
haughtoni, whilst a significant presence is Amphicytherura tumida, which in
Zululand ranges Santonian III to Campanian I. None of the exclusively Cam-
panian or Campanian—Maastrichtian species that first appear in the Campanian I
of Zululand has been recorded, but the heavy decalcification to which these beds
have obviously been subjected militates against obtaining a representative fauna.
With regard to palaeoenvironment, the populations are not suitable for plotting
on a CCBC diagram. However, the preponderance of Brachycythere longicaudata
and Haughtonileberis fissilis strongly suggests that they belong in assemblage
fields 2 or 3 (<100 m, inner shelf). This in turn suggests that shallow-watér
conditions persisted in the Umzamba area at least until Campanian I times,
when the sea had already deepened to 200 m or more in the Richards Bay area,
but had not yet transgressed into the Needs Camp—Igoda region (see Fig. 73).

SOUTHERN AREA: AGULHAS BANK

The presence of uppermost Cretaceous strata on the Agulhas Bank was first
established by sea-floor sampling in a slump-exposed outcrop on the upper
continental slope (Dingle 1971, 1978). Further dredging and commercial drilling
have subsequently shown that Campanian—Maastrichtian rocks occur extensively
over the eastern Agulhas Bank in the upper part of the Alphard Formation
(Dingle 1973, 1978; Du Toit 1976; SOEKOR 1976; McLachlan & McMillan

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 157

TABLE 14
Ostracods from the Alphard Formation (Agulhas Bank, sample TBD 818), Maastrichtian III.

no.specimens % total

Bairdiacea : . Bairdoppilata andersoni _. : 3 l
Bythocypris richardsbayensis _ . ps i
a
Cypridacea : . Pontocyprella nibelaensis 8 4
Paracypris sp. 2 ]
Gyiereiiidaec . . Wflatellaafricana. . . . l be 10
CYINCFENGSDD we 2 19 of
Cytheracea . , Unicapella sacsi . | l
Dutoitella mimica ' 21 10
Phacorhabdotus ? anomala Il l
Agulhasina quadrata 65 30
Paraplatycosta reticulata 9 4 83
Trachyleberis schizospinosa 73 33
Parvacythereis spinosa 4 3
Krithe nibelaensis ; 2 ]
Apateloschizocythere laminata . 5 2
Dominant (> 20%) , Trachyleberis schizospinosa
Agulhasina quadrata
Secondary (10-20%) , : . Dutoitella mimica

= Tertiary (5-10%) . . Cytherella sp.
Cytheracea: total of 184 specimens = 84% fauna. Trachyleberididae = 96%

1979; Klinger et al. 1980). So far, however, the only description of their ostracod
fauna is that given by Dingle (1971) of sample TBD 818 (Figs 1, 3) which is of
upper Maastrichtian (probably Maastrichtian III) age.

15 species of ostracod, assigned to 15 genera have been identified from
sample 818 (Tables 1, 14), and of these, 9 genera (60% by number of species,
18% by number of specimens) have been recorded from the Campanian—
Maastrichtian of Zululand. The fauna is overwhelmingly cytheracean (83%) with
Trachyleberis schizospinosa (33%) dominant, and Dutoitella mimica (10%) in a
secondary role. On a CCBC diagram (Fig. 68) the population of sample 818 lies
adjacent to the field of assemblage 3, as defined in BH-9, but in several ways the
population make-up of 818 differs significantly from those of assemblage 3. In
addition to specific differences accountable by the age difference, 818 contains a
small number of Bairdiacea (which assemblage 3 does not), a higher percentage
of Cytherellidae (10%) than most of the assemblage 3 popuiations, whilst the
genera Krithe (1%), Unicapella (1%), Dutoitella (10%), and Trachyleberis
(33%), which are all absent from assemblage 3, are typically moderate to
deep-water taxa. Furthermore, five species of the nine cytheraceans in sample
818 are blind forms, and numerically blind forms make up 88 per cent of the
cytheracean population (Table 14). In contrast, only one blind cytheracean type
occurs in assemblage 3 (Rayneria nealei, which is a minor constituent, up
to 7%). The trachyleberid component of the cytheracean fauna

158 ANNALS OF THE SOUTH AFRICAN MUSEUM

in sample 818 is 93 per cent (number of specimens), which is higher than in any
other sample from the Campanian—Maastrichtian rocks of south-east Africa.
Significantly, however, the next highest values recorded (76%, mean of 6
samples) were from the populations of assemblage 3.

60

%/o BAIRDIACEA

TRACHYLEBERIDIDAE

Fig. 74. Fields for ostracod assemblages 1-7 on a percentage Bairdiacea (total fauna) v.
percentage Trachyleberididae (of Cytheracea) plot. Sample 818 from the Agulhas Bank lies
near the Bairdiacea baseline, on the RHS of the diagram. See text for explanation.

Contrasts with typical populations in assemblage 3 makes the assessment of
a palaeosedimentary environment for sample 818 uncertain. Despite the differ-
ences mentioned above, however, the overall composition of the population of
sample 818 places it firmly within the shallow-water area of the CCBC diagram.
Nevertheless, the overwhelming blindness of the fauna and the presence of
elements which are typically indicators of deeper water show that sample 818
was not deposited in a near-shore or even mid-shelf environment, as were the
assemblage 3 populations in the BH-9 borehole. This uniqueness can be further
emphasized by comparing the trachyleberid percentage of the Cytheracea of all
the Zululand Santonian—Maastrichtian samples to some parameter that is inde-
pendent of the cytheracean component, such as the Bairdiacea percentage.
Figure 74 shows a progressive increase in the trachyleberid component of the
Cytheracea with increase in water depths and open oceanic influences up the
BH-9 borehole to a point where moderate water depths replace shallow ones.
Movement of the assemblage fields representative of deeper-water environments
is in the reverse direction, with the bulk of the deep-water assemblages (5b, 6
and 7) lying to the left of the 60 per cent trachyleberid value, and showing a
general decrease in trachyleberid values with an increase in Bairdiacea percen-
tage. Sample 818 lies in the shallow-water areas, but well to the right of other
assemblage 3 populations, indicating what may be termed a ‘modified assem-
blage 3 sedimentary environment’.

Although the evidence is inconclusive, we suggest a quiet, shallow-water
environment (as for assemblage 3), but with dominant open-ocean influences.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 159

Considering the geographical position of sample 818 (in the vicinity of the Upper
Cretaceous shelf break where the shelf is about 135 km wide), this suggests an
outer shelf/upper slope setting in shallow water (?~100 m). Such an unusual
sedimentary environment was presumably created during a regression that may
be correlated’ with the major regression that we have identified as commencing
in Maastrichtian II in our Zululand samples (Fig. 73).

SUMMARY

Within the context of the continental shelf and upper slope environments
encountered in the Santonian—Campanian—Maastrichtian of south-east Africa, it
is possible to summarize the palaeosedimentary environments of the ostracod
faunas studied in terms of a CCBC diagram. The various fields’ (assemblages
1-7, as shown in Figure 68), as well as the data from isolated samples from
Needs Camp, Igoda and the Agulhas Bank are shown in Figure 75. Environ-
ments deeper than 100 m (i.e. mid-shelf and deeper, assemblage fields 4-7) lie in
the central part of the diagram, with the deeper-water areas (assemblages 5b, 6
and 7) lying towards the CBC base line (i.e. below the 30% Cytheracea line).
The anti-clockwise closure of the 100 m line in the high Bairdiacea/Cypridacea
populations is somewhat speculative and is based on the Needs Camp popula-
tions.

Bac

Fig. 75. CCBC triangular diagram used to predict depths (in metres) of palaeosedimentary

environments of ostracod assemblages in the Campanian—Maastrichtian strata of south-east

Africa. Constructed from Figure 68 and Table 6, it includes data from Zululand, Igoda, Needs

Camp, and Agulhas Bank. Within the dotted areas (>100 m depth), the field boundaries are
those shown in Figures 68 and 69 (insert). See text for explanation.

160 ANNALS OF THE SOUTH AFRICAN MUSEUM

Large areas, especially within very low value cytheracean populations, as well as
high Cytherellidae populations are completely unknown, but in conjunction with
Tables 6 and 7 it is hoped that Figure 75 will prove of use in future studies in
discriminating palaeosedimentary environments in pre-Santonian south-east
African sequences where more extensive shallow-water environments are antici-
pated. Clearly, Figure 75 will be ineffective in abyssal populations where an
‘overlay’ for such environments will need to be made.

BIOSTRATIGRAPHY
SOUTH-EAST AFRICA

Because of the relatively limited spatial distribution of individual species,
benthic ostracods are not used in worldwide stratigraphic zonal schemes. They
can, however, be effective locally, and several notable attempts have been made
to extend zonation over moderate-sized regional studies (e.g. Simon & Barten-
stein 1962; Oertli 1963). Data on the temporal distribution of species can be
grouped in various ways to highlight the evolution of local ostracod faunas, and
three of the most useful of these have been effectively employed in a recent
survey of the British Phanerozoic ostracods (Bate & Robinson 1978): phylogeny
emphasizing first appearances of individual species (which illustrates the evolu-
tion of the faunas as a whole), phylogeny emphasizing extinctions of individual
species (the method of plotting ‘tops’ of species ranges), and generic phylogeny.
Only in the Zululand area do we have sufficient data to attempt a similar study,
although the faunas from Agulhas Bank and Transkei Swell outcrops can be
compared individually. Figures 67, 76-80 show data on the Campanian—Maas-
trichtian ostracod faunas in Zululand plotted in the three modes mentioned
above to emphasize their biostratigraphic distribution.

Phylogeny: species distribution (appearances)

Firstly, we shall look at the overall temporal population trends using the
stage subdivisions of Kennedy & Klinger (1975) as reference points. It is clear
from a comparison of Figures 67, 76-77 and Table 15, that the response to
environmental change was largely accomplished by changes in the proportions of
the various species present, rather than wholesale invasion or withdrawal of
large numbers of different types. For instance, the similarity between the species
composition of stage subdivisions in the Campanian I to Maastrichtian II
sequence does not fall below 60 per cent, and rises steadily through the
Campanian from 61 per cent between Santonian III and Campanian I to 77 per
cent between Campanian IV and Campanian V (Fig. 76, Table 15). This is
despite the fact that several of the subdivisions (Campanian IV for example)
contain populations that represent sedimentary environments ranging from
moderate depth inner-mid shelf (assemblage 4b) to deep-water outer shelf/upper
continental slope (assemblages 6-7). This basic continuity in species presence is
further borne out by the curve which plots percentage of species in a stage

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 161

mn
a |
Pes / \
80 ] : oie / \
| ie / i, » i
ak ca aa “|
‘| 7 ~.inherited
! A | spp.
60 ! =e similarity
|
\
40 \
" : ee _- population
ci : Se appearin
§ ! ee pe a | pp g

20

Mili Mil Mi CV CIV Cill Ctl Cl Sti Stl

Fig. 76. Ostracod population trends plotted by stage subdivision in Santonian II to Maastrich-

tian III of Zululand and Agulhas Bank. Curves for inherited species, similarity, and population

appearing (residual of inherited species curve) are expressed as percentages of total ostracod

populations. Number of species extant is given for total ostracod population. Data from Figure
77. See text for discussion.

TABLE 15

Biostratigraphic data (species distribution) on Campanian-Maastrichtian ostracoda
from Zululand (BH-9, and Monzi, Mfolozi and Nibela) and Agulhas Bank.

Sant. III Camp.I Camp. II Camp. III Camp. 1V Camp. V Maas.I Maas. II Maas® III

no. species extant. : 21 24 24 17 22 24 32 29 15
inherited (%) . : ; 14 (67) IA GAD) 19 (79) 16 (94) 16 (73) 20 (83) 21 (66) 23 (93) 8 (53)
restricted (%) . : ‘ 1 (5) 2 (8) 2 (8) 1 (6) 0 2 (8) 2 (6) 2 (10) 7 (47)
appearances (% total) . 7 (32) 7 (29) 5 (21) 1 (6) 6 (27) 4 (17) 11 (34) 2 (7) 7 (47)
disappearances (% total) 4 (19) 5 (21) 8 (33) 1 (6) 2 (9) 3 (13) 5 (16) DTD) i
Similarity

no. species F : : 28 29 25 Py 26 35 34 36
common species ; : V7) 19 16 16 20 21 27 8

% similarity . F . 61 66 64 73 We 60 79 22

no. species i ; ; 45 60

common species ° F 17 P|

% Similarity ; : ; Sant./Camp. similarity 37% Camp./Maas. similarity 35%

no. species extant a 40 41

inherited (%) . : : 2 17 (43) 21 (51)
restricted (%) . : : ? 10 (25) 20 (49)
appearances ( % total) 2358) 20 (49)

disappearances (% total) 19 (48) at least 26 (>63 %)

162 ANNALS OF THE SOUTH AFRICAN MUSEUM

subdivision inherited from the subdivision below (Fig. 76)—a steady increase
from Santonian III (67%) to between 83 and 94 per cent in Campanian III to
Campanian V—and is matched by low values (10-30%) for species restricted to
one stage subdivision, and the low number of species (ten, 18%) that are
restricted to two or less of the various ostracod assemblages.

The largest decline in similarity and inherited species content, and the main
event that is the exception to the general trend outlined above, occurs across the
Campanian—Maastrichtian stage boundary (Fig. 76), where both values fall from
80 per cent to around 60 per cent, and then climb rapidly to their former values
higher in the Maastrichtian. This discontinuity is caused by the appearance of a
relatively large number of new species (~30%) that is not accompanied by any
marked change in the overall rate of extinction (~15%). It is likely that several
of these species, which give assemblage 7 its characteristic composition, repre-
sent special types that became established in response to stable, deep-water
conditions and whose local ‘extinction’ followed the reversion to shallower
environments in Maastrichtian II (Figs 67, 77). In all probability, therefore, the
local temporal ranges of species such as Pariceratina hirsuta, Hermanites? arcus,
Cytheropteron cf. westaustraliense, indet. sp. 2, and Cativella? dubia do not
reflect their ‘absolute’ temporal ranges, i.e. they represent a facies fauna. On the
other hand, some of the new species that established themselves early in
Maastrichtian I did not become extinct with the re-establishment of assemblage
5a, 5b and these types (e.g. Klingerella aranearius, Pedicythere fragilis, Ponticu-
locythere biremis, and Cytheropteron brenneri) may, therefore, be more age
diagnostic than the former, short-range group. Species in the Campanian whose
short ranges and associations with particular assemblages similarly suggest strong
environmental control, and therefore lessen their use biostratigraphically, are
Trachyleberis minima (Campanian I), Unicapella reticulata (Campanian II), and
Curfsina monziensis (Campanian V).

In line with the high percentage species similarities between stage subdivi-
sions, the percentage similarity between stages is stable, decreasing only very
slightly up the succession—Santonian/Campanian 37%, Campanian/Maastrich-
tian 35%—with 58 per cent of the Campanian species appearing within that
stage compared to 49 per cent in the Maastrichtian.

One notable feature of Table 15 is the increase in number of species per
stage from 22 in the Santonian to 41 in the Maastrichtian. Plotted by stage
subdivision (Fig. 76), this increase has two peaks: Campanian I and II (24) and
Maastrichtian I (32), with an intervening low (16) in Campanian III which
follows a high in species extinction (30%) in Campanian II and from which the
faunas did not start to recover until Campanian IV. Whether this phenomenon is
environmentally controlled, or reflects overall phylogeny is not known, but it is a
point we shall return to later.

Of the total number of species in the Campanian—Maastrichtian strata of
Zululand and the Agulhas Bank (60), 17 are inherited from the Santonian III, 10
are restricted to the Campanian, 20 restricted to the Maastrichtian, and the

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 163

remaining 13 restricted to Campanian—Maastrichtian. The ranges of these are
shown in Figure 77, on the left of which are plotted the species inherited from
the Santonian III. There is a sharp decrease in the numbers of these species in
Campanian II (four become extinct), but thereafter their numbers show a slow
but steady decline so that four only are known to extend above Maastrichtian II.

Of the 10 species restricted to the Campanian, 7 had appeared by the end of
Campanian II, and of the 20 restricted to the Maastrichtian, 11 had appeared by
the end of Maastrichtian I. The curve showing number of appearances as a
percentage of total population (which is a residual of the ‘inherited species’
curve, Fig. 76) gives an assessment of evolutionary activity. It can be seen that
whilst the Maastrichtian I was marked by an evolutionary ‘burst’ (33%), the
early part of the Campanian seems to have been part of a downward cycle from
Santonian III (32%) that reached a low of evolutionary activity (6%) in the
Campanian III. :

Ostracod zonal scheme

It is accepted as sound stratigraphic policy to establish, wherever possible,
local biostratigraphic zonal schemes that are based on benthic organisms, but
that can be tied in with an established regional zonation based on planktonic
and/or necktonic taxa (such as ammonites, planktonic foraminifera etc). A
prime justification for this is that it frequently allows a good assessment of age to
be made in other local lithofacies in which the internationally accepted zonal
fossils are absent or poorly preserved. Using the distribution of the ostracod taxa
in Zululand, and selecting species which, as far as possible, show palaeo-
environmental tolerance, we propose an ostracod zonal scheme for south-east
Africa. The zonal boundaries are shown on Figures 77-78, and 80 and the zones
defined below. Kennedy & Klinger’s (1975) ammonite zonations are used as a
framework for the scheme. It should be noted that in the following definitions
only particularly characteristic associations are mentioned, and that the com-
plete associations can be identified from Figure 77. The locality numbers given
below are after Kennedy & Klinger (1975).

Amphicytherura zululandensis Zone—range lower and middle part of
Campanian I. Definition: period defined on presence of index species Amphicy-
therura zululandensis. Remarks: the lower part carries the association Rayneria
nealei, Amphicytherura tumida, and Cytherelloida contorta with Amphicytherura
zululandensis. In the upper part, the associated ostracods are Haughtonileberis
haughtoni, H. nibelaensis and Oertliella africana. So far recognized entirely from
strata within the BH-9 borehole.

Un-named Zone—range uppermost Campanian I. Definition: period be-
tween the last appearance of Amphicytherura zululandensis and the first appear-
ance of Hermanites kennedyi. Remarks: no short-ranging species have been
identified in this period, which includes the top of the range of Haughtonileberis
haughtoni. So far recognized entirely from strata within the BH-9 borehole.

ANNALS OF THE SOUTH AFRICAN MUSEUM

164

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CAMPANIAN AND MAASTRICHTIAN OSTRACODA 165

Hermanites kennedyi Zone—range Campanian II to middle Campanian IV.
Definition: period which begins with first appearance of Hermanites kennedyi
and ends with the first appearance of Dutoitella dutoiti. Remarks: can be divided
into three subzones and has been recognized in strata from the BH-9 borehole
and at outcrops in Nibela Peninsula (localities 110 and 113).

Haughtonileberis nibelaensis Subzone—range Campanian II. Definition:
period marked by the presence of Hermanites kennedyi and Haughtonileberis
nibelaensis. Remarks: the lower part carries the association of Oertliella pennata,
Unicapella reticulata, and Haughtonileberis vanhoepeni with Haughtonileberis
nibelaensis and Hermanites kennedyi. The middle and upper parts carry the
association Unicapella sacsi and Haughtonileberis vanhoepeni with Hermanites
kennedyi and Haughtonileberis nibelaensis. Recognized in the BH-9 borehole
and at outcrop on the Nibela Peninsula (locality 110).

Un-named Subzone—range Campanian III. Definition: period marked by
the presence of Hermanites kennedyi between the last appearance of Haugh-
tonileberis nibelaensis and the first appearance of Oertliella maastrichtia.
Remarks: no short-range species have been identified in this period which
includes the upper part of the range of Haughtonileberis vanhoepeni and the one
record of Apateloschizocythere mclachlani. Recognized at outcrop on the Nibela
Peninsula (locality 110).

Oertliella maastrichtia Subzone—range early and middle part of Campanian
IV. Definition: period marked by the presence of Hermanites kennedyi, begin-
ning with the first appearance of Oertliella maastrichtia and ending with the first
appearance of Dutoitella dutoiti. Remarks: the lowermost part of this zone
coincides with the appearance of several new species: Pontocyprella nibelaensis,
Krithe nibelaensis, Platella africana, which form a distinctive association with
Hermanites kennedyi and Oertliella maastrichtia. Recognized at outcrop on the
Nibela Peninsula (locality 113).

Dutoitella dutoiti Zone—range late Campanian IV to early Maastrichtian II.
Definition: period defined on the presence of index species Dutoitella dutoiti.
Remarks: can be divided into three subzones, and has been recognized at
outcrop on the Nibela Peninsula (locality 113), in the Monzi road section
(locality 21) and along the Mfolozi River (locality 20).

Un-named Subzone—range upper Campanian IV. Definition: period marked
by the presence of Dutoitella dutoiti and Hermanites kennedyi that begins with
the first appearance of Dutoitella dutoiti and ends with the first appearance of
Parvacythereis monziensis. Remarks: carries the association of Hermanites ken-
nedyi, Dutoitella dutoiti and Oertliella maastrichtia. Recognized at outcrop on
the Nibela Peninsula (locality 113).

Parvacythereis monziensis Subzone—range Campanian V to early Maastrich-
tian I. Definition: period marked by the presence of Dutoitella dutoiti and
Parvacythereis monziensis that begins with the first appearance of Parvacythereis
monziensis and ends with the first appearance of Klingerella aranearius.

166 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks: carries the association of Dutoitella dutoiti and Parvacythereis mon-
ziensis. In addition, in the lower part (Campanian V) it includes the uppermost
range of Hermanites kennedyi as well as the typical Campanian V species
Curfsina monziensis. The early Maastrichtian I part sees the appearance of
Pterygocythere lanceolata and the extinction of Oertliella africana. Recognized at
outcrops in the Monzi road section (locality 21) and along the Mfolozi River
(locality 20).

Klingerella aranearius Subzone—range middle Maastrichtian I to late Maas-
trichtian Il. Definition: period defined by the presence of index species Kling-
erella aranearius, together with Dutoitella dutoiti. Remarks: this zone, whose top
cannot at present be located precisely, is typified by rich and varied faunas that
include the association of Pterygocythere lanceolata, Ponticulocythere biremis,
Pariceratina hirsuta, and Hermanites? arcus, with Klingerella aranearius and
Dutoitella dutoiti. Recognized at outcrop along the Mfolozi River (locality 20).
For a finer subdivision of the Maastrichtian strata it would be desirable to erect
further subzones at the top of the Dutoitella dutoiti zone (i.e. to shorten the
Klingerella aranearius subzone). This may be possible with further work, but at
present all the short-range species available (e.g. Cytheropteron spp., indet spp.
3 and 4) are either too poorly known, or seem too environmentally-bound to be
suitable.

Agulhasina quadrata Zone—range ?Maastrichtian III. Definition: period
defined on the presence of index species Agulhasina quadrata. Remarks: the rich
and varied cytheracean assemblage of Agulhasina quadrata, Dutoitella mimica,
Trachyleberis schizospinosa, Paraplatycosta reticulata, and Phacorhabdotus?
anomala is typical of a zone whose top and bottom cannot yet be precisely
defined. Recognized at outcrop on the sea-floor of the outer Agulhas Bank
(locality 818).

Phylogeny: higher taxa

Figure 78 and Tables 16 and 17 reveal that, although the Cytheracea are
numerically dominant only in Campanian I (Fig. 67), they are the most diverse
group both in species and genera throughout the Campanian and Maastrichtian,
and that they show considerable evolutionary activity during the last 13 m.y. of
the Cretaceous in south-east Africa. Expressing this in terms of distribution and
appearance of new species by family (Table 17a), the Trachyleberididae are seen
to be the most diverse and phylogenetically active family within the Cytheracea.
This diversity is greatest in Campanian II, where the number of species belong-
ing to the Trachyleberididae reaches 82 per cent of the cytheracean component
(Fig. 79). This value falls steadily to 42 per cent in Maastrichtian II, indicating a
relative reduction in evolutionary activity within the family in progressively
younger strata. (The sharp rally to 67 per cent in Maastrichtian III is based on
one sample only and cannot be taken as representative.) This decline in
phylogenetic activity within the Trachyleberididae is also evident if the figures
for number of species appearing per stage (expressed as a percentage of the total

zone

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CAMPANIAN AND MAASTRICHTIAN OSTRACODA

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Fig. 78. Temporal distribution of ostracods, arranged by higher taxa, in Campanian—Maastrichtian strata of Zululand (BH-9, and
Monzi, Mfolozi and Nibela outcrops) and Agulhas Bank. See Figure 77 for explanation of left and right-hand columns.

168 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 16

Extant cytheracean species in Campanian-Maastrichtian of Zululand
(BH-9, and Monzi, Mfolozi and Nibela), Agulhas Bank, and Igoda.

no. cytheracean total ostracod % cytheracean

spp spp spp
Maastrichtian III. : : 9 15 60
Maastrichtian II. 19 29 66
Maastrichtian | Cows. . 23 32 TZ
Campanian V : 17 24 7
Campanian IV ; 14 22 64
Campanian III ; : 11 1, 65
Campanian II ‘ ; : 16 24 67
Campanian I . , : 16 24 67
Igoda : : : : 5 2 is 71

80

40

Mili Mil Mi CV CIV Clill Cll Cl

Fig. 79. Number of extant species of Trachyleberididae as a percentage of total number of
cytheracean species in Campanian [—Maastrichtian III strata of south-east Africa. Data from
Zululand and Agulhas Bank.

cytheracean element) are considered: 63 per cent in the Campanian and 44 per
cent in the Maastrichtian. The only other diverse cytheracean families are the
Brachycytheridae and Schizocytheridae (<20%) in the Campanian, which are
joined by the Cytheruridae (~12%) in the Maastrichtian. However, of these,
only the Schizocytheridae (21% Campanian) and Cytheruridae (25% Maastrich-
tian) show modest levels of evolutionary activity (expressed in terms of number
of species appearing).

Whatley & Stephens (1976) made a literature survey of the worldwide
Mesozoic development of the Cytheracea, and it is interesting to find that
several of their conclusions are corroborated by our work in south-east Africa.
At the specific level, they found that whilst the total number of new cytheracean
species was higher in the Maastrichtian (375 spp compared to 360 in the
Senonian, their fig. 2), expressed as a percentage of total species (their table 2),
there was a decline in evolutionary activity across the Senonian—Maastrichtian
boundary (76% to 65%). We have data only for the upper part of the Senonian

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 169

TABLE 17

Data on cytheracean ostracods by family in Campanian-Maastrichtian
of south-east Africa.

a. Number of species appearing per stage by family (evolutionary activity of families):
Data from Zululand (BH-9, and Monzi, Mfolozi and Nibela outcrops)

Campanian Maastrichtian total

Brachycytheridae . Pp 0 1-6, iy,
Cytheruridae . ‘ ee ee A EDA 5° 14%,
Schizocytheridae . oe eee | BA he 160 Y 14%
Trachyleberididae : s G2. 627, 7 447, 19: 347,
Cytherideidae a oh Or, a | ees ba
Xestoleberididae 1 0....0 | Ee be
Cytherettidae . Ue O yA t 3O0
Bythocytheridae -. -- .° 0 0 | Yn 58 | aire eA
Indet. . PALS 0 0 be S677 ee 2 ss
19 16 35

b. Number of extant species per stage subdivision by family (diversity of families).
Data from Zululand and Agulhas Bank

Camp.I Camp. II Camp. III Camp.1¥V Camp. V Maas.I Maas. II Maas. III

Brachycytheridae ele IS 2 13% DSi. ON AMF 2 hey 313%, BD VGNIA 0 0
Cytheruridae : 7 307 0 0 0 0 O 0 0 SSA SORIA Danlilee/ 0 O
Schizocytheridae se AREA lL 6% 2 18% lL BY, 2 12, 2 OY PY INA ne,
Trachyleberididae . 12 74% 13 82% 7 64% 9764-7, 10 59% 10 43Y 8 42% 66774
Cytherideidae 0 0 0 0 OO Pig he 1 6F ia 7/4 1 ESTA ea,
Xestoleberididae 0 0 0 O 0 0 ll 24% I tA L o4Y, sy ey
Cytherettidae , oO 0 0 0 O 0 O 0 O t ay, ile SA OF 0
Bythocytheridae 0 0 0 O 0 0 0-0 0 O fe, ay, Ik Se 0. 0
Indet. . 0 0 0 O 0 0 0 0 0 0 RCA ie SA 0 O
16 16 11 14 17 23 19 i 9

c. Total cytheraceans: Zululand and Agulhas Bank

Campanian Ma astrichtian

Total number of species : . 30 29
New species appearing : J 08D. 63 14 48%
Total number of genera . a 23
New genera appearing . 2 : l.. 6% Ce be A

but these suggest a similar decrease: 63 per cent (Campanian) to 48 per cent
(Maastrichtian) (Table 17c). Whatley & Stephens (1976) also found that at the
family level, the Trachyleberididae exhibited the highest worldwide level of
cytheracean evolutionary activity during the Senonian—Maastrichtian, but de-
tected a decrease in this activity across the Senonian—Maastrichtian boundary
(~300 spp to ~250 spp, their fig. 1), which is precisely what our data show
across this boundary (63% to 44%) (Fig. 79, Table 17a). The high level of
cytheracean evolutionary activity, with a decrease across the Campanian—Maas-
trichtian boundary, and the dominance of the trachyleberids in this activity are,
therefore, local manifestations of worldwide phenomena.

170 ANNALS OF THE SOUTH AFRICAN MUSEUM

Other trends relating to evolution at the generic level have been noted in
the taxonomic section, and suffice to summarize here by reiterating that the two
most diverse genera are Haughtonileberis and Oertliella with four species each.
Haughtonileberis is at its most diverse in Campanian I-II when all four species
are extant, and, although this is the period when it reaches its greatest numerical
importance in the Campanian—Maastrichtian strata, it is in reality at the latter
stages of a numerical decline that commenced in the Santonian II. Oertliella, on
the other hand, reaches its local acme much later (Campanian IV to Maastrich-
tian I).

Phylogeny: species distribution (extinctions)

From a commercial point of view, microfaunal data are of limited use when
presented in conventional phylogenetic tables that:emphasize appearances (e.g.
Fig. 77). Because exploration boreholes mainly produce chippings which are
liable to contaminate downhole sections, the only satisfactory way of plotting
species distribution is to identify their ‘tops’ (extinction points, first downhole
appearances). Whilst having limited scientific value, such charts are of consider-
able practical application, and Figure 80 shows an exploration range chart for
the Campanian—Maastrichtian ostracods of south-east Africa based on their
‘tops’. The ammonite stage subdivisions and the proposed ostracod zonal
scheme are included.

Of the 45 species that ‘appear’ in the downhole sequence (i.e. locally
become extinct), 26 of these do so in the Maastrichtian I and II, although the
position of 11 of these cannot at present be located precisely without further
sampling across the Maastrichtian II-III boundary. The ostracod zonal scheme
proposed herein does not provide a more sensitive time base in this part of the
stratigraphic column, and clearly further subdivision is desirable.

COMPARISON WITH OTHER GONDWANIDE LOCALITIES

Ostracods of Campanian—Maastrichtian age have been described from va-
rious Gondwanide localities in the southern hemisphere and these have been
plotted on a palaeogeographical reconstruction of Gondwanaland at 65 m.y.
(late Maastrichtian) (Fig. 81). Several previous workers have made comparisons
of the Upper Cretaceous ostracod assemblages between the various regions of
Gondwanaland (e.g. Bertels 1977; Kr6mmelbein 1972; Dingle 1969; Bate &
Bayliss 1969; Bate 1972; Neale 1975) but all were hampered by lack of sufficient
detail from south-east Africa which lay in a central position between the South
Atlantic and Indian Ocean areas.

Because there are few, if any, species in common between south-east Africa
and the other areas on Figure 81, comparisons of faunal similarity have to be
made at the generic level. This may have serious drawbacks, not the least being
taxonomic problems and the comparison of unequal statistical detail, but for a
general commentary it proves useful. The percentage similarity between Cam-
panian and Maastrichtian ostracod populations of south-east Africa (Zululand

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 171

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172 ANNALS OF THE SOUTH AFRICAN MUSEUM

W Africa Tanzania
C-9°/o M-7°/c C-14°/o, M-17°/o
J \
SE Africa
sen Perth
°356 Carnarvon
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Fig. 81. Similarity at the generic level between Campanian and Maastrichtian ostracod faunas

of south-east Africa (BH-9, and Monzi, Mfolozi and Nibela outcrops, Zululand) and other

Gondwanide localites. Reconstruction for 65 m.y. after Firstbrook et al. (undated). Ostracod

population data after various authors cited in the text. Abbreviations: C = Campanian,
M = Maastrichtian, IM = lower Maastrichtian, mM = middle Maastrichtian.

and Sample 818) and rocks of similar ages elsewhere at generic level is shown in
Figure 81. We shall briefly review these areas.

Western Australia

Campanian—Maastrichtian sediments are represented in the Carnarvon
Basin of Western Australia by the Mirig Marl, Korojon Calcarenite, and
Toolonga Calcilutite. Bate (1972) described the Campanian fauna of the area
but gave no data on the Maastrichtian assemblages. Several of the species that
Bate recorded in the Campanian were also recorded by Neale (1975) in the
Santonian Gingin Chalk of the Perth Basin farther south. There is a 30 per cent
inheritance of species across the Santonian—Campanian boundary in the Carnar-
von Basin, with 24 per cent and 38 per cent similarity at the specific and generic
levels, respectively. Table 18 shows the genera grouped into families present in
the Carnarvon Basin Campanian (averaged for three samples described by Bate
(1975) ). Although no species are common with south-east Africa, the two areas
show a relatively high (27%) generic similarity.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 173

TABLE 18

Genera recorded in the Campanian of Carnarvon Basin, Western Australia.
°% total fauna, mean of 3 samples recorded by Bate (1972).

Cytherellidae . : : +Cytherella, +Cytherelloidea and *Platella . 27.
Bairdiacea . t+Bairdoppilata and +Bythocypris ; , 9%
Cypridacea . +Paracypris and +Pontocyprella_. Ay,
Cytherideidae . ; : +Krithe . : ‘ : : : : : =< A 5°/
Eurotundracythere SA .

Cytheruridae . . +Cytheropteron A : 10%,

Oculocytheropteron : : il yA
Cytherura , ; ' 4 a A 12%

Paracytheridea : : ' : at (yA

+Pedicythere . , : i i é 21%

Bythocytheridae : : B ythoceratina ; 3 : ' , A
CytheralisOn 2 . . .: A Leif SA

Monoceratina ' , : ; : : ry,
Progonocytheridae .. Tickaloracyinere 4) 0 4% 4%
Schizocytheridae . +tApateloschizocythere . ; : 127, 12%,
Pectocytheridae : Premunseyella ; : ; : Pee fe Spey

Trachyleberididae . : Cosia . 5 : : iA

+Hermanites . : : ; ; : : ao

Karsteneis , : : ; : ; ‘ 49,

Toolongella . : i 6%
Anebocythereis : é eA ZT 7,

+Curfsina : , ; aig A

Limburgina _ . : : | eA

¢ +Oertliella : ; 4%

+Trachyleberis ; 5 ae i oe ; | A

Orthonotocythere . : : : =,

Hystricocythere . : : : . <1%

Genus B ; |e

Similarity with south-east Africa (+):
32 genera Australia, 24 genera SE Afr., 56 total, 15 common = 27%

In addition to the cosmopolitan genera such as Cytherella, Cytherelloidea,
Bairdoppilata, Trachyleberis, and Cytheropteron, notable similarities are Pedicy-
there, Pariceratina (Santonian of Perth Basin), Apateloschizocythere, and Curf-
sina. In addition, three species found in Western Australia, Hermanites saggita
(Santonian—Campanian), Oertliella exquisita (Campanian), and Bythocypris
chapmani (Santonian), have very close relatives in south-east Africa: Hermanites
kennedyi (Campanian), Oertliella maastrichtia (Campanian—Maastrichtian), and
Bythocypris richardsbayensis (Santonian—Maastrichtian). As Neale (1975) has
pointed out, the subfamily Pennyellidae is confined to the South African—
Australian—West Pacific areas, albeit at various stratigraphic levels, in the Upper
Cretaceous.

Notable differences between south-east Africa and Western Australia are
lack of the genera Brachycythere and Haughtonileberis in Australia, and the fact
that Majungaella/Tickalaracythere and Rostrocytheridea, which are common in
the Lower Cretaceous of south-east Africa, have not yet been recorded from the
Upper Cretaceous of this area, whereas they do occur in the Santonian—

174 ANNALS OF THE SOUTH AFRICAN MUSEUM

Campanian of Western Australia. Overall, seventeen species (numerically 53%
of the Australian fauna) belong to genera not recorded from south-east Africa.

As can be seen from Figure 81, the Campanian of south-east Africa at the
level of generic similarity is faunally closer to Western Australia than it is to any
of the other areas of Gondwanaland.

East Africa

Ramsay (1968) and Bate (in Bate & Bayliss 1969) have described Cam-
panian—Maastrichtian ostracods from Tanzania (Table 19a). Judging by the small
number of specimens recorded, it would seem that the faunas have been only
partially examined, and because of this, comparisons with other areas must be
tentative.

The late Campanian assemblage of the Runyu inlier area shows a 14 per
cent similarity at the generic level with south-east Africa, but all Bate’s positive
identifications were in the cosmopolitan types. His Genus B bears some resembI-
ance to Agulhasina, but his sketch (Bate & Bayliss 1969, pl. 7 figs 11, 14) is not
sufficient for a positive comparison. However, the Maastrichtian, with a 17 per
cent generic similarity with south-east Africa, has a more varied fauna with the
notable presence of Dutoitella mimica. Other points of similarity between the
two areas are the presence in the late Turonian of Tanzania of Brachycythere
aff. B. sapucariensis which Bate & Bayliss (1969) compared to B. longicaudata
(Santonian—Maastrichtian, south-east Africa), and the similarity of three other
Tanzanian Turonian species to younger south-east African counterparts: Curf-
sina turonica to Parvacythereis monziensis; Cythereis luzangaiensis to C. klingert;
and the general similarity of the genus Akrogmocythere to Gibberleberis.

West Africa

Apostolescu (1961, 1963) and Reyment (1960) have documented the ostra-
cods from various parts of west Africa (Gambia to Nigeria), and Table 19b lists
the genera and species recorded by Apostolescu in his two reviews. We have no
numerical population data, but the assemblages are characterized by numerous
species of Brachycythere, Buntonia, Ovocytheridea, Cophinia, and Nigeria.
Similarity, at the generic level, with south-east Africa is low (Campanian 9%,
Maastrichtian 7%) and the only point of correlation between the two areas
recorded so far is the relative diversity and local abundance of Brachycythere.
Ovocytheridea may be represented in south-east Africa by Pondoina, and one
species of Veenia (in the Santonian at Umzamba) is the only relative of Nigeria.
Bate & Bayliss (1969: 164) have also remarked that the only point of reference
between east and west Africa in the Upper Cretaceous is the presence of
Brachycythere, though it should be noted that they also recorded Ovocytheridea.

Dingle (1969) concluded, after a preliminary examination of the south-east
African Santonian faunas, that there is very little similarity between this area
and west Africa. This conclusion is substantiated by the present, more extensive
studies.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 175

TABLE 19

Campanian-Maastrichtian ostracod genera from east and
west Africa.

a. genera recorded in Tanzania (after Ramsay 1968; Bate & Bayliss 1969)

Campanian Maastrichtian
Prerygocythereis : : : %
+Trachyleberis : :
+Genus C (Dutoitella)
*+Cythereis
+Phacorhabdotus
+Krithe
Genus B ‘ : : F
+ Bairdoppilata F ' : , *
+Cytherella
% +Cytherelloidea

xX &¥ & &* ¥

Be ee ES Re

Similarity with south-east Africa (*):
Campanian: 5 genera E. Afr., 24 genera SE Afr., 29 total, 4 common = 14%
Maastrichtian: 7 genera E. Afr., 29 genera SE Afr., 36 total, 6 common = 17%

b. genera recorded in west Africa (after Apostolescu 1961, 1968)
(number of species in parenthesis)

Campanian Maastrichtian
#(1) +Brachycythere . ; : > *(3)
#(3) Ovocytheridea , ;

a (6) Buntonia ; ; : , : (4)
#(10) t*Cythereis’ . : : : ; *(2)
*(3) Cophinia : : : #(1)
#(1) Nanocythere ; : *(1)

Dactylia : ; : : *1(1)
Protobasslerites . ; : : *(1)
Soudanella . : : : : #(1)
Isobuntonia . : , ; : #(1)
Anticythereis : ; *(1)
=(1) ‘Clithrocytheridea’ : ;
*(3) Veenia (Nigeria) ; : f *(5)
*(2) +Cytherella . ‘ ; *(2)
*(1) Sphaerolevberis ~. sks *(1)

Similarity with south-east Africa (+):
Campanian: 10 genera W. Afr., 24 genera SE Afr., 34 total, 3 common = 9%
Maastrichtian: 13 genera W. Afr., 29 genera SE Afr., 42 total, 3 common = 7%

South America

The only detailed documentation from the western South Atlantic margins
has been from Argentina, where Bertels (1974, 1975, 1977) has described the
Lower and Middle Maastrichtian ostracods of the Jagtiel Formation and its
equivalents (the Campanian is represented by a non-marine facies). Two distinct
faunas are evident: a very shallow marine Lower Maastrichtian fauna at the
base, and an overlying Middle Maastrichtian, deeper water (?mid-shelf) assem-
blage (Table 20). Neither shows any strong similarity with south-east Africa, but
the deeper water fauna has greater affinities (as would be expected): Lower

176 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 20

Genera recorded in Maastrichtian of Argentina.
Data after Bertels (1974, 1975).

a. Lower Maastrichtian (Lower Jagtiel Formation):

+Cytherella . f : ; ‘ é 6%
Jonesia ? : ‘ ‘ ; f : : 8%
Cytherura : : : : : : 8%
Paracytheridea : : : 1%
Trachyleberididae .. Alatacptheré 4 i A OO aoe
*+Trachyleberis ot i ORs 5 aie ee aA E
Platacythereis? . : ‘ : : : Pai ges 1%
Wichmannella ; : : : : : As ge
Similarity with south-east Africa (*):
8 genera Argentina, 29 genera SE Africa, total 37, 2 common = 5%
b. Middle Maastrichtian (Upper Jagtiel Formation):
+Cytherella . ; j : : : : 4%
+Bythocypris ? : : : : : ‘ A
+Paracypris . ; ; ; ; : : 4%
Trachyleberididae . E Togoina ; ‘ ; ; Le
+Trachyleberis ; , : a
Actinocythereis . : : : : : ee
Acanthocythereis . : : : : : eA
+Cythereis? . ; ; : ; Wy 48%
Henryhowella : : ; ; . ‘ py
Protocosta . : : ; SA
Veenia (Nigeria) . : : : , : eye
Wichmannella : : ! ‘ Pex
Hemicytheridae : ; Bradleya ? : . : : . 2 : 4a”% 21°
Anticythereis . ponds Ship waiatenet: Sage ame oe C
Pectocytheridae 3 : Munseyella . : : 2 : , ; ry
Progonocytheridae .. Mosacleberis?- 4 4.5 ar ee oe 3%
Tumidoleberis : , ‘ Le 6%
Sphaeroleberis? . : 2
Cytheruridae . : . +Cytheropteron : : : 4 : ‘ i
Hemicytherura : : : : : ; Le 12%
Cytheromorpha? . : : : : : 10%
Schizocytheridae . +tAmphicytherura? . : : ; : eA

Similarity with south-east Africa (+):
22 genera Argentina, 29 genera SE Afr., total 51, 7 common = 14%.

Maastrichtian 5 per cent and Middle Maastrichtian 14 per cent. It should be
noted, however, that even this low level of similarity may be misleadingly high
because it is based largely on the cosmopolitan types, Cytherella, Paracypris and
Cythereis. Even here uncertainty is introduced by Bertel’s use of ‘Cythereis’ as a
sack term (1975: 100): ‘true Cythereis forms ... do not occur in the known
South American fauna.’ Of the Argentinian trachyleberids (which numerically
make up 48% of the population), only Trachyleberis occurs in south-east Africa.
The similarity between Argentinian and south-east African uppermost Cre-
taceous ostracod faunas is, therefore, even lower than the 14 per cent suggested
by Figure 81. This is in strong contrast to the lower and middle Cretaceous

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 177

faunas, where several species are common to the two areas (e.g. Musacchio
1979). Conversely, there is 14 per cent similarity at the generic level between the
Maastrichtian faunas of Argentina and west Africa. Although this is the same as
the Argentinian and south-east African connection, it does in reality represent a
much stronger link because it occurs in characteristic trachyleberid genera such
as Togoina, Anticythereis, and Nigeria. Significantly, the latter is numerically
and specifically the most diverse and important single genus in both areas.

A similarity in deep-water faunas between south-east Africa and South
America is indicated, however, by the presence of the closely related genera
Unicapella and Paleoabyssocythere, and Dutoitella and Atlanticythere. Benson
(1977) recorded the South American types from the Rio Grande Rise (DSDP
sites 356 and 21) in the western part of the ocean basin, which at the time of
deposition lay relatively close to the mid-Atlantic ridge. The South African types
were deposited in water shallower than the 1 000 m suggested by Benson (1977),
but whether Unicapella and Dutoitella were restricted to the continental margins
is not known. If they were, then communication could have been via the Rio
Grande Rise—Walvis Ridge archipelago. Conversely, they could have evolved
separately from common early or mid Cretaceous ancestors.

ACKNOWLEDGEMENTS

~ It is a pleasure to thank Dr H. C. Klinger of the South African Museum for
guidance during sample collecting in Zululand, for providing the sediment
samples from Igoda and Needs Camp, and for valuable discussions on the
stratigraphy of south-east Africa. I thank Dr R. Benson of the Smithsonian
Institution for permission to publish sketches of some of his unpublished
ostracods, and the Director of the South African Museum for the loan of
Chapman’s types from Needs Camp. The Management of SOEKOR kindly
made available samples from the JC—1 borehole. Permission to collect samples
from Umzamba was kindly granted by the Secretary to the Minister of Agricul-
ture, Forestry, and Fisheries in the Transkei Government. Mr G. Lowcock and
Dr D. Crawford of the University of Cape Town SEM unit are thanked for their
assistance with photography. A University of Cape Town staff research grant is
gratefully acknowledged for fieldwork and SEM expenses. The Editorial Board
of the University of Cape Town provided a generous grant towards part of the
publication costs for this work.

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6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
comma separates author’s name and year
“semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.

In describing new species, One specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:

Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid- tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. ‘Smith, 15 January 1973.

Note standard form of writing South African Museum registration numbers and date.

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. *... the Figure depicting C. namacolus ...’; ‘. .. in C. namacolus (Fig. 10)...’

' (b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene

(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should be expressed in the third person

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book or article, such as
‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation
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Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

R. V. DINGLE

THE CAMPANIAN AND MAASTRICHTIAN
OSTRACODA OF SOUTH-EAST AFRICA

es FZ 2.

VOLUME 85 PART 2. JANUARY 1982 ISSN 0303-2515

itis COMP. ZOOL,
oO By
MAR 26 1982

eT 8 a | eee

ANNALS

OF THE SOUTH AFRICAN
MUSEUM

CAPE TOWN

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2. LAYOUT should be as follows:

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(a) Author’s name and year of publication given in text, e.g.:

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‘As described (Haughton et al. 1927)...’

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Examples (note capitalization and punctuation)

BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

FISCHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann, Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

(continued inside back cover)

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

Volume 85 _ Band
January 1982 Januarie
ray. ee!

MICROMAMMALS AS
PALAEOENVIRONMENTAL INDICATORS AND
AN INTERPRETATION OF THE LATE
QUATERNARY IN THE SOUTHERN CAPE
PROVINCE, SOUTH AFRICA
By

D. M. AVERY

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000

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Printed in South Africa by In Suid-Afrika gedruk deur
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS
AND AN INTERPRETATION OF THE LATE QUATERNARY
IN THE SOUTHERN CAPE PROVINCE, SOUTH AFRICA

By
D. M. AvERY
South African Museum, Cape Town

(With 33 figures and 50 tables)

[MS accepted 3 March 1981]

ABSTRACT

Micromammalian remains recovered from several archaeological sites in the southern Cape
Province are examined in an attempt to ascertain the potential of such material for providing
evidence of palaeoenvironmental change. Analyses of the composition and structure of the
small mammal communities represented and of the mean size of individuals in populations of
different ages and locations are checked against existing evidence from other lines of research.
The micromammalian evidence, which here covers approximately the last 80 000 years, is
shown to be both reliable and informative on a fine scale. The possibility of using the
micromammalian evidence to correlate undated or insecurely dated archaeological sequences is
discussed. Consideration is also given to the application of the data to archaeological interpre-
tation and general theories concerning human technological development.

CONTENTS

PAGE

a at RO re hy hse Nd eens Bins 2 evens tedibwin «go anes ee oa 4 184
Wie Ci EEGs 1728 in 0) be! an SRN 186
Te Oe re don ge ah ties ges cen FE RE ee De ohm 189
RM ORC SITES © pane code wee nwa De ae ee anode tes ent. 189

Tae OA ete Ie GoM as! ace ie be Sn Lp ets das een PS 200
eR Rs ee Pe a ks knw oo wd adhe awn eta dpe 207
eS eS SST a ae ince 207
Pam er ECU @UNNM 22 oo Sis Acs ess See ss hans SEY ele os 209
ee tee MR nn a nla ci n ah Shs) nt La aenin wok Cte as os > 210

Sie manure -Of Cxeavated Samples . 2 fk ee See ae 213

ie Matire Ol Mucromatinalian evidence... . 2... 5... ee eee 217
PO EE ety es Gat RE Rn 228
PeamMputacrOniOn MMi MUMIDETS ... 6. oe te ees 228
Multivariate statistical analysis of primary data....................... PB,
Peceeeces (OD iMbcrpretine alalySeES .. 2... 6 ee ce pet eee bene 234
Mensiration and calculation Of indices. >..........60 0000. b eke des een. 239

i nN eG AS os Sek ss Seg wine ee sie Oe wage wae Se a» 238
Pm NOUIOM) 2 oe cv Sen ee ee ee et ee a hae eee ee 238
Se ROI TS fo icsi i100) re 243
Puicipreraion Of comparative Samples ...... ....5...-2660 6250s ene. 246
Micromammalian evidence for palaeoenvironmental change................ 258
Evidence Onl COMMUNILY COMPOSITION ............. 05.0 Je ob fe ees 255
Pride Gecsmomimeati size Variation . ... 22... hoc ode ce tee wae ate 207
adie armOnln SMPECICS WIVETSILY . O02... 5. ce ree ee ae ee ees 307

183

Ann. S. Afr. Mus. 85 (2), 1982: 183-374, 33 figs, 50 tables.

184 ANNALS OF THE SOUTH AFRICAN MUSEUM

PAGE

Palaeoenvironmental reconstruction based on micromammalian evidence .... 314
BOOM aS: Ain os ie ss 5 2b oe 3 ees re ae ee eee 314
Byneskyanskop. 1 5 occ ois-0 <0 SGN 5 gd rake ted tee bla senegal Re ee 318

Die Keiders 1: Middle Stone: Age Samples... ic tat ua am ve ae ee ere 320
Cnastal samples... ¢..0. cyan ones < Cok woe va trek oe Sete cn came een eee ee 322
FAOIOCENE. BNC MOGETN SAIIPNES 54.3 h i se es Wek were Rea ene ae ee 323
Upper Pleistocene and modersamples, 5-1 sy.4<..4.s ear oe cheney ae ae 324
Existing eviderice for Late Ouaternary environments 52 \tuc3 1...ao eee ees 326
Marine CVINenee 5 5.2 fof 04: os. saat tls me abate « Monee oer ee 327
EOTFEStEA: CVIGENCO eh oe hs bee sails awk ea eo ee ee 330
Correlation of micromammalian and existing evidence..................... 335
General ramework-<-3, <2. oo-s stn Saeed oneal Poca Sette ee ee 336
SOMeheHY Airican CviIdenee: > 255/24 7. 2 ly coe ele ho conte 338
Moacronwmammmiatian Cvidemes® «4... ius for gu ls. Salat c eeete set onde eee 539
Micromammalian evidencefor site correlation. «04 ics. bei nee. fa See cee 347
Prehistory and environmental chanee 245 sc... nna eae t Bee ee 351
Creperal CONSIDERANONS” «5.5135. eases wt Oke oe ee SSL

‘fhe southern (Cane Prowiniecy . 04.1415 dates Bip eeaclocnas is ak ta ee 356

CO CUISOUNS ss. > 5 inte anc oe gga teas cee ee ata sc eae ce 362
SSUBAIR RARER YE F000 fc ol, gs bees Sreesh erties ates tog tad Scat he Ren eae tec crete ea 364
ACKROMCGEOMNEMES 5.53565 ki ane 5. ORE Te Pe Paleniod Bink tenner ene See 365
FRERCION EES) 5 ha 4 yhoo ti ey Sead Yakicg eit at eed ae pee nae eee 365

INTRODUCTION

Micromammalian remains were first used as indicators of past environmen-
tal conditions in South Africa by De Graaff (1960). His preliminary investiga-
tion of the Lower to Middle Pleistocene rodent faunas from the Krugersdorp
(Transvaal) australopithecine-bearing deposits included a short section on the
climatic variations between the different sites, referring in particular to the
degree of aridity or humidity. Cartmill (1967) also interpreted the micromam-
malian evidence from these sites in terms of annual rainfall, providing support
for the hypothesis that Paranthropus was adapted to moister conditions than
was Australopithecus. Brain (1974) discussed the use of microfaunal remains
from archaeological sites as habitat indicators. He postulated (Brain 1974: 58),
for instance, that the presence of Eremitalpa granti in the Mirabib deposits in
the Namib Desert could be used to indicate the advance of dune fields across
the Kuiseb River. This is because E. granti has been shown to be restricted to
soft sand, whereas at present the Mirabib Hills are surrounded by a gravel plain
for 20 km in all directions. In fact, Brain & Brain (1977) subsequently con-
cluded that, in the absence of E. granti, the dune fields were blocked by the
river. They have further suggested (Brain & Brain 1977: 293) that fluctuations
in the proportions of gerbils and geckos reflect changes in the amount of
rainfall and, consequently, of vegetation. Following on this line, the present
study comprises a comprehensive examination of micromammalian remains
from a number of southern Cape archaeological sites containing Upper Pleis-
tocene and Holocene deposits. The purpose is to examine the nature and

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 185

potential of this material and to establish the extent to which the evidence
obtainable from the material supports or refines current knowledge of climatic
and vegetational fluctuations during approximately the last 80 000 years.

The principal interest in the present work, as in that of Brain (1974; Brain
& Brain 1977), is to contribute detailed information concerning the conditions
under which people lived in prehistoric times. This is a basic aspect of
prehistoric studies since it has become increasingly clear that human cultural
adaptation and development must be viewed in the context of the contempor-
ary natural, that is physical and biological, environment of the people
concerned. In pursuance of the general aim, the collection for study of
micromammalian remains from archaeological sites has two main advantages.
These sites comprise one or more discrete units containing durable remains of
human activities in the past. Where several such units accumulate in a stratified
sequence they may well represent a considerable period of time. Micromamma-
lian remains, although apparently not collected by humans, frequently become
admixed with the archaeological material, either in the same or in intervening
units. They, therefore, provide evidence of conditions pertaining over the same
general period of time as that covered by the archaeological samples, but with
the important quality of being independent of the archaeological evidence. For
archaeology, a potential application of the detailed micromammalian evidence
for environmental change is that of assessing the possible effect of such change
on cultural development. This is particularly the case where units can be shown
by absolute dating to represent relatively short periods of time since the
resolution will then be correspondingly tight. Another potentially useful aspect
of the study is the provision of a means of dating sites relatively. Where
absolute dates are lacking, it may be possible to use detailed evidence of
environmental change to provide relative dates.

The southern Cape was chosen as the region of study partly because, being
here defined as that area bounded by the sea and the Cape Folded Mountains,
it forms a convenient natural unit. Primarily, however, it was chosen because
the number of samples of micromammalian material available from this natural
unit provided maximum potential for investigation. Good sequences of samples
have been collected from three sites in the area. These are Boomplaas A in the
Cango valley, and Byneskranskop 1 and Die Kelders 1 near and on the Walker
Bay coast. Other samples come from Klasies River Mouth 1A and from Nelson
Bay Cave, both on the south coast. The available samples allowed conditions
during the Upper Pleistocene (Last Glacial) and Holocene (Present Intergla-
cial) to be examined. The Boomplaas A data are unique in that they provide
evidence of glacial maximum and full interglacial conditions and thus probably
illustrate the range of variation in response by micromammalian communities
which is to be expected for this area. Modern micromammalian material was
examined both for control purposes and in order to see whether change had
occurred during approximately the last 2 000 years since the end of the subfossil
record available from the archaeological sites.

186 ANNALS OF THE SOUTH AFRICAN MUSEUM

The question of the agency responsible for accumulating the micro-
mammalian remains in caves and any possible idiosyncrasies of that agent
which may bias composition of the samples, was investigated in some detail.
The nature of the evidence forthcoming from the micromammalian remains was
also examined. This concerns factors pertaining to micromammals as well as
factors connected with samples recovered from archaeological or palaeontologi-
cal sites. All these considerations will have a potential bearing on the interpre-
tations which it is possible to make from the material. Any inherent strengths
or weaknesses must be clearly recognized. Another integral part of the study
involved establishing as precisely as possible the environmental significance of
individual species as well as the possible reasons for fluctuations in population
size Or community structure. In this exercise it was necessary to generalize to a
certain extent because data concerning specific factors governing distribution
and reproduction are not available for most of the species.

Data from other disciplines which provide evidence for climatic and
vegetational change were examined in order to provide a context for the study
and also to check the validity of the interpretations made from the micromam-
malian evidence. The general framework is known and has been confirmed
from a number of different lines of investigation. At the local level, however,
much remains to be done. The micromammalian evidence provides detail at
this level and suggests that differential change may be detectable on quite a fine
scale within one region.

Two main lines of investigation were pursued in attempting to interpret the
material. The major of these involved examining variations in the structure of
the small mammal community as a whole. Changes in the dominant species, as
well as fluctuations in the proportions of numerically less important species,
suggest changing environmental conditions which it is possible to interpret
within the limits of the evidence. In most cases changes in proportions of plant
life-forms will be reflected directly whereas evidence for climatic change will be
indirect. Changes in species diversity appear also to give some general indica-
tion of changing conditions. At the population level, differences in the mean
size of individuals in different populations of selected species indicate physical
response to varying climatic conditions of which they are thus evidence.
Although interpretation is currently hampered by lack of comparative modern
data, the extent to which this study is yet able to augment present knowledge of
environmental change during the Upper Pleistocene and Holocene confirms the
potential of these investigations.

THE SOUTHERN ‘CAPE PROVINCE

The southern Cape Province is here defined as that part of the Cape
Province which lies south of 33°S (Fig. 1). At this latitude the east-west branch
of the Cape Folded Mountains bounds a coastal plain between Cape Hangklip
in the west and Cape St Francis in the east. The region under discussion has

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 187

(fe) 100-200 mm NX 400-600

& BOO=R Oe Za 600 +

SSNS eee

DS = SSK oY

fynbos _ renosterveld strandveld
Ne ay aa mountain/arid & coastal ee
ay mn temperate forest
Rises anak coastal S mountain
BS

cal succulent scrub &
Karroid broken veld

Fig. 1. The southern Cape Province. A. Location of described sites. (Position of Cape Folded
Mountains based on Hendey 1974.) B. Mean July temperature (based on Schulze 1965).
C. Rainfall (after Van Zinderen Bakker 1976). D. Vegetation (after Kruger 1977).

been ascribed to parts of geomorphologic provinces 14, 15 and 16 by King
(1963), and to part of region 11 by Wellington (1955). The east-west branch of
the Cape Folded Mountains comprises two principal subparallel ranges, the
northern Swartberg and southern Langeberg Mountains. These are extremely
rugged with deep river-cut gorges crossing them at intervals. Boomplaas A is
one of a series of caves, including the famous Cango Caves, which is formed
along a fault zone in the foothills of the Swartberg Mountains (Roussouw et al.
1964: 87). Between the two mountain ranges lies the Little Karoo which,
although flatter, is still hilly and rocky. The coastal plain tends to be undulating
with some low hills into one of which Byneskranskop 1 and 2 are cut. The caves
that are situated on the present coastline, namely, Die Kelders 1, Nelson Bay
Cave and Klasies River Mouth 1A, were incised by the sea during past
transgressions.

The western part of the region has a Mediterranean climate with winter
rainfall (Schulze 1965). Further east the rainfall is year-round, gradually giving
way to a summer rainfall régime in the south-eastern Cape. Annual rainfall
varies greatly, from as much as 3 000 mm in some mountain valleys to as little
as 250 mm in the Little Karoo and rain-shadow areas such as the Breede River
valley. Extremes of temperature are greatest inland, in the mountains and
Little Karoo, where there is very marked variation in temperature at different
times both of the day and the year. Such a continental régime is in contrast to
that of the coastal region, where temperatures are considerably less variable
and generally mild, with frost being almost unknown.

The natural vegetation of the southern Cape principally comprises fynbos
(Fig. 1). This is basically restricted to the winter rainfall area where the mean

188 ANNALS OF THE SOUTH AFRICAN MUSEUM

annual rainfall is at least 250 mm. In general, fynbos is characterized by a lack
of single species dominance and/or conspicuous presence of Restionaceae, and
physiognomically by restioid, ericoid and proteoid elements (Taylor 1978: 174).
Three major subdivisions have, however, been recognized (Kruger 1977; Tay-
lor 1978). Mountain fynbos tends to have several layers, particularly on the
lower slopes which are characterized by proteoid shrubs, usually 1,5 to 2,5 m in
height. Arid fynbos, which occurs as a narrow belt along the inland lower
slopes of the Cape Folded Mountains (Taylor 1978: 199), tends to be more
open than mountain fynbos, with a simpler structure and a preponderance of
ericoid forms. The coastal fynbos includes many more grasses than the moun-
tain fynbos, with the south-coast variant tending to comprise a lower ericoid
layer and a taller proteoid layer.

Strandveld is another type of vegetation that occurs in the coastal regions
of the southern Cape. Elements of fynbos and forest play a part in the
succession of this type. From pioneer grasses the succession proceeds to a
climax of coastal scrub, the highest development of which is represented by
small forest patches occurring near Stanford (Taylor 1961). Low scrub or
sward occurs in moist depressions near the coast. Coastal renosterveld, domi-
nated by the renosterbos (Elytropappus rhinocerotis), separates the vegetation
of the coastal plain from that of the mountains. Its rich flora includes many
species of grasses (Taylor 1978: 216). Mountain renosterveld occupies a simi-
lar transitional position between the mountain fynbos and the Karoo vegeta-
tion. It occurs at medium altitudes and contains narrow-leafed dwarf shrubs
(Werger 1978: 284). Unlike the coastal renosterveld which has fynbos affini-
ties, the mountain variety contains a considerable Karoo element. Between
the mountain ranges the Little Karoo bears a vegetation of shrubs and dwarf
shrubs with a dominance of succulents and few grasses. Afromontane temper-
ate forest occurs as enclaves in the southern Cape, principally near Knysna
but also on suitably watered and sheltered southern slopes as far west as
Table Mountain (White 1978: 507). The forest varies from being comparable
to Afromontane rainforest in structure to scrubforest and thicket in less
suitable conditions.

The intergradation of one type of vegetation into another illustrates the
dynamic nature of the delicate equilibrium between the various types. The
region of the Cango valley provides a very good example of the complexity that
may exist and of the interaction of different vegetations. In this case the rich
flora reflects the intermediate position of the valley between the moist Swart-
berg Mountains and the drier Little Karoo (Moffett & Deacon 1977: 128). Five
major phytogeographic elements have been recognized in this one area and
vegetation assignable to four of Acocks’ (1975) veld types occurs in the valley.
It is to be expected, particularly in such a complicated situation, that even
minor changes in climate would alter the balance, and there is now direct
evidence from botanical data and indirect evidence from the macromammalian
material of such past changes.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 189

THE SITES

The sites from which the material for this study was collected are all caves
or rock shelters. Within such caves and rock shelters there is a tendency for
debris to build up as a result of the use of these places as shelters by animals
and humans. Over a period of time such debris may accumulate in layers to a
considerable thickness. According to the picture of human occupation patterns
which is emerging from archaeological study, it would appear that occupation
of the caves by people was periodic (e.g. Parkington 1972, 1976; Deacon &
Brooker 1976). Relative increase in the proportion of micromammalian
remains in levels not connected with human occupation indicates the alternate
use of the caves by owls (see below for discussion on the predator involved)
and presumably other animals, particularly birds. The process of accumulation
of layers of micromammalian-rich matrix can be observed in operation in caves
and rock shelters today. Collections were subsequently made from such sites in
an attempt to elucidate conditions prevailing at various times in the past.

ARCHAEOLOGICAL SITES

The sites are described below in alphabetical order. In each case the
location of the site is given, together with the period represented by the
deposits and the cultural sequence as interpreted by the archaeologists who
have studied the human occupation of the site. Some indication is also given of
the quality of the micromammalian samples and the horizons from which they
were extracted.

Boomplaas A (BPA)

Boomplaas A (33°23’S 22°11’E) is situated in the Cango valley about
40 km north of Oudtshoorn, which lies in the Little Karoo (Figs 1-2). It is part
of a fissure system in the Drupkelderkop, some 600 m above sea-level in the
foothills of the Swartberg Mountains. There are approximately 5 m of deposits
(Fig. 3). The time range covered by these deposits is from about 80 000 B.P. to
about 1 500 B.P. (Deacon et al. 1976). There is a period of leaching and wash
about half-way down the sequence, somewhere between 14 200 + 240 B.P. and
21 000 + 420 B.P., which probably represents a short hiatus (Deacon &
Brooker 1976: 209; H. J. Deacon 1976 pers. comm.). Deacon & Brooker
(1976) give a general description of the sequence, noting that contributions to
the deposits were made at various times by material entering through a fissure
at the back of the cave, through the front of the cave by human and other
agency, and as a result of roof spalls and rock tumble. Deacon et al. (1976) give
a more detailed description of the top third of the sequence. Figure 3 is based
on the most recent description by Deacon (1979). The cultural sequence begins
with Middle Stone Age (M.S.A.) at the bottom. This is succeeded at about
32 000 B.P. by an upper M.S.A. industry with long quartzite blades. This is, in
turn, replaced first by an as yet undescribed industry of non-Levallois character

190 ANNALS OF THE SOUTH AFRICAN MUSEUM

Walker
Bay

34°34'S

19°23 'E

Fig. 2. Detailed location of sites. A. In the Cango valley. B. On the Walker Bay coast.
(After Avery 1977.)

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 191

years BP

CBM historic occupation

burnt sheep dung

l m
= 1630750 UW337
DGL prehistoric herder
ISIO* 75 UW307] occupation
= 1700755 UW338
+
BLD 1955765 UW336
= Wilton
BLA 640075 UW306
BRL 91007135 UW4I0
2 104257125 UW4Il
2 124807130 UW4I2
12060 Pta
CL Robberg
= 142007240 UW30I
GWA/
HCA
LP early Late Stone Age /
ia 21100420 UW300
LPC 22000 Pta late Middle Stone Age
3 YOL—
BP
= 32400t700 UW304
a ee __®}»40000 UW305
OLP 42000 Pta Middle Stone Age
vvvvVVYVY¥YVyY
vvvvvyy ‘vy Vv
4 »>40000 UW308
BOL
Boa es ee fe. fb oO
Bree eo OF oo Bo
OCH Poe ep 8 oO Oo DB
ea oe ooo Go oO GB
Bebe ae a ae eae DB
5 LOH

below datum

Fig. 3. Diagrammatic section of Boomplaas A with lithological units, 'C determinations and
cultural units (after Deacon 1979).

ANNALS OF THE SOUTH AFRICAN MUSEUM

192

cer. GES 86 LOI LZ 6 bl OL8I ¥9hh «O01! §=OLZI = 89 8081 86 SII €8€ IZE =N
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650

6r'0

se‘0

VILNAGOu

]e10}-qns

I a ee
780 610 — = ae = = = Z0‘0 £0°0 = L7‘0 90°0 a a ae — * * * $s4aqiaayos *W
a aa = =z =~ = = S00 +00 700 — L7‘0 = = = = ae ae sisuadvo “q
== ase = = — _ — —_— — —_ —_ — 90‘0 = = = =a © * §njojuajjoy “7
= 6D a ‘an ae a a 0 6 STO ST 0ST = L1‘0 == = 970 T€‘0 40109141 “W
= Ex = — <s ei = ae = z0‘0 ar = ox 25 a me oa janansa] ‘We
ie a ae a pa? - eet IIo §=6: 600 s—s«: 900-800 sre 90°0 = = 970 sal sisuaddd “Y
zv‘0——s«éST'D aa = aad a rs 910 «©6910 = LOO S800 — 90°0 a oe a a * snsoatjo “Y
Vaud LdOUlHo
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VuUOALLOASNI
HOT HOO s10a +108 £108 710M I108 TOM C410 @d1IO IdIO dIO ?tdh# ftdh# tdh Idd ‘IOA

‘TI d oienbs 3s9} y sevjdwioog 94} Ur Ss[oAo] JOMOT WOIJ so[dutes ur so1oeds Jo UOTeJUESaIdaI o8RjUIIIOd

[ alav

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS

leaf mould

partly disturbed

marine shell

concentration

compoct ashy loyer

tortoise shell
concentration

ay —

— ean i ai is a ei ee

bedrock

O 0,5 m

193

black ash

ash, brown soil
ash

ash, brown soil

Fig. 4 Section 028/29 at Byneskranskop | (after Schweitzer & Wilson 1978).

194 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 2

Percentage representation of species in samples from upper levels in main excavation at Boomplaas A,

DGLI DGL2 BLD2 BLD2A BLD3 BLD3A_ BL BLA BRL BRL2

INSECTIVORA
C. duthiae as Cae ee 0,42 1,08 — 0,78 0,71 1,06 0,16 0,48 0
E. edwardi sce Feet. ote ELS 1,26 1,08 0,99 0,78 0,71 1,42 0,99 2:40 6a
M. varius a. ets) ae tee ieee ee O. 8,82 4,30 3,47 6,07 4,48 3,90 6,73 5,29 4,05
?Myosorex . . «© + «© «© « «+ 9,59 — — 0,50 0,47 0,47 0,35 0,16 — 1,62
Se PART ee ee wee a OS FOES 0,42 — — — 1,18 0,71 1,64 1,92 0,81
GlCvpaneds are a. es ee ALS 4,20 3,23 1,98 2,18 2,36 4,61 1,64 1,92 2,02
Go avescens 2 si. rok, Ve me nel E24 13,87 9,68 14,85 12,15 11,56 13,12 11,00 10,58 7,69
sub-total) % 7? jo 2s Peedi ee 5 9 2485 28,99 19,35 21,78 22,43 21,46 25,18 22,33 22,60 20,24
CHIROPTERA
R. clivosus — — 1,08 — 0,16 0,24 — —_ 0,48 —_
R. capensis —_ — — —_— 0,31 — — 0,16 _ —
?M., lesueuri . — — —_— — — —_— — — _ —
IMEC PIC OlOb is Se Bie tole baa ee — — — _ — — 0,16 _— 0,40
E. hottentotus _ —_— — — 0,16 — —_— — — =
E. capensis _— _ —_ _— _— _— — — — —
M. schreibersi — — —_ —_— — — — — — —_—
sub-total — — 1,08 — 0,62 0,24 — 0,33 0,48 0,40
RODENTIA
C.hottentotus . . ... ..1 1,24 11,76 17,20 24,26 18,69 16,27 12,41 12,97 6,73 9,31
A. SUDSPINOSUS. =. Cs Sw, «SSD 0,84 — — 0,16 0,47 2,13 0,66 0,48 2,43
A; namaquensiS. «95 = « & ws « M24 10,50 9,68 7,43 7,01 8,02 7,09 9,03 14,42 18,22
ID INCONMUS es" Geo ie Leh he ts — — — 0,50 0,31 0,24 — 0,16 — 0,40
Miminutoides.” «0 ¢ «2. 3 sw we CU COS 0,42 — 0,50 0,47 0,94 0,35 0,99 0,48 0,81
P. verreauxi SR tee eo ok ee oe LS — 1,08 2,48 0,62 0,47 1,06 0,82 3,37 3,24
Ra pUumilio® oS és eee cs 8 P28 2,94 — 2,48 2,34 4,01 2,48 0,99 0,96 1,20
M. albicaudatus: . « SF os sl Ce) «GS 5,04 6,45 4,95 5,76 6,60 8,51 6,40 SAT) 3,64
DiGfea ares UR Re. wulget ee — — — 0,16 0,47 0,71 0,49 — 0,40
G. paeba . ee 5” Ph een ec — — — 0,16 0,24 — 0,16 0,48 —
D.imelanotis 3 6% Re Se es 0,42 — — 0,16 0,24 0,35 0,33 0,48 0,40
D. mesomelas re te emo ean 26 — — — — — —_— —_ — — -_
2Dendromus 3-5 5 2 «= « «1 « 4 0359 0,42 1,08 0,50 0,16 — 0,35 0,33 — =
SSAReDStL ot as. ba es “Ue, Pee eee 0,42 3,23 0,50 1,25 2,36 3,19 1,64 2,40 1,62
Sucampestris:s, 2 3 6 4 ue OS ES 1,68 1,08 0,99 1,40 1,42 1,77 0,49 0,96 —
Olaminatus 96 Viens. se ee ey LS 0,84 1,08 — 0,31 0,47 0,71 0,49 0,48 0,81
O. saundersae Bees’ Ah, beh Sheet LALO 5,88 10,75 6,93 8,10 7,55 6,74 8,21 6,25 8,50
QO irroratus, 2% e-em ee es he SAO 2, 2 228199 26,88 25,74 27,26 26,89 25,89 31,36 32,69 25,10
Oo untsuleatus™ %- oe «he we ALES 0,84 1,08 0,99 2,49 1,65 1,06 1,81 0,96 3,24
Goculanis.| 2145. 3 ty = = ae — — — 0,16 — — — — —
sub-total so «. gee, chic, = TOS 71,01 79,57 78,22 76,95 78,30 74,82 77,34 76,92 79,35
ING =e a aire, Mee et ok ee 169 238 93 202 642 424 282 609 208 247

at about 24 000 B.P., and then by the Robberg industry at about 20 000 B.P.

Thereafter, at the end of the Pleistocene or the beginning of the Holocene,

about 11 000 B.P., the Albany industry succeeded the Robberg industry and

was itself replaced by the Wilton industry by about 6 500 B.P. at the latest.

Whilst it is presumable that the makers of all of these industries were hunter—

gatherers, domestic stock appears to have been introduced into the site about

1 700 B.P. This phenomenon was accompanied by a significant reduction in the

number of stone tools recovered (Deacon et al. 1978: 47).

The samples of microfauna from much of the lower half of the sequence |

are large even though they were available only from the test square metre

(Table 1). The sample from the upper half is also good now that it has been

augmented by material from the extended excavation (Tables 2-3). This latter

sample comprised a considerable number of subsamples based on lithological

units observed in the excavation.
|
:

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 195

t

BRL CL3 GWA LP LP LPC
4UA BRL4 BRLS5 BRL6 BRIT CLI CL2 BG TBF GWA GGU GGL LPM 2B

nes t0l 860,52 «=s(«i,72—é—é‘zi1«94«=Cfsi‘i3,60st—ié‘“SS;23—(i‘8sC3,13i3,27 Ss ,5 a9 2
am) 3,70 = 2,03 1,04 1,44 = 072 623 O14 ce ae ag matt o
Meee 3,91 «86810 648 6,22 5,81 13,67 15,00 26,17 25,28 30,39 22,99 2211 39,61
Bis io ao iv fe th fh 38 1B te aR ER eS
— , VJ) > ’ > ’ ’ ’ 1, 9 2,61 0, , 3
Sees 2,78 «= 2,330 3,11 2,58 22,16 ~~ :1,36 pa Rae pane Berd hone
eee 405 «7,51 «39 5,74 0S «7,53 «10,79 «= «44,32,S's«i12,79''s«*d4,28—Ssi«i,2si2,30s«i53 «1,95

16,05 «19,78 = 19,24 19,95 18,90 19,14 34,53 27,50 36,50 = 39,33 42,81 32,76 35,05 53;z5

me 022 «#2025 «026 — xy es Le ode Ge 043 “oe” sean ee
“a a i Eat Abe ee as ee Ay ak Wwe a
a =, = os 3 we p72. 023: — GiGi 014. OAds 2
. ‘ea = oe = as =o sas = oe is 0,1 ee a
et - 02 — oa ae ae = = ue ra) Olin ee
oa = es se. en, = a Z = ri a Shiels. Osi og a8
ue o2s)««(Cw78—i24 CC SCti‘i«iSTst«‘i SSSC«iC SAC‘ Citi STi —
$64 11,96 11,39 14,51 16,51 20,43 10,79 22,73 7,30 4,25 3,92 5,60 4,00 1,95
m0 842076 | 120 022 07 as 26 9. OO (fea 2.628 - OC2l.amOes
675 1761 15,44 1088 789 1075 1007 614 179 161 261 172 «1,79 1,95
123 022 025 026 048 0; a on pia. One ieee ode. Baie ec
go? (051 #42026 O48 022 — =) gm” 09. 4 ou 629 | O53 ~ 2065
Moe 495 «253 «250 sisd1'91's«i43—«‘i “Citi 4sCGSiéi838—i29—i«iS3——«é GS
123 O87 127 «291781 «2120 086 1:44 182 0355 0732 0.65 0:43 0,84 1,95
ase 304 «#2337 «6311 426194 6« 144 iia'14i(iHeiCi 029 0.42 0,65
aa ey rr ‘Z 8 <a is val as 5 5
oa me 52s «200206 Cl ze al = a = ee a we
mer o4s «(101 «s«26 «024 Si43si«é«72—«“‘ t«‘istié‘“iSSt«it(itiéiiwsdSC«*dYSG—~C*«*«*3
no -— o72 024 — cane fos Mes es ig | guide Suis ee
oo 025 — 04 — re = eas” 080-033" 629 «(0,53 °° 495
— 087 02 — iat = ae ast is oe co = rae ues
ier 0; 155 048 108 072 136 O14 016 — O14 — a
TAL 6,97 1494 11,40 933 13,76 17,27 14,09 39,12 39,57 30,72 «41,38 «39,89 24,68
29563 28,48 + 25°32 «3083 «36,60 «2903 20;14.S «2159 7702,S'—:«(C«B~—«12-42~=«s10,20S-=«9,89—S« 49
Meso 177 «#20.260«C««i24 08 CC ti<C~i sii «4,32 3,90
meas 625 — 024 022 072 023 O14 O27 - O29 O44 —

82,72 79,78 80,51 79,27 80,86 80,86 64,75 71,82 63,36 60,35 56,86 66,67 64,42 46,75
8 460 395 386 418 465 139 440 726 1246 306 696 950 154

Byneskranskop 1 (BNK1)

Byneskranskop 1 (34°35’S 19°28’E) lies some 9,5 km south-east of Die
Kelders. It is situated 60 m above sea-level in the side of a small limestone
hill near the Uilenkraal River (Figs 1-2). There are approximately 3m of
deposits spanning the last 12 500 years without major breaks. The stratigraphy
(Fig 4) comprises mainly dark-grey soil interspersed with ash bands and a
prominent band each of shell and tortoise bones (Schweitzer & Wilson 1978).
The cultural sequence is divided broadly into two industries, a ‘pre-Wilton’ in
levels 19 to 10, and a Wilton in levels 9 upwards (Schweitzer & Wilson 1978).
Further subdivisions of these major divisions into a total of five subgroups are
possible but have not yet been refined. Pottery was recovered from the
highest levels.

The microfaunal samples from the pre-pottery levels is adequate (Table 4);
that from the pottery levels was too small to warrant examination.

196 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 3

Percentage representation of species in excavation units in the main excavation at Boomplaas A.

DGL BLD3 BL BLA BRL CL GWA LP LPC

INSECTIVORA
C. duthiae 0,38 0,75 1,06 0,16 0,62 2,74 3,04 3,13 3,25
E. edwardi 2 ris 0,75 1,42 0,99 2,59 0,55 0,05 0,0 —
M. varius a Eee Debt 5,44 3,90 6,73 5,63 9,44 25,61 23,72 39,61
?M yosorex : 0,26 0,47 0,35 0,16 0,28 0,34 6,49 4,87 7,14
SVarlas:.. <a. . 0,26 0,47 0,71 1,64 1,24 1,30 1,62 1,28 1,30
C. cyanea 6 3,83 225 4,61 1,64 2,48 2,33 — — ==
C. flavescens. . . 13,54 11,91 13,12 11,00 0 5,68 1,47 2,41 1,95

sub-total. ........ 24,52 22,05 25,18 22,33 19,92 23,05 38,29 35,45 53,25
CHIROPTERA
ROCHVOSHS Sc. 0,13 0,19 — — 0,28 — 0,15 0,20 —
R. capensis ; — 0,19 — ,16 — All — 0,05 —
M. lesueuri = — — — — — — —
M. tricolor ‘ — — — ,16 0,11 ,14 0,10 0,10 —
E.hottentotus . . — 0,09 — — — — — 0,05 —
E. capensis — — — — ll — — 0,05 —
M. schreibersi — — — — — — — 0,05 —

sub-total... . O13 0,47 — 0,33 0,51 0,34 0,25 0,51 —_
RODENTIA
C.hottentotus . . 16,35 eds 12,41 12,97 11,25 19,08 5,38 4,56 1,95
A. subspinosus . . 0,51 0,28 2,13 0,66 0,90 0,62 0,25 0,20 0,65
A. namaquensis . . 9,32 7,41 7,09 9,03 15,48 8,48 1,67 1,90 1,95
Doinecomtus: = =. « 0,26 0,28 — 0,16 0,28 0,27 0,10 0,10 —
M. minutoides . . 0,38 0,66 0,35 0,99 0,39 0,21 0,30 0,36 0,65
BP. verreauxt 5 3 GS) 0,56 1,06 0,82 3,32 1,03 0,15 0,41 0,65
RiecpUTIHIO =a re) Px ATI 3,00 2,48 0,99 1,24 1,30 0,41 0,67 1,95
M. albicaudatus . . 4,09 6,10 8,51 6,40 3,71 1,98 0,25 0,31 0,65
Et OftGe eo a — 0,28 0,71 0,49 0,17 0,07 — — —_—
Gi paeba Sas. — 0,19 — 0,16 0,17 0,07 — — —_—
D.melanotis. . . 0,13 0,19 0,35 0,33 0,51 0,48 0,91 1,33 1,30
D.mesomelas . . — — — — 0,23 0,14 — 0,05 —
?Dendromus . 0,51 0,09 0,35 0,33 0,11 0,07 0,81 0,41 195
S. krebsi . ‘ae 0,77 1,69 3,19 1,64 0,79 — — — —_
S. campestris > pe 1,28 1,41 1,77 0,49 0,11 — — — —
O. laminatus . : 0,64 0,38 0,71 0,49 0,96 0,96 0,15 0,05 —
O. saundersae , 6,77 7,88 6,7 8,21 9,85 12,93 39,40 38,99 24,68
QO: irroratus = 3s 3 «29576 27,11 25,89 31,36 28,36 28,11 6,49 10,40 6,49
O.unisuleatus . . ETS 2,16 1,06 1,81 1,58 0,48 4,97 4,10 3,90
G.ocilaris? «59 %- 0,13 0,09 — — 0,17 0,27 0,20 0,20 —

sub-total’ “7 2. ., " 75;35 77,49 74,82 77,34 79,57 76,61+ 61,46 64,04 46,75
Ne 783 1066 282 609 1777 1462 1972 1952 154

+ Includes Muridae indet. 0,07 %.
BLD3 = BLD3 and BLD3A
BRL = BRL — BRL6

CL = BRL7 — CL3BG

GWA = GWATBE and GWA
LP = LPGGU — LPM

Die Kelders 1 (DK1)

Die Kelders 1 (34°32'S 19°22’E) is situated on the Walker Bay coast some
5 km north-west of Gansbaai (Figs 1-2). The cave is presently 8 m above
sea-level. There are more than 7 m of deposits but approximately 1 m of these
represents a period of non-occupation between the Middle Stone Age and the
Late Stone Age. Tankard & Schweitzer (1974, 1976) have suggested that the
former occupation lasted from about 80 000 B.P. to about 35 000 B.P. There
are two published dates for the M.S.A. (Schweitzer 1970: 136) which were
obtained using the apatite fraction of bone; one of these seems unlikely and the
other is inconclusive. The former, GX—1716, from level 3, is 11 200 + 700 B.P.
which by comparison with other sites is far too young. The latter, GX-1717,

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS iy
from level 5, is 31 800 + ae B.P., which could fit the circumstantial dating
given by Tankard (1976a: 156). There is a series of dates from the L.S.A.
occupation, ranging from 2 020 + 95 B.P. (GX-1686) to 1456 + 100 B.P.
(GX-1685). Tankard & Schweitzer (1974, 1976) discuss the geology of the site
and Figure 5 is based on their sections. Accumulated upon quartzite boulders
and sand are six M.S.A. occupation levels interleaved with layers of sand. Near
the top of the M.S.A. sequence a massive rock-fall was probably caused by an
earthquake (Tankard & Schweitzer 1976: 310). Thereafter, at the height of the
Last Glacial, the cave was uninhabitable, first because of standing water and
then because of blockage of the entrance by a sand-dune. Only at about 2 000
B.P. was the cave re-occupied, this time by Wilton people who subsisted largely
on shellfish and other marine resources.

Although adequate microfaunal samples were obtained from most of the
M.S.A. levels, there is a tendency for larger numbers of individuals to have
been accumulated at times when the cave was thought not to have been
occupied by people (Tankard & Schweitzer 1974: 367) (see Table 5). This is
similar to a tendency for rodent bones to be present only when human
occupation was not intense, which was noted at Redcliff Cave in Zimbabwe by
Brain (1960: 132).

Klasies River Mouth 1A (KRMI1A)

Klasies River Mouth 1A (34°07'S 24°24’E) is one of series of seven caves
containing archaeological material which is situated within a distance of about
2 km along the coast some 40 km west of Cape St Francis (Fig. 1). KRMIA is
presently 6-8 m above sea-level (Wymer & Singer 1972: 207). According to
Butzer (1978), levels 33 to 22 comprise alternating well-stratified hearth zones
and light grey silty sand. Levels 21 to 10 comprise silty sands with some coarse
sands, while levels 9 to 1 again comprise silty sands but with a greater
abundance of artefactual debris. KRM1A contains both M.S.A. and L.S.A.
levels, of which the former date from perhaps 125 000 B.P. to about 65 000
B.P. (Bada & Deems 1975), while the latter were deposited during the last
5 000 years (Klein 1976). The M.S.A. has been divided into five industrial
units, M.S.A. I and II, Howieson’s Poort, and M.S.A. III and IV, of which the
first is the oldest. The L.S.A. has been divided into I and I with the latter
being the younger.

One of the two numerically adequate micromammalian samples from this
site comes from level 15 which is a Howieson’s Poort level; the other comes
from level 32 which is an M.S.A. II level (Table 6).

Nelson Bay Cave (NBC)

Nelson Bay Cave (34°06’S 23°22’E) is located on the west face of the
Robberg peninsula just west of Plettenberg Bay at about 19-21 m above
sea-level (Fig. 1). The deposits reach a depth of about 5m and have been
discussed in detail by Butzer (1973) and the section given here (Fig. 6) is based

198

TABLE 4

ANNALS OF THE SOUTH AFRICAN MUSEUM

Percentage representation of species in samples from Byneskranskop 1.

INSECTIVORA

C. asiatica .
E. edwardi .
M. varius
S. varilla
C. cyanea
C. flavescens

sub-total

CHIROPTERA

R. clivosus .
R. capensis .
M. tricolor .
E. hottentotus .
E. capensis .
M. schreibersi .

sub-total

RODENTIA

C. hottentotus .
G. capensis .

A. subspinosus .
A. namaquensis
D. incomtus

M. minutoides
P. verreauxi

R. pumilio .
M. albicaudatus
T. afra .

D. melanotis
D.mesomelas .
?Dendromus

S. krebsi

O. laminatus
O. saundersae .
O. irroratus

sub-total

N =

1

2

3

20,00
20,00

4

1,67
16,67

35,00

0,26

20,41
0,90
12,07
aye 9

61,87
179

0,27
0,55
2,47
1,65
3,30
1,65
0,82
0,82

19,51
0,82
12,91
5,49

55.75
364

58,78
131

on this work. The lower half of the sequence consists mainly of iron humate-
stained loams while the upper half comprises mainly shell material in a sandy
loam matrix. M.S.A. material was found in the lower third of the deposits
below a rubble horizon which Klein (1972a: 184; 1974a: 272) considered at one
time to have been accumulated during a period of about 10 000-15 000 years
when the cave was not occupied. The existence of comparable industries at
Nelson Bay Cave and at Klasies River Mouth suggests, however, that the
Nelson Bay Cave M.S.A. occupation might have lasted from about 120 000
B.P. to perhaps 90000 B.P. Above the rubble horizon, the late Upper

=~ ———————————————

\IICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 199

9B 10 i 12 13 14 15 16 17 18 19

3,90 2,56 351 3,90 1,14 1,75 4,02 1,39 3,85 2,70 paYIs

24,68 12,18 13,10 AWD 9,09 17,28 71 9,72 14,42 8,11 15,24
6,49 at 4,76 3519 Lyd 2,92 oe | 4,17 2,88 2,70 111

6,49 962 10,71 7,79 13,64 20,47 97 = A250 10558 5,41 13,85
esceeesols 32,14 2468 26,14 37,43 33,33 27,78 31,73 18,92 32,41

= 0,64 = = re G57 Vi 30.) = a 0,83
a On. 119 += = = =e Hs o = 0,28
_ = -_ ma an = a a 0,28
os as Be MAG hy As 1 = = = 0,28
= = = _ at SGOr = Jes ss 0,28
- a Oe.) Hor ae = 1,94
1,3 1,28 i, = A 0,28
io 7,05 833 10,39 5,68 —$,85 5,56. 865 8,11... -6,65
O5e- 116 > 30 | 192 1,39

= = = = as as oa G96 pee ec
= af as = = = = ee a = 0,28
hao 1.92 Poe. iid. O68 is 0.0600 ° = 0,83
fo. 1,28 + Foo. 14 O38. O57 1,39, 21,92 ~~ 2:70. - 194
Pets «60038 06Ci‘iK ie 230 278-385 2700 *, as0s
memes 5.05 (3903.41 i351 iia — ar ki 222
meoeeeo S056 «(C1969 )0S12,50 12,28 «= «aG32s—i33—i‘«wS 3B 6,65
a wae 130 «#414 «26058 «#8057 — 096 — 0,28
ion, bi fia: (O58 -.0,57 .° 1239. 0.96 Se 0,28

— £92 239 1,30 —
Zest. «t/5! 11,90 ive. 4,55 co) Tae FE BP 4,17 3,85 2,70 5,26
2,60 — 2,38 2,60 yA | LA7 2,30 2,78 0,96 2,70 1,39
mieoeetst «696 96,67 15,58 25,00 17,54- 19,54 18,06 17,31 24,32 19,94
5,19 8,97 Bo3 4612:99 8 8=6135,91 ere . 2399 222522. A923- 27,038.) 15,24

58,44 68,59 66,67 74,03 73,87 61,99 66,09 69,44 68,27 81,08 65,65
77 156 84 77 88 171 174 12 104 37 361

Pleistocene Robberg industry underlies shell-midden deposits associated with
the Holocene Albany and Wilton industries.

Two Robberg levels, YSL and YGL, produced good microfaunal samples
(Table 6). These were accumulated between about 18 000 B.P. and 12 000 B.P.
(Klein 1972a: 203). As such they are interesting because no other samples of
this age have yet been found on the coast. On the other hand, their usefulness
for palaeoenvironmental interpretation is reduced by the fact that they do not
form part of a sequence and cannot, therefore, provide evidence of changing
conditions over a period of time.

200 ANNALS OF THE SOUTH AFRICAN MUSEUM

MODERN SITES

Where possible, comparative modern samples were collected from close to
the archaeological sites. The Cango valley provided a wealth of material from
two main sites and one minor locality, namely Boomplaas, Nooitgedacht, and
Osgat respectively. A sample was obtained from Byneskranskop but, apart
from this , the nearest collections to Die Kelders 1 were made at Stanford. This

LSA shell midden
pink shelly sand

sond deposited _ into

standing water

|
collapsed roof rock, loam

3

4
Im 5 loom

6

Ti

8

9

10 e@boulis secs

1/12 loam, eboulis secs

13
0 14

1-14 MSA levels

Fig. 5. Stratigraphy of Die Kelders 1 (after Tankard & Schweitzer 1974).

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ANNALS OF THE SOUTH AFRICAN MUSEUM

latter material was obtained by other workers and was not examined personally
by the writer. The most easterly collection was made at Glentyre near Wilder-
ness. In this case the aim was to sample a different environment, namely
Knysna forest bordering lakes and coastal dunes, rather than to collect close to
an archaeological site.

Again the sites are described below in alphabetical order. Some indication
is given of the vegetational setting as well as of the samples collected.

Boomplaas B and C (BPB and BPC)

Boomplaas B and C (33°23’S 22°11'E) are adjacent rock shelters set in the
same cliff as, and within 100m of, Boomplaas A which is described above

TABLE 6
Percentage representation of species in samples from Klasies River Mouth 1A and Nelson
Bay Cave.
KRMIA 15 KRMIA 32 NBC YSL NBC YGL

INSECTIVORA
C. dathiae % se. % — — 0,28 0,26
A.hottentotus . . . 0,95 0,93 — —-
NE. varias 0 eS! 2s: 2,86 0,93 1,87 1535
iS: MORI Sh eS — — 0,52 0-13
C. eyanea 4 6." — _ 0,03 —
C. flavescens. = =. % 11,43 5,61 3,92 3,62

sub-total 3; 2... . 15,24 7,48 6,63 5,56
CHIROPTERA
Fe. CLEVOSES oo On oe ad 0,95 —- 0,03 —
KR. COpensi§s <5 0,95 — = 0,13
CEDICSICUS’ 2 gs — — — 0,13
M. schreibersi . . . — — — 0,13

Sub=total’*2- 2 ..% 1,90 — 0,03 0,39
RODENTIA
C.hottentotus . . . 2,86 2,80 1,63 3,10
G: CQDCEMSIS 5 5. x 4,76 4,67 1,53 1,68
D..tieomtus 3. ss 2,86 — 0,07 0,26
M.minutoides . . . = — 0,03 —-
P. VCFFEQUAE =. yn ks 4,76 = 0,21 O5a2
Ri PUTANIO | ok 5.11 2,80 0,59 0,91
M.albicaudatus. . . — — -9,40 8,15
TGHEGS | ee is, Ss Me Pe — — 25,70 22
D..melanotis.. . . . 0,95 — 0,31 0,26
O.laminatus. . . . 7,62 11,21 0,35 1,16
O.saundersae . . . 21,90 13,08 50,12 50,19
O: Hrovatus . = x. s 31,43 57,01 3,09 3,49
O.unisulcatus . . . — — 0,31 1,81
G.iOCHIGHIS: (3 es — 0,93 — ==

sub-total 2. 8.* «0 & 82,86 92.52 93,34 94,05

N= va 2 eee. Bae 105 107 2883 773

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS

B/|

BSBB/|
H/G
BSBH/G

Ra

Rb

GSL

BSL
YSL
YGGL

Grey Loam
Rubble Horizon

Black Loam III

Black Loam Il

Black Loom |

Pale Brown Loam

Basol Loam

Fig. 6. Stratigraphy of Nelson Bay Cave (after Butzer 1973).

y micromammalian samples

203

204 ANNALS OF THE SOUTH AFRICAN MUSEUM

(Fig. 2). They are about 50 m above the valley floor. The surrounding vegeta-
tion is Limestone Vegetation with Mixed Bush (Moffett & Deacon 1977: 130),
with some large trees near the Grobbelaars River and cultivation on the valley
floor. Only the loose surface-material of these shelters was examined; it is
consequently not possible to tell what depth of deposit there might have been
under the surface. The surface material largely comprised Procavia capensis
(rock hyrax or dassie) dung, supplemented to a certain extent by that of Papio
ursinus (chacma baboon). Owl pellets were scattered throughout the shelters,
although there was a tendency in Boomplaas B towards a concentration nearer
what was presumably the roost. There was also a trail of pellets on various
ledges below what appeared to be the roost, judging by the quantity of
droppings found there. In Boomplaas C, perhaps because the area was more
exposed, there seemed to be no consistent roosting-spot, although half the first
collection was made from a ledge some 5 m above the main floor. The owl
which frequents these shelters has been identified by G. Avery (1976 pers.
comm.) as Tyto alba (barn owl) on the basis of shed feathers collected in the
shelters. Both 7. alba and Bubo africanus (spotted eagle owl) have been seen
in the valley (H. J. Deacon 1976 pers. comm.). Other birds such as Onychogna-
thus morio (red-winged starling) and Columba guinea (Cape rock-pigeon)
inhabit the rocks around the shelters.

Owl pellets were collected by the writer from these shelters in January and
July 1976 and 1977, the site being cleared on each occasion. Other collections
have, however, been made since 1974 and some pellets have also been collected
from Boomplaas A (H. J. Deacon 1976 pers. comm.). From these a sample of
50 pellets was analysed by V. A. Scott, together with a sample from Boomplaas
A level OLP2, in a student project that represented an early attempt to assess
the use of micromammalian remains in palaeoenvironmental studies. These
samples were subsequently re-examined by the writer and the results incorpor-
ated in the present study. A bulk sample was also collected from the floor of
each of the two shelters. Table 7 lists the total from these rock shelters.

Byneskranskop 2 (BNK2)

Byneskranskop 2 (34°35'S 19°28’E) is a rock shelter situated just below
and in the same cliff as Byneskranskop 1, which is described above (Fig. 2). In
the vicinity of the cave, both on the hillside and along the nearby Uilenkraal
River, coastal scrub (Taylor 1978: 213) with Sideroxylon inerme (milkwood)
occurs. Further patches of similar scrub also occur some 4 km north-west of the
site (Schweitzer & Wilson 1978). Vegetation along the river is dense and
includes Phragmites reeds, rushes, and alien Acacia species. The vegetation of
the sandstone hills if predominantly restioid, whereas that on the limestone
ridges is mainly proteoid (Schweitzer & Wilson 1978). Considerable areas
below the rock shelter and, particularly, on the coastal plain, have been farmed
and are either under cultivation or lying fallow. Elsewhere expanses of alien
Acacia, coastal fynbos and some grassland contribute to the mosaic.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS

INSECTIVORA

C. asiatica

C. duthiae

A. hottentotus
E. edwardi

M. varius
?Myosorex

S. varilla .

C. cyanea

C. flavescens .
Crocidura sp .

sub-total

CHIROPTERA

N. thebaica

R. capensis

M. tricolor

E. hottentotus

E. capensis

M. schreibersi
Chiroptera indet.

~ sub-total

RODENTIA
C. hottentotus
G. capensis
A. subspinosus

A. namaquensis .

D.incomtus .
M. minutoides
P. natalensis .
P. verreauxi .
R. rattus .
R. pumilio

M. albicaudatus .

T. afra

D. melanotis .
D. mesomelas
?Dendromus .
S. krebsi .

S. campestris
O. laminatus .
O. saundersae
O. irroratus .
G. ocularis

sub-total

i oe

Percentage representation of species in modern samples.

BPB-C

TABLE 7

BNK2

205

GLEN NGA-B
2,19 0,98
0,73 e

Biss 0,25
7,83 24.02
4,01 1,96
oh: 2.21
1,46 0,49
a 0,12
16,21 30,02
0,36 0.12
0,55 0,12
0.18 ue
0,18 ey
1,28 0,61
at 0,61
5,10 a
oi 5,27
ide 13,36
we 0,25
9,84 4.29
ae 5,51
0.73 8.46
0,73 ce
3,28 2,33
is 0,12
1,82 135
6,38 1,59
as 135
a 0,61
0,91 0,86
0.36 4.04
53,37 19.00
- 0,37
82,51 69,36
549 816

206 ANNALS OF THE SOUTH AFRICAN MUSEUM

There were no fresh pellets in the shelter, only a concentration of micromam-
malian bones which was particularly noticeable on the northern side of the shelter.
A bulk sample (Table 7) was collected from this area, but it was not possible to
examine the shelter in any detail as it was occupied by a swarm of bees.

Glentyre Shelter (GT)

Glentrye Shelter (33°59’S 22°38’E) is located about 100 m off the road
which skirts the north-east corner of Onder-Langvlei and approximately 6 km
east of Wilderness (Fig. 1). The rock shelter, which is an archaeological site
excavated by A. J. H. Goodwin in 1938-40, published by Fagan (1960) and
mentioned by Schrire (1962), is situated about 100 m above lake-level on the
side of the ridge bounding the string of lakes to the north. The shelter is
situated in Knysna forest, while the lower ground is cultivated and the lake is
surrounded by Phragmites reeds. The owls (Tyto alba) appear to roost on a
ledge north-east of, and above, the shelter. The result is that, when not caught
in vegetation, the pellets roll and are washed down the slope into the nearer
half of the shelter. There were a great number of micromammalian bones both
here and on the talus slope below the shelter (Table 7). This fact, in itself,
throws some interesting light on the mechanics of accumulation of these bones
in archaeological sites.

Nooitgedacht A-—C (NGA-C)

Nooitgedacht A—C (33°22’S 22°10'E) are adjacent caves and shelters
situated 2,5 km north-west of Boomplaas about 100 m above the valley floor
(Fig. 2) in a Limestone Vegetation area. Nooitgedacht A is by far the largest of
the caves and seems to be the main roost since most of the material has come
from there. It consists of a main cave with an elevated tunnel or fissure
extending back at least 20 m from the northern corner of the cave. On the
second visit, pellets were collected from this tunnel so that it is clear that the
owls roosted far up it at least part of the time. Nooitgedacht B and C are very
small shelters or overhangs. As at Boomplaas, the main surface material
comprises Procavia capensis dung together with some faeces of Papio ursinus.
Onychognathus morio and Columba guinea inhabit nearby rocks as they do at
Boomplaas. The bodies of three Tyto alba have been found in the cave.

In January and July 1977 collections were made at Nooitgedacht where the
main cave, A, yielded a great deal of material. Caves B and C yielded a few
pellets each and were presumably used only as occasional alternative roosts to
Nooitgedacht A. Nooitgedacht B did, however, provide a useful bulk sample.
The total for caves A and B is given in Table 7.

Osgat (OG)

Osgat (33°22’S 22°11’E) lies 1,3 km north-east of Nooitgedacht and 5 km
north of Boomplaas, only just above the valley-floor (Fig. 2). It is a small
shelter with a cleft in the rock parallel to the back wall in which the pellets were

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS ZO

found. The shelter was obviously well used; not only were there Procavia
capensis and Papio ursinus droppings but also those of domestic cattle. Bats
were found roosting in the cleft and it was interesting that the owl pellets
contained an unusually high proportion of bat remains. These remains were
identified as belonging to Nycteris thebaica (Egyptian slit-faced bat) and,
although it was not possible to identify the live specimens, it is likely that they
were the same species. Osgat yielded only a few pellets in January 1976, and
since the site was obviously deserted it was not visited again.

Stanford (ST)

Stanford village (34°26'S 19°28’E) is situated about 15 km south-east of
Hermanus beyond the head of the Hermanus lagoon. The material which was
published by Grindley et al. (1973), but is no longer available (C. J. Vernon
1977 pers. comm.) was recorded as having been collected from ‘a hole 3 m
above the river in the bank of the Klein River, about 1,5 km west of Stanford’,
that is, towards the lagoon (Fig. 1). “The surrounding habitat was a fringe of
Phragmites and macchia along the river encompassed by agricultural lands,
rooikrans thickets and gum plantations’ (Grindley et al. 1973: 266). (It should
be noted that ‘macchia’ has been called coastal fynbos in the present study on
the basis of current nomenclature. )

_ The second collection in the area was made at Windheuvel (34°28’S
19°32'E) which is about 8 km east-south-east of Stanford (Fig. 1). The roost
was in a hole in a limestone cliff along a tributary of the Modder River, which
itself drains into the Klein River. Unfortunately an attempt to relocate the site
at a later date failed and it was thought that the cliff must have partly collapsed.
The material from this collection is being held privately by T. N. Pocock of
Vanderbijl Park, who provided the writer with a copy of his analysis.

THE MATERIAL

The material consists of large quantities of bones of rodents and insecti-
vores, together with a small number of bat remains. The identity of the agent
responsible for their collection is first considered. Thereafter, the methods used
to collect the samples and questions of identification are discussed. Finally, the
relationship of the excavated sample to the original living community from
which it was derived, as well as the type of interpretations it is possible to make
therefrom, is examined.

AGENCY OF ACCUMULATION

It has generally been assumed (Cartmill 1967: 171) that owls are respon-
sible for the accumulation of quantities of microfaunal remains in caves. While
this does, indeed, seem to be the most likely explanation, it is as well to
consider the question in rather more detail in order to ascertain whether or not
other small predators might have been responsible. Given the large numbers of

208 ANNALS OF THE SOUTH AFRICAN MUSEUM

small animals and diversity of species involved, it seems reasonable to accept
that the majority of the microfauna was accumulated in the cave by some
outside agency. A few animals such as bats might have been living in the caves
and have died there, but this cannot have been true for most of the animals
which would not live in or enter a cave voluntarily. For instance, limited
trapping at Boomplaas A produced only one species, Aethomys namaquensis,
in March 1979. Nor can it account for the quantities of animals involved. It is
then necessary to employ a process of elimination in order to isolate the agency
responsible. Firstly, only predators can be involved. Secondly, only predators
of a certain size would be likely to catch animals in the size-range represented,
that is, anything up to the size of a half-grown Georychus capensis (Cape
mole-rat). Small carnivores such as Herpestes pulverulentus (Cape grey mon-
goose) are a possibility, as are various birds such as owls and eagles. Humans
are also known to eat rodents. Shortridge (1934: 240) notes that in summer the
Okavango of Botswana catch and eat large quantities of Ofomys irroratus
(vlei-rat) and Dasymys incomtus (shaggy swamp-rat), as well as the much larger
Thryonomys sp. (cane-rat). Vesey-Fitzgerald (1964: 66) records the fact that
the Wanyika of Tanzania catch and eat Rhabdomys pumilio (striped field-
mouse) which is surprisingly small. Also small is Steatomys pratensis (fat
mouse) which Coetzee (1971: 2) records as being eaten throughout its range.
Lizards, frogs, tortoises and insects are also said to be eaten on occasion by
Bushmen in Botswana (Schapera 1930: 141; Service 1966: 101, quoted by
Yellen & Lee 1976: 42).

In the matter of eliminating the possibilities, various points may be
considered. For instance, the time of day at which prey and predator are active
must coincide. This line of argument may not seem very strong since two of the
main prey items are apparently active throughout the twenty-four hour cycle.
Davis (1973) found, however, that Otomys irroratus was primarily crepuscular
and concluded that this activity pattern is probably common to all Otomyinae.
Thomas & Schwann (1905a: 264) noted both nocturnal and diurnal activity in
Myosorex varius (forest-shrew) but Roberts (1951: 39) stated that the Soricidae
in general are nocturnal. The great majority of species represented in the
samples is nocturnal and must have been taken at night. This eliminates
humans who are not nocturnal and are unlikely to have developed snares
capable of catching huge numbers of animals. Likewise, the diurnal raptors
such as the eagles are ruled out on this ground as well as by the fact that they
digest the bones of their prey (R. A. C. Jensen 1977 pers. comm.). H.
pulverulentus is also mainly diurnal (Dorst & Dandelot 1970: 122). Of the
remaining possibilities, only animals which frequent caves and regularly excrete
the more or less complete bones of their prey in one cave can have been
responsible. By this token the various carnivores may be ruled out. Some, such
as Canis mesomelas (black-backed jackal), may tend to defaecate in restricted
places (Stuart 1976: 204) but not in caves. This species has been shown to have
a very varied diet (Stuart 1976: 201) which only sometimes includes moderate

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 209

proportions of rodents. Felis libyca (wild cat) and Genetta genetta (common
genet) eat large quantities of rodents (Stuart 1977: 240; Smithers 1971: 125,
165) but, again, they would defaecate in the open and, in the latter case at
least, the prey bones tend to be very broken up (Stuart 1977: 241). Possibilities
are now reduced to the nocturnal birds of prey which, in practice, means the
owls since only these eat small mammals in large quantities. Of these Tyto alba
(barn-owl), Bubo africanus (spotted eagle-owl) and perhaps Bubo capensis
(Cape eagle-owl) would be likely to roost in caves (McLachlan & Liversidge
1970: 246 ff). Bubo. capensis would, however, probably take larger prey; little is
known of the local race, B. capensis capensis, but it cannot be expected to take
appreciably smaller prey than the slightly larger B. capensis mackinderi, which
is known to eat animals the size of hares in Zimbabwe (Steyn & Tredgold 1977:
41). Although the diet of T. alba and B. africanus is similar, the pellets of the
latter typically contain highly fragmentary prey remains (Grindley et al. 1973:
266). This finally leaves Tyto alba as the most likely creature to have accumu-
lated microfaunal remains in caves. In addition, TJ. alba is known to hunt
occasionally on dull days (McLachlan & Liversidge 1970: 247) which could help
to account for the small proportion of diurnal species represented. A further
indication is that the range of prey species in modern collections of pellets, all
of which are thought to be from 7. alba, are similar to those in the archaeologi-
cal samples.

METHODS OF COLLECTION

This study was initiated after most of the archaeological material had been
excavated. For this reason the collection of microfauna has taken place under
differing circumstances. In some cases the excavators routinely kept the micro-
faunal remains which were extracted during the general sorting after the
material had been screened through fairly finely-meshed sieves. A 3 mm (9”)
mesh was used at Nelson Bay Cave, at Die Kelders 1, and at Byneskranskop 1.
This has proved adequate for the collection of microfauna. At Klasies River
Mouth 1A, 12 mm (3”) and 6 mm (4”) mesh sieves were used for the most part
and only selected samples were screened through a 1,5 mm sieve for the
recovery of small finds, including microfauna. The result is that the sample
from this potentially very rich site is extremely small. At Boomplaas A, the
material from the test pit, square P12, was screened through a 3 mm meshed
sieve and the microfauna was extracted during the general sorting, except in the
case of levels OLP2 and OLP3. Here the sample was so large that it remained
unsorted until the writer examined it. During the main excavation at Boom-
plaas A, a system of sampling for microfauna and other small finds was
employed in order to speed up the process of sorting. Material from one sample
Square in each line of squares was screened through a 2 mm meshed sieve as
opposed to the usual 3 mm meshed sieve. In the upper levels, the microfauna
was sorted from all the squares although obviously more was retained by the
finer screening. Thereafter it was decided that a sufficiently large sample could

210 ANNALS OF THE SOUTH AFRICAN MUSEUM

be obtained from the sample squares alone and the extraction of microfauna
from the other squares was discontinued.

The modern comparative material derives from both bulk samples and
complete owl pellets. A bulk sample was collected from an unmeasured area at
Byneskranskop 2. From both this and a similar sample from Nooitgedacht B,
the bones were subsequently extracted by hand; the high proportion of bones
and lack of fine matrix made sieving unnecessary. At Glentyre as much
diagnostic cranial material as possible was collected from the floor of the site,
care being taken not to overlook the smaller species. At Boomplaas B and C,
bulk samples were collected from a measured square metre, an area which it
was thought would provide an adequate sample. The area was marked out to
cover places where the pellets tended to accumulate and it was assumed that
there would consequently be a high proportion of bones to general matrix. This
mostly comprised Procavia capensis (rock hyrax) dung which was whole on the
surface, disintegrated below and compacted beneath that. The top two categor-
ies were collected and screened in the 2mm meshed sieve. The bones were
then extracted and the dung discarded. Pellets were first measured and
weighed. They were then pulled apart when dry and the bones were extracted.
This system was found to be the most efficient, although it is known that some
workers prefer to soak the pellets first (Vernon 1972: 11). It was not thought
necessary to develop a mechanical method such as that used by Southern (1954:
389). An account was kept of the contents of each pellet, including the
presence of birds, frogs and the occasional beetle, although non-mammal
material was not further considered.

IDENTIFICATION

The identification of small mammal remains from archaeological sites is
hindered by the broken nature of the material. Most distinguishing characters
employed by zoologists are missing. These include external characters such as
colour of pelage, relative tail and body length, and numbers of mammae and
internal characters such as chromosome counts. Sometimes cranial and dental
characters are invoked in keys, but this is usually after basic distinction has been
made on other characters not preserved under archaeological and palaeontologi-
cal conditions. The need for a key based solely on cranial characters was
recognized by Hanney (1962), who published such a key to the small mammals of
Nyasaland (Malawi). In 1972 Coetzee published a similar key for southern Africa
expressly for the identification of remains from owl pellets. Such keys are
extremely useful, especially if one is dealing with complete or nearly complete
skulls. Unfortunately, this is seldom the case in archaeological contexts so that
the pioneer work of Davis (1965), who distinguished species by their alveolar
patterns, a logical extension of the established practice of counting tooth roots, is
of great importance. More recently Knox (1976) has examined the alveolar
patterns of Muridae in Australia. Misonne’s (1969) comprehensive study of the
Muridae is based on the teeth and as such can be usefully employed.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 211

In practice it was found that distributional data such as those of Davis
(1974), Meester (1958, 1961, 1963), and Coetzee (1972) gave a basic indication
of the species likely to occur in the southern Cape. It is emphasized, however,
that this merely provides a starting point; it is in no way considered to be rigidly
inclusive or exclusive since there is, after all, every reason to believe that
changes in distribution have taken place (Meester 1958; Brain & Meester 1964;
Avery 1977). Subsequently a key such as that of Coetzee (1972) is useful for
identifying the more obvious species and perhaps for eliminating other possi-
bilities.

Although it is possible to identify definitely the majority of specimens,
there remain certain cases where doubt is likely to arise, particularly in closely
related species. The following remarks refer to such areas of possible confu-
sion:

1. Praomys spp. and Saccostomus campestris

The mandible of S. campestris may be distinguished from those of Praomys
spp. by the facts that the body is relatively much deeper and that the anterior
border of the muscle attachment is not contiguous with the mental foramen as
it is in Praomys. It was not found possible to distinguish the mandible of P.
natalensis from that of P. verreauxi with any degree of certainty. The teeth of
the latter are relatively smaller than those of the former, but that is not a
distinguishing factor due to overlapping ranges.

2. Acomys subspinosus and Steatomys krebsi

Because the region of M; is frequently broken, it is often necessary to
resort to other characters to distinguish the two species. For instance, Acomys
has a broad zygomatic process and no masseter knob, whereas Steatomys has a
narrower zygomatic process and a masseter knob; the anterior palatal foramina
penetrate between the roots of the upper first molars in Steatomys but not in
Acomys. Several differences are apparent when the mandible is viewed in the
buccal aspect. In Acomys the anterior border of the muscle attachment is level
with the anterior alveolus of M, and is, therefore, situated closer to the mental
foramen than is the case in Steatomys, where the anterior border of the muscle
attachment is level with the midpoint between the two alveoli of M,. In
Acomys the symphysis forms a visible angle with the ventral border, which
itself forms an almost straight line. In Steatomys, on the other hand, the
junction of the symphysis with the ventral border is not visible but is, instead,
incorporated in the general curve of that border.

3. Dendromus spp. and Mus minutoides

Davis’s (1965: 145) potentially useful point that M. minutoides possesses a
three-rooted M, should have provided the key to distinguishing the mandible of
this species from those of Dendromus spp. Unfortunately, however, it was

zie ANNALS OF THE SOUTH AFRICAN MUSEUM

found that this feature was not constant in the material examined. While it
seems that the posterior alveolus of M, in M. minutoides is at least broader
than the corresponding alveolus in Dendromus, this fact is not demonstrable
with any degree of certainty. Consequently, where doubt existed, the speci-
mens were assigned to the category ?Dendromus as these species generally
occurred with greater frequency. It is implicit in the above that the mandibles
of the two species of Dendromus were also indistinguishable when the teeth
were missing.

4. Myosorex varius and Crocidura cyanea

Although it is generally possible to distinguish between these two species,
there remains a small body of material that cannot be identified. This consists
of the central portions of the mandible and P*'to M? regions of the maxillae
when the teeth are missing. In these cases, specimens were assigned to
?Myosorex since this genus was invariably found to be numerically superior on
the basis of identified specimens.

5. Crocidura cyanea sspp.

The only dental character given by Meester (1963: 48) to distinguish
Crocidura cyanea cyanea from C. c. infumata concerns M3. Since this tooth is
frequently missing it is rarely possible to distinguish the two subspecies in
subfossil material The only reason for attempting such a distinction is that the
two subspecies live in different environments and may, therefore, be useful in
the interpretation of past environments.

6. Otomys spp.

Identification of the various species of Otomys has proved difficult. The
teeth are easily lost, especially the M° which, together with the M,, is the most
diagnostic tooth. Moreover, there is a great deal of variability in the numbers
of roots and, therefore, of alveoli. General trends are noticeable which allow
assessment of specimens with reasonable but not absolute certainty. Very large
specimens are likely to be O. laminatus, especially if there are a large nuniber
of alveoli since the number of these seems generally to be proportional to the
number of laminae. In this way O. unisulcatus, which has fewer laminae, tends
to have fewer alveoli. This latter species is, however; roughly the same size as
O. saundersae and as such can only rarely be distinguished with any degree of
certainty if the teeth are missing. The most frequently found species in most
cases are O. irroratus and O. saundersae, of which the latter is considerably the
smaller. There are, however, specimens of an intermediate size which must
necessarily be assigned somewhat arbitrarily to one of these species and it is
possible that some of the larger specimens may be O. /aminatus and not
O. irroratus. It is felt, however, that while the more frequent species may tend
to be over-represented, the proportions are generally accurate.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 2s

THE NATURE OF EXCAVATED SAMPLES

The excavated sample bears a complex relationship to the biocoenose (life
assemblage) from which it is derived. Examination of the taphonomic processes
involved provides some understanding of that relationship and its consequent
effects upon interpretation. It is also of importance to consider the quality of
the sample in order to assess the type of deduction it is possible to make from
the data. Moreover, the fact that the excavated samples generally comprise
material accumulated over a considerable period of time raises a question as to
the extent to which periodic natural fluctuations will be noticeable.

Depending upon the type of investigations being made, consideration of all
these points may not be necessary. For instance, for the purpose of making
relative environmental interpretations it may be necessary to establish the
relationship of one sample to another but not to the original communities.
Thus, establishment of the fact that similar biases were affecting the samples
would be important but not necessarily the exact nature of those biases. If,
however, an attempt were to be made to establish absolute changes in condi-
tions with, perhaps, a quantitative approach, then the relationship of the
sample to the population would become important. Equally, any work on
population structure would demand such a base. In general it would appear
advisable to examine such questions as an aid to better understanding of the
overall subject, whether or not it has any immediate relevance.

Taphonomic factors

The theoretical sequence of events from life assemblage to the collected
sample is discussed by Clark et al. (1967). Behrensmeyer (1975) considers the
taphonomic processes involved, with particular reference to large mammals in
open situations. Brain (1967) has examined differential preservation of bone
near a Hottentot (Nama) village in South West Africa, as have Binford &
Bertram (1977) for some American sites. Both the latter sets of data are of
theoretical importance, although they are of cultural origin.

Given a life assemblage whose composition is regulated by various biotic
factors (Clark et al. 1967), the initial factor governing which bones will be
included in the deposit is the mode of death (Behrensmeyer 1975). In the
present case this would appear to be almost entirely due to predation by Tyto
alba, as has been discussed above. The possible biases caused by T. alba are
considered below. In the present context, however, it is necessary to examine
the possibility of further biases due to differential damage of the bones by the
predator. In fact it appears that 7. alba causes very little damage to small
mammal bones (Grindley et al. 1973; pers. obs.), so it seems unlikely that this
will cause any significant bias. Although T. alba is said to feed only the body of
the prey to its young, it apparently eats the head itself (Vernon 1972: 109).
There has, however, been no indication that there is any selectivity in prey fed
to young birds. Failure of the adult to drop its pellet at the roost site should,
therefore, have the effect of reducing numbers indiscriminately if the skulls

214 ANNALS OF THE SOUTH AFRICAN MUSEUM

alone are counted. Glue (1967) noted one case of an owl not eating the head of
its prey but was of the opinion that this did not happen often and that a count
of skulls and mandibles gives a reliable count of prey taken. This is an
important point since in the present study only mandibles and maxillae were
counted. A small addition to the sample collected by the predator may be made
by small mammals dying in the cave from other causes. The identity of such
animals is mentioned elsewhere.

The thanatocoenose (death assemblage) thus accumulated is_ further
reduced by factors of preservation related to weathering, transport and burial. In
the present case, the fact that the bones were deposited in rock shelters or caves
should have the effect of protecting them from the extremes of weathering. They
will also be protected initially by being enclosed in the owl pellet which
effectively reduces the period of time between exposure and burial. Transport
must necessarily have been limited in the cave situation, but one real source of
damage will have been trampling by people and animals. Of the taphic or burial
factors listed by Clark et al. (1967: 117) the nature of the sediment could be
relevant in the present context. It is possible, for instance, that the coarse matrix
provided by shells or stone artefacts will adversely affect the preservation of
small mammal bones. Similarly, a rock fall or high degree of spalling could well
damage or destroy such bones. Of the post-depositional factors, the actions of
burrowing animals and the effects of permeating solutions may possibly affect
small mammal bones in caves. In the case of the former, however, it may be that
the bones will generally tend to be disturbed rather than destroyed, although the
digging of storage pits by humans could have a more serious effect. Damage by
erosion within a cave is almost certainly a minor consideration, but that caused
by leaching is likely to have been greater. The lighting of fires in the caves could
destroy bones on or even below the surface.

In general it would seem that these various factors will act more or less
indiscriminately. This is because there is relatively little variation in the size of
the specimens, which suggests similar reactions to destructive forces. Some
correlation of size with susceptibility to destruction or damage is suggested by
both Clark et al. (1967) and Behrensmeyer (1975), who note that the effects of
weathering are less on small bones than they are on large ones. The chances of
disproportionate preservation are probably reduced in the present case by the
fact that only maxillae and mandibles have been considered. In particular the
body of the mandible, which is constructed of dense bone with a consequently
high survival rate (Brain 1967), will perhaps have a similar chance of being
preserved in animals of a similar size. It would seem, therefore, that in the
present case the fossil assemblage will constitute a reasonably unbiased sample
of the thanatocoenose.

Sample quality
Clark et al. (1967) discuss the factors responsible for the differences
between the total fossil assemblage and the collection ultimately available for

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS DA

interpretation. The first group of factors is connected with collecting. In the
present context this could mean that there will be a bias against the smaller
species either because the material was sieved through an insufficiently fine
screen or because the specimens were overlooked in the subsequent sorting.
Jaws which are prone to losing their teeth are also at greater risk of being
overlooked. Personal biases such as training, persistence, psychological
fluctuations, visual acuity and physical comfort are all considered by Clark et al.
(1967) to affect collecting which, in this case, constitutes retrieval of the
specimens from the excavated matrix. Where a group of collectors is involved
even these factors will become variable, although they may balance out.

The data which are available for interpretation may be different again from
the material collected. Here the factors relate to identification and the estima-
tion of minimum numbers of individuals represented. In the present case the
question of differential ease of identification (Clark et al. 1967: 120) does not
apply because, as is explained below, each individual is represented only by its
jaws. This means that all individuals are, in effect, identified at the same level
despite the fact that it might have been possible to identify some species on
much less evidence. Grayson (1978) has further pointed out that rare species
tend to be heavily over-represented because the fewer the elements present the
greater the chance that each element will represent another individual. Simpson
et al. (1960: 116) have also noted that animals with more bones have a
potentially greater chance of being represented in the sample. In the present
context, however, the representation of each individual by the same four bones
means that such biases are eliminated and each individual has an equal chance
of being included.

Excavated samples cannot meet the usual specifications for homogeneity of
faunal sampling of live populations (Simpson et al. 1960). They must therefore
necessarily always be more heterogeneous. Thus the sexes will not usually be
distinguishable and age distinctions may not always be possible. This fact could
have an important bearing on attempts to interpret interpopulation differences.
For instance, it will be difficult, if not impossible, to establish with certainty the
cause of variation in mean individual size. The change may be due to a
difference in sexual or age composition of the sample rather than to any real
change caused, perhaps, by a response to climatic variation. Whilst it may be
possible to control the age factor, it will not be possible, in small mammal
cranial material at least, to control the sexual factor.

Neither is the available sample always of a size to represent adequately the
population from which it was drawn. This means again that it is not always
possible to come to reliable conclusions about the degree of similarity or
difference between different populations. In another context the size of the
sample may affect its composition. The question of the size of the sample
necessary to include all available species is discussed below. If, as is suggested,
the size of the sample must increase with the number of possible species, it
follows that the number of species that may be omitted from a small sample

216 ANNALS OF THE SOUTH AFRICAN MUSEUM

must also increase in relation to the possible number of species. Therefore, the
absence from a small sample of a species which never forms a large proportion
of any sample cannot be regarded as significant because it could be due to
sampling error. Only where species normally form a large proportion of a
sample may their absence be ascribed to some real change in conditions.
Generally, however, presence is more significant than absence and changes in
proportional representation may provide useful data for interpretation.

Levels of inference

It has been shown that some rodent populations are subject to periodic
and, in some cases, spectacular fluctuations in size (Honer 1963). Such
fluctuations are of a cyclical nature and may be due to a number of causes
(Krebs 1972). They may occur approximately every 3 years (Honer 1963) or
over longer periods such as every 8 or 10 years (Davis 1966). Moreover,
trapping programmes in the south-western Cape have demonstrated that
seasonal variation in population size may also occur (R. C. Bigalke 1975 pers.
comm.). Other irregular fluctuations may be caused by events such as flooding
(Davis 1973; Brooks 1974) or fire. The effects of the latter appear to be quite
variable. In Ugandan grassland all species had returned to an area 7 months
after burning and by 11 months there was little difference between the burnt
and unburnt areas (Neal 1970). R. C. Bigalke (1975 pers. comm.), on the other
hand, has found that in the south-western Cape a period of 3 or 4 years elapses
before burnt ground is fully recolonized.

In none of these cases is the cycle of fluctuations longer than a decade. For
this reason one would not expect to be able to detect evidence of such
fluctuations in material from archaeological sites. This is because each exca-
vated sample of micromammalian fauna is likely to comprise material accumu-
lated over a period of some hundreds of years. It would seem most unlikely,
therefore, that the short-term fluctuations described above will be discernible in
excavated samples. The only possible exception is the effect of fire on long-
term averages. This is not because the effects of each burn are long-term, but
because the effects of possibly frequent burning could be cumulative. Kruger
(1979) is of the opinion that natural fires in the fynbos were likely to have
occurred randomly at about 6 to 30 or 40-year intervals. It is also presumably
possible that the frequency might have varied at different times in the past.
Moreover, fires caused by humans are likely to have become increasingly
common. In the first instance, people might have fired the vegetation in order
to encourage growth of geophytes (Deacon 1976: 174). Later, pasture control
by burning became prevalent after the introduction of sheep into the southern
Cape around 2 000 B.P. (Deacon et al. 1978). It seems unlikely, however, that |
fire will have had a differential effect on the micromammalian fauna, for two
reasons. The first is that, even if the diurnal species are most likely to be
reduced by fire (Neal 1970), these species are least frequently preyed upon by
Tyto alba (see discussion below). If, however, fire may be assumed to destroy

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 2Y7

food resources indiscriminately, then all or most species will be equally
affected. The second reason is that, depending on the extent of the burn, the
owl may either extend its hunting territory or move its roost temporarily. In the
first case the effects of a short-term change in prey composition would probably
not affect the longer-term average. In the second case the overall size of the
sample and not its composition would be affected. It would appear, therefore,
that such long-term fluctuations as are evident in excavated samples must have
been caused by large-scale events or trends outside the range witnessed on a
short-term basis.

While it is clear in general that excavated samples can only monitor major
trends or fluctuations, the actual size of these may vary. This is because
different samples represent the average situation over varying periods of time.
It is normally not possible to regulate samples so that, within a sequence, they
all represent an equal period of time. This is, however, unimportant if there is
some indication, preferably a suite of radiocarbon dates, of the actual or
relative period of time represented. In this way it will be possible to assess the
relative importance or amplitude of a trend. Thus, an isolated fluctuation
shown by a sample accumulated over a period of 100 years is minor compared
with a generalized trend evident over a period ten times as long.

THE NATURE OF MICROMAMMALIAN EVIDENCE

The use of micromammalian remains as evidence of environmental change
requires the prior assessment of the nature and possible limitations of the
evidence. In the first instance it is necessary to establish whether, in fact, the
evidence can show that environmental change has taken place. Thereafter, in
general terms, consideration must be given to matters relating both to the
micromammals themselves and to the predators that are thought to have been
responsible for accumulating the samples.

Micromammalian evidence and environmental change

It has been assumed that micromammalian remains may be used to
indicate environmental change. The validity of this assumption must now be
tested. It may be, for instance, that changes in the composition of the small
mammal fauna represent an autogenous succession to a stable climax such as is
known to occur in plants (Krebs 1972). Cyclic change, again as observed in
plants, is another possibility. At the population level, fluctuations about a mean
density level are known to occur under stable environmental conditions (Klomp
1962). However, all these changes tend to operate towards the establishment of
homoeostasis, or a state of equilibrium, which is said to operate at all levels of
the ecosystem (Odum 1971: 34). This would suggest that any extension to the
regular range of fluctuation must represent disruption of the equilibrium. In
other words, since climate is the overall factor controlling the vegetation of an
area (Krebs 1972: 432) and animals are directly dependent on plants (Krebs
1972: 416) it follows that any unusual change in the rodent community must

218 ANNALS OF THE SOUTH AFRICAN MUSEUM

Rodentia (Praomys verreauxi )

premaxilla

incisor

diastema

anterior palatal foramen

cheekteeth (m!-3) = upper tooth row (UTR)
alveolus

posterior palotal foramen

zygomatic arch

left mandible - lingual aspect

cheekteeth (M)_3) = lower tooth row (LTR)
—— _______diastema

incisor

body of mandible

Insectivora (Crocidura flavescens)
length of tooth row ce

5 lower alveolar length (LAL) ao
4

=)

right mandible - buccal aspect

j'-3
C

5CBLC BD
b>

{ coronoid spicule

O 10mm coronoid process

I, (procumbent)
eee eect
approx. condyle

length of mandible

| plus incisor

height of ascending ramus

Chiroptera (Rhinolophus clivosus)

Aa premaxilla
2)

f ae
\

PINA ae , infraorbital conal

I Nera) right mandible - buccal ospect

Fig. 7. Diagrams of species representative of micromammalian orders with terminology used.
(See Meester 1963 and Rosevear 1969 for further explanations. )

rostral trough

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 219

reflect similar change at some higher point in the chain of influence. At the
same time it must also be borne in mind that not all species may react, or they
may react differently at different rates, to a particular change, especially in
areas of intermediate climatic change (Tchernov 1975: 346).

The type of change found in the community is not, in fact, illustrated by
the subfossil small mammal material. The pattern of change does not show the
progressive stages characteristic of a succession. It is to be expected that such
stages would be characterized by one or more dominant species. These would
be replaced or succeeded by other species after a period of time and there
would not be evidence of a resurgence of a species once it had been replaced.
However, the changes visible in the sma!l mammal record (Figs. 8-9, for
example) represent fluctuations which are not consistent with the succession
model. Equally, there is no evidence that the changes are truly cyclic as
described by Krebs (1972: 432). The data do not show that one species is
regularly replaced by and, in turn, replaces another.

Two facts are observed about population density. One is that is does not
rise indefinitely and the other is that it varies in different environments (Chitty
1960). This implies that there are mechanisms which prevent unrestricted
increase in density and others which control mean density, all of which tend to
maintain a state of equilibrium. There has been considerable discussion as to
the nature of these mechanisms. It has been suggested that climate, enemies
(predators, disease, etc), self-regulation or a combination of all three could be
responsible (Krebs 1972: 287). It seems likely, in fact, that the dominant factor
will vary depending upon the favourableness of the environment, as suggested
by Huffaker & Messenger (1964, quoted by Krebs 1972: 280). Thus, in
marginal areas climate may be the most important factor, whereas in optimal
areas natural enemies or decline in the quality of the individual (Chitty 1960)
may be responsible for regulating numbers. Equally, such mechanisms as
territoriality may control numbers below a level at which the food supply would
be destroyed.

It has been suggested above that the fluctuations which are visible in the
subfossil record are on a scale far larger than those which occur in a stable
environment. If this is the case, it would appear that the mean population
densities of different species must have altered at various times in the past.
Mean population density has been observed to be higher in optimal areas than
in marginal areas. If, therefore, the mean population density is shown to vary
from time to time in one geographical area, it must mean that the suitability of
that area has varied. It would not be possible, given all the regulatory
mechanisms discussed above, for such changes to occur unless a major para-
meter were altered. In the present case, since many of the animals are
herbivorous mammals, it would appear that changes in the vegetation would
have the most important effect on the favourableness of the environment. In
the case of the insectivorous mammals it may be that a change in the insect
fauna and/or change in the vegetation was the cause. In both cases it is to be

ANNALS OF THE SOUTH AFRICAN MUSEUM

C.flavescens

20 C.hottentotus

5S _S.campestris

16) — ee —
5 _S.krebsi
O

SE San I ee ee ee ee)

10 M.albicaudatus

5 _Q.laminatus

e@eeeoete e

O.irroratus

30

20

A.namaquensis

|

5S _Rverreauxi
S .R.pumilio

e e

abcdefghi j k Imnopq rs tuvwx

ss Oo. OP a ore 2

DGL i LPC q- OLP2

BLD3 ) TOE r OPS

BL k, BPI s BOL

BLA Li aBPRe t BOLI-3

BRL mi BRS u BOL4

CL n BP4 v BOLS

GWA oO. JOLP w OCH

LP p OEP x LOH
FACTOR 3

%o

5 Q.unisulcatus
eel eee
FACTOR 2

Yo

5 C.duthiae

O

40 Mvarius

30

20

10

O

39 S.varilla

7,

5 _D.melanotis

O Je. sogme

40 O.saundersae

5 A.subspinosus

————————

M.minutoides

ol

s . seas 2

abcdefghijkIimnopqrstuvwx

Fig. 8. Variation in percentage representation of species loading highly
in the analysis of species from Boomplaas A (. = less than 0,5%).

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 221

A.namaquesis
M.minutoides

O.irroratus
O.unisulcatus

FACTOR |! a C.duthiae k Pverreauxi
b E.edwardi | R.pumilio
c Mvvarius m M.albicaudatus
d Svvarilla n D.melanotis
e C.cyanea o S.krebsi
f C.flavescens p S.campestris
g C.hottentotus q O.laminatus
h A.subspinosus r O.saundersae
i s
if

FACTOR! 2
30) OP
“fo
20
10
FACTOR 3

fo) _

40. OLPI

30_ BLD3

10

fe)

abcdefghijkIimnopqrst abcdefghijkIimnopqrst abcdefghijkIimnopgqrst

Fig. 9. Proportions of species from levels loading highly in the
Varimax solution of the analysis of levels at Boomplaas A (. = less
than 0,5 7%).

222 ANNALS OF THE SOUTH AFRICAN MUSEUM

expected that a change in the climate would be the initial factor responsible for
altering the quality of the environment. In fact, considerable changes are
known to have occurred in the climate during the Upper Pleistocene and the
Holocene, and consequent changes in the vegetation must have taken place
(Van Zinderen Bakker 1976). It is these long-term changes which are most
probably being reflected in changes in species composition of the micromamma-
lian fauna. Significant variation in mean individual size in different populations
of the same species would tend to support the view that the evident changes
could not have resulted from minor, albeit regular, fluctuations since such
physical responses take time to come into effect.

Direct micromammalian evidence

Theoretically, direct information from micromammals is available at the
individual, population and community levels. At the individual level changes in
average size may indicate changes in prevailing climatic conditions. This is
because Bergmann’s Rule states that within a given species those members
living in colder climates will tend to have a greater body mass than those living
in warmer climates (Coon 1962: 59). Guilday (1971: 252) points out, however,
that this rule is not universal, some forms being unaffected and some exhibiting
a negative response. If it is possible to establish how a particular species reacts
with respect to this rule, it may be possible to furnish fairly precise information
concerning past temperature fluctuations. The possibility also exists of corre-
lating variation in average rainfall with differences in body mass, as was
demonstrated in Israel for Spalax ehrenbergi by Tchernov (1968). Physiological
studies on the moisture and temperature tolerance or requirements of live
animals could perhaps provide information on the usefulness of different
species as indicators of these phenomena, as well as the actual limits repre-
sented. The accumulation of such data would, however, require extensive
long-term investigation.

At the population level it is important to ascertain as precisely as possible
which aspect of the habitat is responsible for governing distribution. It will
largely be this aspect that the population or species may be said to represent for
the purposes of interpretation. It is also likely that changes in the density of a
species will indicate changes in the parameter with which that species is
correlated. Information concerning habitat preferences of individual species
largely comprises general field observations by various collectors. The data
provided generally give an indication of plant life-form and state of ground-
surface preferred by the species in question, as well as some basic information
on food preferences and activity patterns. Detailed distributional data such as
those of Lynch (1975) for the Orange Free State are very useful in that they
itemize the constituent parts of the habitats in which different species occur. All
such information, however, suffers the major disadvantage of being purely
descriptive. What is needed is a quantitative assessment of the relative impor-
tance of different elements of the habitat in determining the distribution of

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS pa

individual species. In this respect the work of Rosenzweig & Winakur (1969) is
of great theoretical importance, and that of Bond et al. (1980) on local species
is of direct relevance. These studies consider the factors governing distribution
of different species. In both cases the conclusion is that plant life-form and
foliage density exert major influences on small mammal distribution, and in a
variety of ways as is discussed by Saint Girons (1977). Plant species diversity,
on the other hand, appears not to be of great importance. This would appear to
confirm the statement by Schulz (1953, quoted by Davis 1973) that density and
nature of cover are far more important than the plant species involved. The fact
that plant life-form is apparently so important it would suggest, in fact, that even
available general ecological information may allow some acceptable interpreta-
tion of past vegetation changes. It is important, however, to note that micro-
mammalian evidence cannot now, and probably never will, provide information
concerning the floristics of a given area.

Qualities of ground surface were also assessed for their role in determining
species distribution. Rosenzweig & Winakur (1969) concluded that these abio-
tic variables were of secondary importance. Bond ef al. (1980), on the other
hand, showed that specific size and abundance of stones or rocks were impor-
tant habitat components for Aethomys namaquensis and Acomys subspinosus,
although in the former case the correlation might have been fortuitous. Nel &
Rautenbach (1975) found that the structure of the substrate restricted distribu-
tion of burrowing species. In so far as the nature of the substrate and the
amount of rainfall affect vegetation, these factors may also be said to influence
the distribution of species reliant on type of vegetation cover.

Changes in population density will appear as changes in community com-
position in the subfossil evidence. This is because this evidence is such that it
shows only the relative abundance of the constituent populations of the small
mammal community. It would be difficult, if not impossible, to demonstrate
absolute change in individual mean population density because the relationship
of the sample to the living population is not known. In fact, it is likely that
changes would have the effect of changing community structure whether they
resulted from the simultaneous independent reaction of individual populations
or from interrelated reaction. It may be possible, and it is certainly desirable, to
distinguish the two types of reaction. The latter, for example, may appear as an
inconsistency in the evidence and, as such, should be recognizable and explain-
able. Such a situation, which could be due to interspecific competition, may be
illustrated by the relationship of Otomys irroratus and Rhabdomys pumilio.
Both have a preference for dense vegetation, but R. pumilio has wider
tolerance. Brooks (1974) concluded that when there is competition in the area
of overlap the narrow-niche species, O. irroratus, has the advantage over the
broad-niche species, R. pumilio, because of greater specialization. R. pumilio
would then be forced to make use of less suitable habitats where, presumably,
its density would be reduced. In this situation the increase in O. irroratus and
decrease in R. pumilio would be interdependent and apparently contradictory.

224 ANNALS OF THE SOUTH AFRICAN MUSEUM

An independent parallel response is suggested by the decrease in O. irroratus
and increase in Aethomys namaquensis which would result from reduction in
plant cover and change to more arid vegetation. Work such as that of Neal
(1970), Sheppe & Osborne (1971), and Nel & Rautenbach (1975) shows habitat
utilization in operation at the community level. This suggests that it should be
possible to establish a correlation between community composition and a
known set of conditions. Thereafter, if the relationship of the various species
can be determined, it may be possible to postulate the manner in which the set
of conditions must have been different to support a differently composed
community.

Predator-related factors

An ideal sample will represent exactly the size and composition of the
living population from which it is taken. However, as Simpson et al. (1960: 110)
have pointed out, perfect sampling is an impossible ideal which not even the
most rigorous collecting methods will achieve. In the present case, the sample
does not approach the ideal because the method of its acquisition prevents this.
It is necessary to attempt to determine the manner and extent of its deviation
from the ideal. The extent to which the collecting habits of the predator may
have biased the sample must be investigated. Tyto alba (barn owl), which is
assumed to have been the predator responsible, has been shown to be an
effective sampler of small mammal populations in that it preys on most, if not
all, of the species to be found within its hunting territory (Vernon 1972; Dean
1977). On the other hand, it has also been shown not to take its prey in direct
proportion to relative abundance (Glue 1967; Hanney 1963).

It must be noted at the outset that the methods used to check the
relationship of the sample taken by the owl to the structure of the living
population in the area hunted, may themselves be subject to bias. Trapping is
the most usual method employed since it is the only viable way of controlled
sampling. Here the type of trap used can affect the results (Neal & Cock 1969;
Wingate & Meester 1977). Moreover, some small mammals are known to be
trap-shy and some will not take bait (Hanney 1963; Davis 1973). All these
factors may operate to bias the sample, although, being aware of the problems,
workers generally take steps to reduce their possible effect. There remains,
however, the problem of the extent to which trapping results may be directly
comparable with those obtained from owl pellets. Glue (1967), for instance,
points out that the owl sometimes hunted outside the area trapped and that this
fact could, to some degree, influence the validity of the comparison. Hanney
(1963), on the other hand, suggests that direct comparison of his results is
precluded by the fact that the pellets were accumulated over bi-monthly
periods, whereas the trapping was carried out at the end of each such period.
The pellets thus constituted an average for the period, whilst the trapping
represented the situation only at the end of the period. In view of seasonal
variation this could well affect the comparison of the two sets of data. The

———— a

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 225

composition of prey-remains of other birds of prey or carnivores which hunt in
or near the territory of 7. alba may provide some confirmation of the data. In
general, however, it is to be expected that too many variables will be operating
to make comparison useful for assessing biases inherent in the 7. alba sample.

The effectiveness of T. alba as a small mammal sampler is indicated by the
fact that Vernon (1972) found four more species in owl pellets from the S.A.
Lombard Nature Reserve than did Meester (1955) by trapping. Coetzee (1963)
encountered a similar situation in the Kruger National Park. Glue (1967) and
Dean (1977) have also shown that 7. alba takes a far wider range of prey than
do other raptors with which it was compared. In this context it is interesting to
note that Honer (1963) found that 7. alba in the Netherlands displayed a
preference for hunting-territories that showed a range of elevation. In such
situations there would presumably be more niches available and, therefore, a
greater variety of small mammal species. The number of pellets it will be
necessary to analyse will depend upon the complexity of the community on
which the owl is preying. In some areas where few species occur, a small
collection will suffice. In other areas such as the Cango valley which has a very
rich small-mammal fauna, a large collection will be necessary. This is shown by
the fact that out of a total of 2 126 individuals from pellets and bulk samples,
only two each of Dasymys incomtus (shaggy swamp-rat) and Steatomys krebsi
(Cape fat mouse) have been recovered (see Table 7). Moreover, the only
specimen of Crocidura flavescens (red musk-shrew) from Nooitgedacht A was
recovered from pellet no. 343. The presence of what one might term trace
elements does, however, suggest that over a period of time virtually all the
species occurring in an area will be sampled by T. alba.

There are, of course, physical limits to the size of prey which the owl will
be capable of taking, so that only that part of the mammal community within a
certain size range will be sampled. The limits of prey taken by 7. alba have
been discussed above but it is important to note that these limits include all the
Insectivora and virtually all the Rodentia that are likely to be found in the
southern Cape. Within the limits, however, there is the possibility of a bias in
favour of animals at the larger end of the scale. This is because, as Sparks &
Soper (1972: 76) have pointed out, all things being equal, it would be more
economical in terms of the bird’s energy budget for the owl to catch the larger
animals. If, therefore, a relatively small species is a particularly abundant prey
item, it would appear either that sparse cover and/or an abundant population
make capture easy or that very small numbers of larger prey species are
available. In this case, a deviation from what may be regarded as the norm
could prove significant for the purposes of interpretation.

A second bias is introduced by the obvious need for coincidence in activity
patterns in the predator and the prey, a point also made by Davis (1958) and
Brooks (1974). Those species which are diurnal are clearly less likely to be
preyed upon by 7. alba, which is nocturnal. In this context Dean (1977)
recorded a higher proportion of diurnal Rhabdomys pumilio (striped field-

226 ANNALS OF THE SOUTH AFRICAN MUSEUM

mouse) in pellets of Asio capensis (marsh-owl), which hunts during the day,
than in pellets of 7. alba. Brooks (1974) recorded a similar situation. It
therefore follows that there is likely to be an unnaturally low proportion of
diurnal species in a sample collected by 7. alba. This, however, is unlikely to
cause a major imbalance in the results because the majority of the insectivores
and small rodents is nocturnal, crepuscular or active night and day. Apart from
R. pumilio, only the Macroscelididae (elephant-shrews) are said to be diurnal
(Roberts 1951: 26).

There is also some evidence to suggest that T. alba is selective in its choice
of major prey item. Glue (1967) analysed pellet contents from a site in England
and undertook a trapping-programme in the area hunted by the owls. This
showed that almost twice the proportion of Microtus agrestis (short-tailed vole)
(79 % as opposed to 40%) was found in the pellets as in the traps. Conversely,
Rattus norvegicus (brown rat) and Arvicola terrestris (water-vole), which were
shown to be present in reasonable numbers, seldom occurred in the pellets.
Glue (1967: 178) concluded that the owl either preferred M. agrestis or found it
easier to catch, even though the two sets of data may not be directly compar-
able, as was discussed above. Hanney (1963) found a similar lack of correlation
between trap results and pellet contents. Although Praomys natalensis (multi-
mammate mouse) was dominant in both, proportions were generally very much
higher in the pellets, which suggests concentration on this species out of
proportion to its relative abundance on the ground. From this it would appear
that, although its actual numerical importance may be exaggerated, the species
favoured by the owl will tend to be numerically dominant on the ground. This
seems reasonable in view of the fact that such a species must normally be more
at risk than a rare species. Indeed, Craighead & Craighead (1956: 364)
maintain that prey density is of primary importance in determining on what
raptors will feed. Furthermore, they are of the opinion (Craighead & Craig-
head 1956: 138) that, in general, the diet of the raptor will reflect prey densities
of those species available within the habitat of the predator and vulnerable to
1G:

In some cases a relationship has been noticed in changing proportions of
major prey items. This not only illustrates the types of reaction discussed
above, but adds the further possibility of the introduction of predator bias.
Craighead & Craighead (1956: 182) make the general point that as density or
vulnerability of the major prey item declines, other species will be taken to a
greater extent by raptors capable of taking them. An example is given by
Hanney (1963), who noted that there was a tendency for numbers of Crocidura
sp. (musk-shrew), the secondary prey item, to vary inversely in relation to
those of P. natalensis, the major prey item. Whether increases in numbers of
Crocidura sp. in pellets reflect increases in actual numbers or whether they
were an artefact of reduced numbers of Praomys natalensis, is not known. Two
further examples suggest that both situations are possible. Dean (1977) noted
that in 7. alba pellets proportions of P. natalensis declined over a three-year

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 22)

period while proportions of Otomys angoniensis (Angoni vlei-rat) rose. In this
case the fact that the same shift in emphasis was observed in the prey remains
of Tyto capensis (grass-owl) and Asio capensis (marsh-owl) may suggest that
there was a real switch in the numerical importance of the two species. Glue
(1967), on the other hand, found that when the number of Microtus agrestis
declined, the owls turned their attention to Rattus norvegicus, a species which
previously they had all but ignored. There was, however, no indication of a real
increase in numbers of R. norvegicus (Glue 1967: 180), so that the increase
apparent in the pellets was due solely to a reduction in the major prey item. It
is unfortunate that it probably would not be possible to detect such a situation
from the subfossil material. It is perhaps, therefore, more correct to speak in
terms of an increase in relative importance of a particular species, and hence
vegetation type, rather than an absolute increase. This, again, would tend to
confirm the suggestion that it is only possible to show changes in relative
abundance in the community.

Even though the major prey item remains the same, it has been shown that
there may be fluctuations in total prey composition. In England, Glue (1967)
has noted that a greater variety of species was taken during the summer, which
may be correlated in some way with the fact that this was the breeding-season
of the owls. In Malawi, Hanney (1963) found seasonal fluctuations in the
proportions of some species which were unrelated to any shortage in supply of
the major prey item. In the Transvaal, minor prey items constituted a steady
proportion of the total prey but the composition of that minor prey element
varied (Dean 1977). Craighead & Craighead (1956: 183) maintain that an
increase in the proportion of minor prey items will result from a decline in the
vulnerability of the major prey item which, nevertheless, will not itself decline
proportionally. Whatever the reason, such fluctuations would appear to have
been short-term and, as such, not visible in the subfossil record. It is thought,
in general, that the potential contribution of the minor prey elements to the
present study will normally be low, and for this reason they have been given
little weight.

The size of the hunting-territory regularly frequented by 7. alba is thought
to have a radius of about 5 km (Kowalski 1971: 473), but the distance covered
must depend to some extent upon the availability of food. Indeed, Coetzee
(1963: 115) quotes Bodenheimer (1949) as saying that the range in Israel is
between 5 km* and 25 km*, depending upon the availability of prey. There-
after, the question arises as to how far it is legitimate to generalize from the
relatively small area of the owl’s hunting-territory to a wider area. It is unlikely
that vegetational or climatic conditions will be strictly limited to the area in
which the owl hunts, except perhaps in the desert where distribution of
vegetation may be definitely circumscribed. This being the case, it is necessary
to establish the present extent of what may be termed a homogeneous unit. In
order to do so it is first necessary to decide on an acceptable level of
generalization. This process may be illustrated by reference to Boomplaas A.

228 ANNALS OF THE SOUTH AFRICAN MUSEUM

Since the hunting-territory of the owl must encompass several vegetational
categories as defined by Moffett & Deacon (1977), such a category must
constitute too low a level of generalization. In practice both here and else-
where, it will probably happen that a topographic division will provide an
acceptable unit, with the added advantage that it is unlikely to have changed
during the period under discussion. Thus, in this case, the Cango valley may
perhaps be taken as representative of the foothills of the Swartberg Mountains,
as distinct from both the mountains themselves and the Little Karoo. These are
distinct from each other and from the foothills in climate, vegetation and
substrate. Byneskranskop, on the other hand, may perhaps reasonably be taken
as representative of the south-western Cape coastal foreland. When sites are far
apart or in very different situations, comparison must necessarily be on an even
more general scale. Provided that this is made ‘clear, such generalities can still
provide useful working hypotheses which can be confirmed or refuted when
more information becomes available.

METHODS

Investigation of changes in community composition involves computation
of minimum numbers of individuals, multivariate statistical analysis of these
data and, finally, the interpretation of the analyses. The methods used in each
stage are described below. The measurements taken to provide basic data for
the examination of different populations and the indices employed to describe
community structure are also explained.

COMPUTATION OF MINIMUM NUMBERS

In accordance with general practice in micromammalian studies, only the
cranial material was examined, although it is intended at some later date, when
time and a comparative collection allow, to attempt to analyse the postcranial
material. In order to estimate minimum numbers of individuals, left and right
mandibles and maxillae were enumerated separately for each species and the
highest number taken. If several stratigraphic units or squares were to be
considered as a whole, the total counts for each jaw were first obtained ‘and
then the highest number was taken. In the problematic cases of Myosorex/C.
cyanea and Dendromus spp., further calculations were necessary in order to
obtain the correct total minimum number of individuals. The minimum number
of certainly identified individuals was obtained in the usual way. The minimum
number of doubtful specimens was then obtained by subtracting the sum of
certainly identified minima from the total minimum number for the combined
group. If the answer was negative, no doubtful specimens were required to
make up the total. The following examples illustrate the process:

Example 1
L. maxillae: 1 M. varius + 1 C. cyanea = 2
R. maxillae: 2 M. varius + 1 C. cyanea = 3

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 229

L. mandible: 10 M. varius + 5 C. cyanea + 4? = 19
R. mandible: 6 M. varius + 5 C. cynaea + 3? = 14
Minimum number for total = 19
Minimum number M. varius = 10
Minimum number C. cyanea = 5
Minimum number ? = 4

Example 2
L. maxillae: 2 M. varius + 1 C. cyanea = 3
R. maxillae: 1 M. varius + 1 C. cyanea = 2
L..mandible: 11 M. varius + 3 C. cyanea + 1? = 15
R. mandible: 7 M. varius + 5 C. cyanea + 2?.= 14
Minimum number for total = 15
Minimum number M. varius = 11
Minimum number C. cyanea = 5
Minimum number ? = 0

Higham (1967: 302) and Clason (1972: 141) have pointed out that various
methods of obtaining minimum numbers have their disadvantages and may
provide different answers. The method used here is considered the most
practical for dealing with large samples since a more detailed examination of
the material would be too time-consuming. On the other hand, certain precau-
tions were taken in order to standardize counting, with a view to preventing the
possibility of raising the numbers artificially. To this end, jaws were counted
only if they included a certain frequently preserved element that was deter-
mined for each species or group of species. Specimens that did not possess this
element were not counted as they could potentially have broken off others that
did. Similarly, even though a loose tooth of a rare species might have been
noticed, it was not counted because this would have resulted in a lack of
consistency and an over-representation of such species. As it is, the method
employed will have tended towards a general under-representation. It is felt,
however, that a conservative estimate is to be preferred.

MULTIVARIATE STATISTICAL ANALYSIS OF PRIMARY DATA

The quantity of material involved (Tables 1-7) made it difficult to isolate
by simple graphic methods alone the underlying patterns which were assumed
to exist. For this reason it was considered appropriate to employ some form of
multivariate statistical analysis as an aid to interpretation of the evidence. It
was decided that factor analysis could prove useful because of its data-reduction
capabilities. Its main aim is to simplify the data by explaining as much as
possible of the variation present in terms of as few patterns or factors as
possible. In effect, each pattern or factor represents a cluster or group of the
original variables. Conversely, the meaning of the factor is established by
reference to the variables which load highly on it. Both Cattell (19655: 424) and
Rummel (1967: 451) have pointed out that factor analysis can serve a useful
function in the exploration of new fields of research by generating hypotheses.

230 ANNALS OF THE SOUTH AFRICAN MUSEUM

Its simplification of the data allows the identification of patterns not otherwise
apparent, and herein lies its advantage in the present work.

Factor analysis is very complicated and, consequently, there may be
problems in its application, more particularly since it was designed for a
different type of research. As Doran & Hodson (1975: 198) point out, factor
analysis was developed for and by psychologists with a different set of problems
in mind from those encountered by archaeologists. Whilst they are inclined to
doubt its usefulness in archaeology, Cattell (1965a: 192) makes it clear that it
has potential in a great many fields. Thus it appears that, although somewhat
different, its use in the present situation cannot be ruled out a priori. In fact,
the problems that arise would seem to be due mainly to inadequacies in the
data rather than to the inherent unsuitability of the method. These questions
must be considered and evaluated before use of the analysis can be made.

The first question that arises is whether or not the data are suitable for
submission to this form of analysis. The use of factor analysis in the present
circumstances could be questioned on the grounds that the basic data comprise
frequencies; these are ordinal-level measurements (Nie et al. 1975), whereas
interval- or ratio-level measurements are required for factor analysis. It may be
argued, however, that the numbers represent the score of each species for the
given level, which would mean that, effectively, they were ratio-level
measurements. Whilst the logic of this argument may be doubtful in terms of
pure statistics, it is considered to be acceptable for the purposes of the present
research. It was, therefore, considered permissible to use factor analysis in
these circumstances, the more particularly since it is, in fact, the most appropri-
ate analysis to the problem at hand. At some later stage it may become possible
to refine the data base to make it more closely applicable to factor analysis.

The suitability of the samples may also be called into question. Some of
the problems concerning excavated samples have been discussed above, but
there are some which are directly relevant to the analysis. Rummel (1967: 452)
points out that research can centre on describing the data matrix alone, in
which case statistical problems such as the type of underlying frequency
distribution, sample size and randomness of selection are not and need not be
part of the research design. However, although factor analysis may be a
mathematical rather than a statistical tool, some statistical considerations are
involved in the acquisition of a suitable or reliable matrix of correlation
coefficients on which the analysis is based. Moreover, it is important to have
the coefficients of variation of the variables as low as possible. Since calculation
of both coefficients depends on the standard deviations of the variables, it is
clear that some attention must be paid to these.

There are two problems, one concerning the absolute size of the samples
submitted to analysis, and one concerning variation in the size of the different
samples being compared. In the first case the problem more specifically
concerns small samples. Small samples, or rare categories, tend to give
spuriously high correlations, a point mentioned by Doran & Hodson (1975:

a

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS Zou

144). This is because frequent values of 0 mark a departure from the condition
of a normal distribution, which is a prerequisite for calculation of accurate
correlation coefficients. Similarly, a small mean will tend to have a relatively
larger standard deviation, with the result that correlation coefficients will be
artificially raised. In order to obviate this problem, small samples were omitted
from analysis in the present study and only data for the most highly represented
species were used in the analysis. In some cases data for different levels were
either omitted or combined, as has been advised by Cowgill (1968: 373). This,
however, was done as infrequently as possible because of the loss of informa-
tion entailed.

Variation in the size of the samples made it difficult to achieve the desired
effect of keeping variability in the scores of the variables on the cases as low as
possible. In practice it generally happens that when the levels are considered as
variables, a small number of species will dominate the fauna while there are
others that occur in very small numbers. This will necessarily result in high
standard deviations. Similarly, when the species are considered as variables
there are also very high standard deviations because of the considerable
difference in the amount of material available from the individual levels of a
site. If the large standard deviations are caused by obvious outliers it may be
advisable to reduce, or increase, these numbers specifically in some way in
order to moderate their effect on the analysis. In the present work, because
there was such a large amount of material from levels OLP2 and OLP3 at
Boomplaas A, the scores of all species on these levels were divided by ten in an
attempt to bring these scores within the general range. Similarly, the scores of
species on level 5 at Byneskranskop 1 were divided by five, and an experimen-
tal analysis for Die Kelders 1 M.S.A. samples was carried out with the scores
on levels 1, 2, and 12 to 14 multiplied by ten. More pervasive variability, on the
other hand, requires more general treatment for its reduction. In the present
study it was decided, after some experimentation, that log, ) transformation
seemed to provide the best solution. Although there was still considerable
variation, it was felt that any further transformation would destroy meaningful
variability in the samples. The use of percentages may appear to solve the
problem since the sample size is thereby standardized and, consequently, the
mean will be the same. However, in the present work this advantage was
militated against by the fact that the standard deviations and, therefore, the
coefficients of variation, tended to be considerably larger than in the logy,
transformation. Moreover, the use of percentages has the effect of increasing
negative correlations because as one category increases one or more others
must be reduced in compensation. Although Doran & Hodson (1975: 145)
mention that this effect is likely to diminish as the number of categories
increases, they note that the detailed effects are not known. In conclusion,
whilst it is probable that the matrices achieved by log;,) transformation are not
ideal, from an empirical point of view the output from the analyses appears to
be logically interpretable in terms of the research at hand. Moreover, experi-

939

_—_

ANNALS OF THE SOUTH AFRICAN MUSEUM

ments to reduce the standard deviations did not substantially alter the results in
most cases.

The analyses were carried out on a Univac 1110 computer at the University
of Stellenbosch, using the subprogramme Factor of the SPSS: statistical package
for the social sciences (Nie et al. 1975). There are various options open to the
user of this programme which are explained by Kim (1975) and which pertain
principally to the method of factoring and the subsequent method of rotation to
the final solution. On the advice of R. G. Klein (1975 pers. comm.), the
matrices were originally submitted to principal factoring without iteration
(PA1) and the resultant principal components were rotated orthogonally by the
Varimax method, which centres on simplifying the columns of the factor matrix
(Kim 1975: 485). These options have been used with interesting results by Klein
(1976, 1977) on macromammalian faunas from Klasies River Mouth and
Border Cave. It was subsequently decided, however, to investigate further
options. Consequently, the method of factoring was changed to principal
factoring with iteration (PA2) and the method of rotation to Oblique. PA2
factoring was suggested by the SPSS manual (Kim 1975) as the most useful
general purpose method. Oblique rotation was chosen because, by allowing
that the factors may be correlated, it is intrinsically more likely to fit the
situation in nature (Cattell 19655). It is also consequently more flexible in
achieving the best possible clustering of variables, particularly where moderate
or high correlations are shown to exist between factors. This means, in effect,
that the higher the correlation between the factors the greater will be the
difference between the Oblique and Varimax solutions. Moreover. the Oblique
solution provides more detailed information than the Varimax solution. This
is because the factor-pattern and factor-structure matrices, which are com-
bined in the Varimax solution, are separate in the Oblique solution. The
factor-pattern matrix is of particular importance for the present study because it
distinguishes clusters of variables. Thus, not only are the clusters better defined
by Oblique rotation, they are also more clearly represented in the factor-
pattern matrix. Here the position of the variable in a cluster is indicated by its
loading; the higher the loading the more central the variable is to the pattern
represented by the factor. The loadings in this matrix more closely correspond
to regression coefficients while those in the factor-structure matrix are correla-
tion coefficients as they are in the Varimax solution. The structure matrix does
not distinguish clusters of variables and, as such, is not of importance for
present purposes. Since Rummel (1967: 467) makes the point that it is permiss-
ible very roughly to interpret the loadings of the pattern matrix as correlations,
it is possible in general terms to compare the results of the Varimax rotation
with the Oblique factor-pattern matrix.

The data which were submitted to analysis consisted of the counts for the
microfauna from different levels at several archaeological sites which have been
described above. For the purposes of analysis the microfaunas were restricted
to the species that were best represented, for the reasons given above. In the

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS ZOD

case of the analyses concerning more than one site, they were further restricted
to species occurring in adequate numbers at all the sites concerned. Grouping
of levels in various sites was kept to a minimum so as not to mask or distort any
inherent evidence of change. Basically there are two ways of considering the
data. The species may be grouped according to fluctuations from level to level
in their contribution to the fauna. The alternative is to group the levels
according to the composition of the microfauna of that level. In terms of factor
analysis the former constitutes an R-mode analysis and the latter a Q-mode
analysis. However, since the programme handled only R-mode analysis, the
data matrix had to be transposed manually before it could be analysed. In
terms of the research in hand, grouping of the species, that is, having the
species as variables, allowed the identification of different environmental pat-
terns. It was possible, for instance, to identify a glacial pattern which involved
extensive grass cover among other things. Grouping of the levels, or using the
levels as variables, allowed, in effect, the previously established environmental
patterns to be assigned to specific periods within the general framework.

For the purposes of interpretation one recognized restriction was accepted.
This allows that only solutions containing factors with an eigenvalue of 1 and
over were selected for rotation and, therefore, interpretation. Since the eigen-
value is a measure of the relative amount of variance explained by a particular
factor, this ensures that only the most important factors are considered. In the
present context a high loading is taken as being one of 0,71 and above. This
represents 50 per cent and above of the total variance explained since the
percentage is the square of the loading. High loadings on the same factor
suggest that variables are varying together in a meaningful way (Klein 1981).
Moderate loadings are defined as being those from 0,40 to 0,70 inclusive, with
low loadings being those below 0,40. These two categories are arbitrary, for use
in the general description, and are not claimed to have any mathematical
significance.

A series of analyses was performed, first for individual sites and then for
various combinations of sites. Individual analyses were performed for the sites
with good sequences, namely Boomplaas A, Byneskranskop 1, and Die Kelders
1 M.S.A., in order to establish the internal patterns of variation within the
sites. Thereafter, a coastal analysis was performed, including the Byneskrans-
kop 1 and 2 samples as well as the Die Kelders 1 M.S.A. and L.S.A. level-12
samples. Only the level-12 sample from the L.S.A. levels was sufficiently large
to be included in the analysis. In effect, grouping these samples from one small
area produces artificially a long sequence comparable to that from one site such
as Boomplaas A. The chances of locating possible cycles of change are clearly
enhanced in a longer sequence so that it is advantageous to create one, where
possible, where it does not exist naturally. The purpose of two further analyses
was to test the amount of comparable change discernible in contemporary
sequences at a distance from each other. Thus, an Upper Pleistocene analysis
was performed using samples from the lower half of the Boomplaas A

234 ANNALS OF THE SOUTH AFRICAN MUSEUM

sequence, the Die Kelders 1 M.S.A. levels, Klasies River Mouth 1A levels 15
and 32, and Nelson Bay Cave levels YSL and YGL. To these were added the
modern samples in order to check for comparison with the present situation at
the same time. An additional aim of this analysis was to establish whether there
was any basis for site correlation through comparison of changing micromam-
malian composition, since radiometric dates are not available for much of the
material. Finally, a Holocene analysis was conducted using the Byneskranskop
1 material and that from the contemporary levels at Boomplaas A. Again the
modern samples were included, for the same reason as before. In all cases
where analyses of multiple samples were conducted, the data base was re-
stricted to those species that occurred in adequate quantities at all sites
involved. It was hoped in this way to remove local factors from the comparison.
In the case of the coastal analysis, a check was effected by running the analysis
with and without Steatomys krebsi, a species present in large numbers in the
Holocene but absent from the Upper Pleistocene. It was discovered that,
although constituting the most obvious difference, it was by no means the sole
distinguishing feature. In general, however, it was felt that it was preferable
merely to omit such species.

PROCESSES FOR INTERPRETING ANALYSES

Interpretation of analyses comprises three parts. The first of these entails
the establishment of the common denominator of the variables (species or
levels) loading highly on each factor. In the case of the species, this will be the
pattern of changing proportions of the individual species in the community.
Thus, species occurring in greater numbers in the Upper Pleistocene will load
on a different factor from those occurring in greater numbers during the
Holocene. The first factor will, therefore, represent Upper Pleistocene condi-
tions and the second Holocene conditions. In the levels analyses, the situation
will be reversed and levels with similar faunal composition will be grouped
together. Thus different factors will be characterized by high, or low, propor-
tions of particular species.

The second stage in the procedure is to interpret the meaning of the
factors. In the species analyses, this will entail translating the species that load
highly on each factor into their environmental equivalent. Thus, having estab-
lished that a factor represents Upper Pleistocene conditions, one can then
proceed to suggest what those conditions comprised. The levels analyses
provide, in effect, a cross-check for the interpretations, since the same picture
is being viewed from a different angle. The two types of analysis are also
complementary; it is possible to establish both which element (species = en-
vironmental equivalent) is most important during a given period and also more
precisely when during that period conditions thus indicated would have per-
tained in their purest form. Naturally, interpreting only the variables that load
highly will provide only a partial interpretation of the meaning of the factors.
This will, however, establish the general patterns, which is the aim at this

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 235

preliminary stage. Certainly Klein (1976) has produced eminently worthwhile
results in this manner and it is doubtful whether there is sufficient information
currently available to make an attempt at a more detailed interpretation
appropriate at this time.

The final stage in the interpretation involves employing the information
acquired thus far to construct a history of past environmental change. To this
end, the species analyses, together with the visual aids (Figs 8-12), are used to
show the general trends of climatic and vegetational change. The levels
analyses, together with the relevant visual aids (Figs 9-13), delineate and
characterize periods of similar climate and vegetation. Beyond this, lesser
fluctuations within periods can be ascertained by examining changes in the
faunal composition in the different levels.

MENSURATION AND CALCULATION OF INDICES

Variation in mean individual size and in species diversity was investigated.
Calculations concerning the former were all based on a series of measurements
taken from individual specimens. These measurements had, for the most part,
to be determined by the part of the specimen that was most frequently
preserved. For this reason the measurements might not have been those most
likely to reflect change in the overall mass, although, taken together, they may
perhaps be indicative of such changes. Differences between populations, as well
as any possible trends, were examined in order to establish the potential of this
line of investigation. Changes in the patterns of community structure were also
examined in order to establish whether or not there was any correlation with
known climatic changes. General diversity, which includes both species richness
and species equitability, provides a basic pattern. Splitting of the general index
into its component parts should give some indication as to which is the more
important element.

Mean individual size

Measurements were taken on mandibles and/or maxillae of individuals
belonging to five species. Of these Crocidura flavescens and Myosorex varius
were chosen because the former shows some indication of a positive reaction to
Bergmann’s Rule, while the latter may well exhibit a negative response
(J. Meester 1978 pers. comm.). It was anticipated, therefore, that these species
might provide some evidence of temperature changes in the past. Cryptomys
hottentotus was chosen because it is known to vary in size through its present
range (De Graaff 1981) and because Spalax ehrenbergi, which is also a
mole-rat, is known to vary in size in Israel according to the annual rainfall
(Tchernov 1968). Tatera afra and Aethomys namaquensis provided some ade-
quate samples for measuring and were therefore examined without any prior
expectations. All measurements, which were taken with Helios needlepoint
dial calipers, were expressed in millimetres and calculated to the first decimal
place.

236 ANNALS OF THE SOUTH AFRICAN MUSEUM

In Crocidura flavescens up to five mandibular measurements were taken
where possible. These are as follows:

A. height of ascending ramus, being the distance between the superior
and inferior surfaces of the ascending ramus, measured perpendicular
to the line of the jaw (see Fig. 7);

B. depth of mandible, being the distance between the superior and
inferior borders of the body of the mandible, measured between M,
and Mj in the lingual aspect and perpendicular to the line of the jaw;

C. length of M,_3, being the distance from the posterior surface of M,
to the anterior surface of M,;, measured along the line of the jaw;

D. length of the lower tooth row (L.T.R.), being the distance from the
posterior surface of M; to the tip of I,, measured along the line of the
jaw;

E. length of mandible plus incisor (M+I), being the distance from the
most posterior point of the articular condyle of the lower jaw to the
tip of I,, measured along the line of the jaw (see Fig. 7).

Parameters D and E were included because they are standard
measurements (Meester 1963:5). However, because these parameters could be
taken too infrequently to allow reliable statistical comparisons to be made,
parameter C was taken as an alternative measurement. Parameters A and B
were taken because it was thought that they might give an indication of shape,
either alone or in conjunction with a length measurement. Tchernov (1968)
used depth of mandible for this purpose.

In Myosorex varius four measurements were taken where possible. These
are as follows:
height of ascending ramus, as in Crocidura flavescens;
length of mandible plus incisor (M+1), as in C. flavescens;
length of lower tooth row (L.T.R.), as in C. flavescens;
length of Py to M3, being the distance from the posterior surface of
M; to the anterior surface of P,, measured along the line of the jaw.

OOS

For Cryptomys hottentotus five measurements were taken, where possible,
on the mandible. These are adaptations of the measurements taken by Tcher-
nov (1968) on Spalax ehrenbergi and represent an attempt to provide data for
estimating size. The parameters are as follows:

A. alveolar length, being the distance between the posterior border of
the alveolus of M,; and the anterior border of the alveolus of Pu,
measured along the alveolar row;

B. M; to symphysis, being the distance between the posterior border of
the alveolus of M; and the posterior border of the symphysis;

C. maximum depth of attachment of the ascending ramus, being the
distance between the superior and inferior borders of the ascending
ramus where it attaches to the body of the mandible in the region of
M,., measured perpendicular to the line of the jaw;

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 237

D. foramen mentale to alveolar margin, being the distance between the
foramen mentale and the superior border of the body of the mandible
at Pa:

E. depth of mandible, being the distance between the superior and
inferior borders of the body of the mandible at M,, measured perpen-
dicular to the line of the jaw in the lingual aspect.

In Tatera afra it proved possible to measure only teeth. Here the lengths ot
M,, M._2>, M!’ and M’* were taken. In each case the distance was
measured from the posterior to the anterior surfaces of the teeth in question,
measured along the line of the tooth row.

For Aethomys namaquensis the lengths of M,_3, M,2, M and
M!'~? were measured, using the same definitions as for T. afra. In addition,
the depth of the mandible was also measured, using the same definitions as
those given for C. flavescens.

In all cases the arithmetic mean (X), standard deviation (s) and 95 per cent
confidence interval for the mean were calculated as a basis for estimating
differences between populations. The coefficient of variability, which is given as
V = 100s by Simpson et al. (1960: 90), was calculated for selected samples. In
the present case the coefficient was used in order to discover which
measurements were most homogeneous and, therefore, potentially most useful
in distinguishing populations. This is because parameters that exhibit little
internal variation should be more effective in distinguishing differences
between populations than those parameters that exhibit greater internal varia-
tion. Student’s ‘t’ test was used to establish whether or not differences between
populations were statistically significant. Such significant differences may indi-
cate periods of accelerated change if a short time elapsed between the accumu-
lation of the two samples. Over a longer period of time the indication would be
of the extent of the change involved.

=

Patterns of community structure

The Shannon index of general diversity, as given by Odum (1971: 144),
was employed to establish the basic character of the communities represented
by the samples from the different levels in archaeological sites and from
modern owl roosts. This index is calculated from the equation H = —=P, log P,
where P; = ni/N = importance probability for each species. The base of the
logarithm is immaterial but in the present work natural logarithms (log.) were
used. As was mentioned above, this index takes into account both aspects of
diversity, namely species richness and evenness of importance of each species.
Odum (1971: 149) points out that this index is very useful for making compari-
sons because it is reasonably independent of sample size, which means, in
effect, that fewer samples are needed to obtain a reliable index. In the present
case this is a definite recommendation because sample size varies quite consid-
erably and some samples are not very large.

238 ANNALS OF THE SOUTH AFRICAN MUSEUM

It may, however, be useful to distinguish which element of diversity is the
more important, especially in an exploratory exercise. Either the number of
species or the evenness of representation of the species may be varying. The
first index of species richness given by Odum (1971: 144) was employed. In this
index d = oN where S = number of species, N = number of individuals. The
index of evenness is e = os which is, in effect, the converse of the index of
dominance c = X(ni/N)*, where the symbols are as given above. In practice it
may not be necessary to distinguish the two components since there is a
tendency towards both greater species richness (Krebs 1972: 509) and equit-
ability (Odum 1971: 144) in the tropics relative to temperate and polar regions.
This would suggest, in the present context, that a change in either may indicate
a response to changing climate. Equally, it may suggest that both parameters
will tend to co-vary. On the other hand, Odum (1971: 145) notes that domi-
nance is concentrated in fewer species where physical conditions are extreme.
This suggests that dominance may prove an independently useful index of
environmental change.

THE MODERN DATA

The basic assumption of palaeoecology is that the ecology of the fossils
may be inferred from what is known about equivalent or related living species
(Odum 1971: 159). It cannot be otherwise or it would be impossible to proceed.
In the present case the propriety of basing research on this assumption is
reinforced by the fact that all the species concerned are extant. It follows that it
is first necessary to establish as precisely as possible the factors governing
distribution and relative abundance of the individual species. In the first
instance their ecological distribution will indicate the conditions which they
favour and of which they may be said to be indicative. Generally speaking, it
would seem that small mammal distribution is influenced by vegetation type.
Correlation with other factors is likely to be coincidental, except in so far as
those factors will probably be affecting the vegetation upon which the animals
are dependent. The principal exception will be the influence which the nature
of the substrate will exert on the distribution of burrowing species (Bigalke
1978: 1005).

Proportions of species in the modern samples may be correlated with
suitable habitat present in the expected area of the owl’s hunting territory. This
should provide some basis for comparison of the samples with those from
archaeological sites.

ECOLOGICAL DISTRIBUTION

The available information concerning the ecological distribution of the
individual species forms the basis for interpretation in the present study. Data
regarding the factors that apparently influence distribution have been assem-
bled from the literature and are given below. The species are listed in
taxonomic order (see Table 8).

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS

TABLE 8

Hi: 8 os)

Taxonomic list of species encountered, together with their English common names.

Order Family

Genus, species

English common name

INSECTIVORA — Chrysochloridae

Macroscelididae
Soricidae

CHIROPTERA Nycteridae

Rhinolophidae

Vespertilionidae

RODENTIA Bathyergidae

Muridae

Cricetidae

Muscardinidae

Chrysochloris asiatica
Chlorotalpa duthiae
Amblysomus hottentotus
Elephantulus edwardi
Myosorex varius
Suncus varilla
Crocidura cyanea
Crocidura flavescens
Crocidura sp.

Nycteris thebaica
Rhinolophus clivosus
Rhinolophus capensis
Myotis lesueuri

M yotis tricolor
Eptesicus hottentotus
Eptesicus capensis
Miniopterus schreibersi
Cryptomys hottentotus
Georychus capensis
Acomys subspinosus
Aethomys namaquensis
Dasymys incomtus
Mus minutoides
Praomys natalensis
Praomys verreauxi
Rattus rattus
Rhabdomys pumilio
Mystromys albicaudatus
Tatera afra

Gerbillurus paeba
Dendromus melanotis
Dendromus mesomelas
Steatomys krebsi
Saccostomus campestris
Otomys laminatus
Otomys saundersae
Otomys irroratus
Otomys unisulcatus
Graphiurus ocularis

Nomenclature according to Meester & Setzer (1971).

Insectivora

Cape golden mole
Duthie’s golden mole
Hottentot golden mole
Cape elephant-shrew
forest-shrew

dwarf-shrew

reddish-grey musk-shrew
red musk-shrew
musk-shrew

Egyptian slit-faced bat
Geoffroy’s horseshoe-bat
Cape horseshoe-bat
Lesueur’s wing-gland bat
Cape hairy bat
long-tailed house-bat
Cape serotine

Schreiber’s long-fingered bat
common mole-rat

Cape mole-rat

Cape spiny mouse
Namaqua rock-rat
shaggy swamp-rat

dwarf mouse
multimammate mouse
Verreaux’s mouse

black rat

striped field-mouse
white-tailed rat

Cape gerbil

South African pygmy gerbil
grey pygmy climbing mouse
chestnut climbing mouse
fat mouse

pouched mouse

laminate vlei-rat
Saunders’ vlei-rat

vlei-rat

bush Karoo rat

Cape dormouse

Chrysochloris asiatica lives in cultivated and uncultivated ground (Short-
ridge 1942: 32). It occurs not only in sandy soil under Karoo scrub (Shortridge
1942: 32; Rautenbach 1971: 138), but also in alluvial soil under open grasslands

(Shortridge 1942: 75).

Chlorotalpa spp. are burrowing species generally to be found in sheltered
mountain valleys or forests where they can burrow in the peaty soil (Roberts
1951: 108). Pienaar (1964: 16) notes, however, that they occur in sandy soil.

240 ANNALS OF THE SOUTH AFRICAN MUSEUM

C. sclateri occurs in areas which experience severe frost and with an annual
rainfall of 700 mm (Lynch 1975: 137).

Amblysomus hottentotus burrows extensively in soft but not necessarily sandy
soil, generally in open ground with plenty of grass cover (Roberts 1951: 189).

Elephantulus edwardi appears to favour rocky outcrops on grassland (Cor-
bet & Hanks 1968: 97), although Roberts (1951: 29) quotes A. Smith as saying
that the type came from ‘a locality bearing little or no vegetation, except a few
dwarf shrubs’.

Myosorex varius frequents dense undergrowth near streams or swamp
vegetation (Shortridge 1934: 37; 1942: 35) in a fairly moist and often montane
environment (Brain & Meester 1964: 337) and is often to be found under fallen
trees (Thomas & Schwann 1905a: 131, 264). It is also to be found on grassy
slopes (Rautenbach 1976: 134). Although it usually exists in wetter areas, it can
be found in drier places where dense vegetation or frequent mists presumably
counteract the lack of precipitation (Meester 1958: 327). It was found in a wide
variety of habitats by Bond et al. (1980: 41).

Suncus varilla: at least one subspecies is said to inhabit deserted termite
mounds (Roberts 1951: 44).

Crocidura cyanea: the two subspecies are almost entirely restricted to the
above and below 500 mm rainfall zones; C. c. infumata inhabits the former in a
wide range of vegetation while C. c. cyanea occupies the latter in steppe and
fynbos vegetation (Meester 1963: 54, 58). Shortridge (1942: 53, 78) records the
fact that C. c. cyanea favours rocky, stony country while Rautenbach (1976:
135) notes that what is presumably C. c. infumata occurs in dense grass along a
stream-edge.

Crocidura flavescens is generally stated to prefer dense undergrowth (Tho-
mas & Schwann 1905a: 130), normally in rocky or broken country (Meester
1962: 77; Sclater 1900-1: 162). It also occurs in and around cultivated land
(Thomas & Schwann 19055: 264). At one time it was thought to be restricted to
the above 750 mm rainfall zone (Meester 1963: 39) but it now seems that it is
able to survive in drier areas where there is sufficient ground cover or mists
(Swanepoel 1975: 118). It does not seem to be restricted to any particular type
of vegetation (Meester 1963: 77).

Rodentia

Cryptomys hottentotus is a burrowing species which occurs both on moun-
tain plateaux and in low-lying areas (Shortridge 1942: 97; Rautenbach 1976:
136), where it may be found on and between dunes (De Graaff & Nel 1970:
180; Rautenbach 1971: 141). It seems to avoid severe frost conditions (Lynch
1975: 137). It apparently prefers more open grassy vegetation, but is also found
in fynbos (Stuart n.d.).

Georychus capensis burrows extensively in sandy or other loose soil,
whether cultivated or not, usually in areas bordering pans and the like (Roberts
1951: 382; Sclater 1900-1: 76).

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 241

Acomys subspinosus is generally said to prefer rocky situations (Roberts
1951: 465; Davis 1962: 62), and Bond et al. (1980) correlated its occurrence
with rocks 1 m in diameter. Shortridge (1942: 92) further noted that it occurred
on heathy slopes and plateaux. Bond et al. (1980: 39) found that it favoured
high-altitude areas with dense phytomass, especially below 60 cm. On the coast
it appears attached to thick undergrowth at the edge of forest (Thomas &
Schwann 1906a: 164) and to forests and woodlands (Andrews et al. 1975: 24).

Aethomys namaquensis is generally described as frequenting rocky hillsides
(Sclater 1900-1: 43; Roberts 1951: 482). Bond et al. (1980: 40) have found its
distribution strongly correlated with the occurrence of stones up to 13 cm in
diameter, but have suggested that this may be fortuitous. It is sometimes to be
found in flatter areas (Shortridge 1934: 295; De Graaff 1974: 178). Although
this is usually in areas where there are rocks (Thomas & Schwann 1904: 179),
it has been recorded from river-banks and even dune areas (Rautenbach & Nel
1975: 197). It appears to prefer scrub (Hanney 1965: 596; Bond et al. 1980), but
Ansell (1960: 92) records it as living in woodland. Stuart (n.d.) also noted that,
although it occurred in indigenous forest, it preferred short to medium fynbos
at higher altitudes. Bond et al. (1980: 40) have found its occurrence correlated
with sparse (less than 75 per cent total shrub cover) succulents. It was also
restricted to areas of less than 10 per cent microphyllous shrubs.

Dasymys incomtus is generally stated to inhabit dense vegetation on the
banks of streams, edges of swamps, reed-beds and the like (Roberts 1951: 494;
Ansell 1960: 97). Thomas & Schwann (1905b: 270) state, however, that it is not
necessarily found close to water and Hanney (1965: 617) found it up to 1 km
from streams at high altitudes but within 20 m of water at low altitudes. It is
commonly found in grasslands (Pienaar 1964: 22; Davis 1974: 152), and Davis
(1962: 62) further states that it is typical of savanna and montane savanna.

Mus minutoides is found in all manner of habitats from dunefields (Rauten-
bach & Nel 1975: 197) to woodland (Vesey-Fitzgerald 1964: 67) and even forest
(Delaney & Neal 1966: 331), on hillsides and on the flat (Thomas & Schwann
1905a: 136). Vesey-Fitzgerald (1966: 117) notes that it occurs where vegetation
is usually in decline or secondary.

Praomys natalensis inhabits a diversity of environments including forest,
grassland and rocks (Vesey-Fitzgerald 1966: 115), and also cultivated land and
houses (Delaney & Neal 1966: 326; Pienaar 1964: 22). It appears, however, to
avoid extremes such as desert or semi-desert (De Meneses Cabral 1966: 194;
Coetzee 1975: 637) and rain forest (Hubbard 1972: 436). It perhaps has a
preference for grassland (Sheppe 1973: 173; Andrews et al. 1975: 24) and seems
to prefer the vicinity of water (Shortridge 1934: 301).

Praomys verreauxi is said to live among the scrub and bush on rocky
hillsides (Roberts 1951: 470; Shortridge 1942: 93) and also in the forest
(Roberts 1951: 470; Thomas & Schwann 1906a: 164). There is some doubt as to
whether it ventures onto the plains, Shortridge (1942: 93) maintaining that it
does not and Thomas & Schwann (1906a: 164) stating that it is found in swamps

242 ANNALS OF THE SOUTH AFRICAN MUSEUM

and grass-filled hollows in open country. Bond ef al. (1980: 41) found this
species Only at mid and high altitudes.

Rhabdomys pumilio is generally stated to prefer dense vegetation, often on
stream banks or in dry river-beds (Roberts 1951: 497; Shortridge 1934: 278),
even in otherwise very dry areas such as the Namib Desert (Coetzee 1969: 30).
The type of vegetation is obviously less important than the amount of cover
afforded to this diurnal species since it is found in scrub (Nel & Pretorius 1971:
107), bracken (Hanney 1965: 606), in cultivated land (Shortridge 1942: 607)
and even in the Tsitsikama Forest (Smuts 1832: 37), but is may perhaps have a
preference for grassland (Smithers 1971: 292; Vesey-Fitzgerald 1964: 66). Bond
et al. (1980: 38) found that its presence was correlated with continuous or
extensive ‘grass’ patches, ‘grass’ including Poaceae, Cyperaceae and Restio-
naceae as well as Gramineae for the purposes of their classification. Brooks
(1974) found that it may prefer grassland and vlei at different times of the year,
or that it may be forced into less favourable areas by a periodic abundance of a
narrow-niche species such as Otomys irroratus. Although it seems to have a
preference for damp areas (Hanney 1965: 606) which afford immunity from the
effects of fire (Vesey-Fitzgerald 1966: 117), it is apparently not restricted to
them (Rautenbach 1971: 141).

Mystromys albicaudatus appears to be restricted to grassland, including
high-altitude montane grassland where it is an indicator of the transition from
savanna grasslands to woodlands (Davis 1962: 64). Roberts (1951: 436) records
that it was found in meadowland along a river bank.

Tatera afra prefers to burrow in sandy coastal plains (Shortridge 1942: 89).
It is to be found in open ground with short grass (Sclater 1900-1: 21), under
bushes and in cultivated land (Roberts 1951: 409).

Gerbillurus paeba lives in sandy areas where the vegetation cover is not
dense, in desert and semi-desert areas (Shortridge 1942: 53; Coetzee 1969: 29).

Dendromus melanotis is generally said to live in grassland, whether near
trees or not (Shortridge 1942: 91; Davis 1962: 63), both near swampy country
and in dry areas (Smithers 1971: 319).

Dendromus mesomelas appears, in contrast to the previous species, to be
found more frequently in bush or woodland (Davis 1962: 63; Smithers 1971:
613). However, it also occurs in tall grass and scrub (Ansell & Ansell 1973: 61)
and Shortridge (1934: 243) states that it favours damp grass, although it is
seldom far from trees and bushes. Kingdon (1974: 532) states that although it
has a preference for wet habitats, it will occupy the driest most terrestrial
habitat when in competition with other species of Dendromus.

Steatomys krebsi apparently lives in more or less open subcoastal grassland
and bush (Shortridge 1942: 91).

Saccostomus campestris is generally said to inhabit woodland (Pienaar
1964: 23; Delaney 1972: 11) or at least grassland with scattered trees and shrubs
(Sheppe 1973: 184), but not open grassland, at least not in South Africa (Davis
1962: 63). De Graaff (1981) notes wide tolerance in this species. It is also

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 243

attracted to cultivated land (Vesey-Fitzgerald 1964: 67; Delaney & Neal 1966:
337). It seems to favour drier country (Hanney 1965: 613; Delaney & Neal
1966: 337), but within reach of water (Shortridge 1942: 59; Thomas & Schwann
1905b: 269). It seems to avoid areas of severe frost (Lynch 1975: 137).

Otomys laminatus is recorded from moist submontane and coastal savanna
(Davis 1962: 63; 1974: 152), while Thomas & Schwann (1905b: 268) state that it
occurs among rocks on hillsides and cliffs.

Otomys saundersae is said to share a preference for high altitude and
montane grasslands with O. irroratus and O. sloggetti (Davis 1962: 64), while it
is recorded by Shortridge (1942: 89) as inhabiting belts of dry rushes in heathy
country on high mountain slopes. According to J. U. M. Jarvis (1978 pers.
comm.), it prefers rockier situations with more open vegetation than does
O. irroratus.

Otomys irroratus is generally stated to inhabit dense vegetation on the
banks of streams and edges of swamps (Roberts 1951: 420; Delaney 1972: 120).
It is, however, sometimes found at some distance from water (Thomas &
Schwann 1905a: 135; 1905b: 266; 1906b: 589; Shortridge 1934: 239) and Bond et
al. (1980: 39) have found it in a range of habitats, none near water. The
occurrence of this species was, instead, correlated with the microphyll layer
between 1,0 m and 2,5 m and the total microphyll layer, and its presence was
apparently restricted to areas with 75+ per cent shrub cover (Bond et al. 1980:
40). Davis (1973) confirmed the occurrence of this species in an area of dense
basal and canopy cover.

Otomys unisulcatus is generally said to inhabit drier areas among the scrub
vegetation of the Karoo, and rocks on the lower slopes of hills and mountains
(Roberts 1951: 427; Shortridge 1934: 244). Nel & Pretorius (1971: 107),
however, recorded it from riverine scrub and forest.

Graphiurus ocularis is said to be basically a rock dweller, although to some
extent it is found in trees (Roberts 1951: 369; Shortridge 1942: 51). It seems to
prefer drier areas with Karoo or ‘karoid’ vegetation (Shortridge 1934: 216).

THE BASIS FOR INTERPRETATION

The data given above require analysing in such a way as to be useful for
interpreting environmental conditions. This involves separating data relative to
individual factors or aspects of the habitat. For present purposes, data relating
to abiotic factors such as horizontal distribution, especially in areas of high
relief, and correlation with different ground-surface and substrate types will
restrict application of data forthcoming from biotic factors. In practical terms
this means that, for instance, the presence of Aethomys namaquensis in a
sample will suggest scrub, probably semi-arid, which is specifically most likely
to be present on the lower slopes of the hills or mountains. Of the biotic
factors, the life form and density of the vegetation may be treated separately. It
is possible to establish some sort of correlation between annual rainfall and the
incidence of various species, based on geographical distribution, which may

244 ANNALS OF THE SOUTH AFRICAN MUSEUM

have some broad predictive value. Being homoiothermic, small mammals may
be expected to exhibit broad temperature tolerance. Moreover, they often exist
in microhabitats which are very different from the macrohabitat. For both
reasons it is not considered appropriate to attempt to correlate temperature
with small mammal distribution.

In the present context, establishment of horizontal distribution largely
consists of determining which species occur on hillsides and mountain slopes.
The majority of species occurs on flat ground which, depending on the area
being studied, may be the coastal foreland, valley floors, or, possibly, mountain
plateaux. Of the six species which Bond et al. (1980) trapped on the slopes of
the Swartberg Mountains, Aethomys namaquensis occurred only on the lower
slopes while Acomys subspinosus and Praomys verreauxi were apparently
restricted to the higher slopes. Otomys irroratus, Rhabdomys pumilio, and
Myosorex varius occurred at all altitudes. Otomys saundersae 1s said to occur
on high mountain slopes (Shortridge 1942: 89) but Otomys laminatus may be
restricted to lower hillsides or, at least, lower elevations (Davis 1962: 63;
Thomas & Schwann 1905b: 268). Otomys unisulcatus is apparently also res-
tricted to the lowest slopes, as well as the low ground (Shortridge 1934: 244).
Mus minutoides has been found on rocky hillsides (Thomas & Schwann 1905a:
136) as well as in a wide variety of other situations. In general, species that
occur on hill or mountain slopes appear to be those that are to be found on
rocky ground (see Table 9). This being the case, a further group of three
species may also occur on hillsides, although this has not been specified.
Graphiurus ocularis is known to be a montane species (J. U. M. Jarvis 1978
pers. comm.) that frequents rocks (Roberts 1951: 369) and presumably, there-
fore, mountain slopes. Crocidura flavescens is apparently restricted to rocky,

TABLE 9

Correlation between distribution of various species and abiotic factors.

Al: upper slopes A2: lower slopes A3: unspecified slopes
A, subspinosus A. namaquensis M. varius
P. verreauxi O. laminatus E. edwardi
O. saundersae O. unisulcatus R. pumilio
O. irroratus
G. ocularis

B1: rocky ground _ B2: loose sand/soil B3: sandy plains

E. edwardi C. asiatica M. minutoides
C. flavescens Chlorotalpa spp. S. krebsi

A, subspinosus A. hottentotus S. campestris
A. namaquensis C. hottentotus

P. verreauxi G. capensis

O. laminatus T. afra

O. saundersae G. paeba

G. ocularis

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 245

TABLE 10

Correlation between distribution of various species and vegetation
type and density.

Al: grass A2: scrub A3: woodland/forest
A. hottentotus E. edwardi Chlorotalpa spp.

M. albicaudatus G. capensis

T. afra A. subspinosus

D. melanotis A, namaquensis

G. paeba P. verreauxi

S. krebsi O. unisulcatus

O. laminatus G. ocularis

B1: dense vegetation B2:spasre vegetation

ee ———————

M. varius C. asiatica

C. c. infumata E. edwardi

C. flavescens C. hottentotus
A, subspinosus G. capensis

D. incomtus A. namaquensis
R. pumilio G. paeba

D. melanotis O. unisulcatus
O. irroratus G. ocularis

~

mountainous country (Meester 1963: 40), although this may be in valleys;
Sclater (1900-1: 162) mentions wooded ravines. Elephantulus edwardi exhibits
a preference for rocky ground (Shortridge 1942: 29), including high plateaux,
but possibly lower hills rather than steep mountain slopes.

Of the vegetation types, grass and in particular ‘grass’ sensu Bond et al.
(1980) appears to support the greatest number of small mammal species (Table
10). A certain amount of variation is, however, discernible. Some species, such
as Rhabdomys pumilio and Dendromus melanotis, prefer dense grass-cover.
Others, such as Chrysochloris asiatica and Gerbillurus paeba, occur in much
more open grass. Indeed, the latter species is known to decline in years when
the grass cover is increased by good rains (J. A. J. Nel 1978 pers. comm.),
whereas the opposite occurs in R. pumilio (Swanepoel 1975: 123). Mystromys
albicaudatus and Dendromus mesomelas, although they are grassland species,
indicate the presence of trees or bushes in the vicinity (Davis 1962: 63). With
other species, grass is only one of the life forms in which they occur. Of the
species occurring in scrub, it would appear that Aethomys namaquensis and
Graphiurus ocularis are found in both fynbos and semi-arid scrub, although
perhaps more often in the latter. In this context fynbos is taken to mean
mountain fynbos and semi-arid scrub to mean either succulent vegetation with
Karoo affinities or arid fynbos. Woodland or forest, probably riverine or valley
floor, is indicated by three species, Chlorotalpa spp., Dendromus mesomelas,
and Saccostomus campestris. Reed-beds may be suggested by Dasymys incom-
tus and Otomys irroratus, although the latter species in particular is not

246 ANNALS OF THE SOUTH AFRICAN MUSEUM

restricted to such conditions. A remaining group comprises broad-niche species
which occur in a variety of vegetation types.

It is possible to find limited correlation between rainfall (Fig. 1C) and the
geographical distribution of individual species. The range appears, in the main,
to be fairly broad, but some distinction is possible between a proportion of
species occurring in drier areas and those to be found in wetter areas
(Table 11).

The main environmental factors indicated by the individual species are
summarized in Table 12. This information will serve as the basis for interpret-
ing the information forthcoming from both modern and excavated micromam-
malian samples. In the present context it should be noted that flats are taken as
indicating valley floors and moutain plateaux as well as more low lying areas
such as the coastal foreland. |

INTERPRETATION OF COMPARATIVE SAMPLES

There are two major collections of comparative material from the Cango
valley, which are taken together, and two from the coast, at Byneskranskop 2
and Glentyre. Details of the location of the sites and the methods of collection
are given above. Chaline (1972: 273) reconstructed the environment of three
modern localities in France in order to show that it was possible to obtain an
accurate result. Since this point has, therefore, been adequately proved, it is
not necessary to repeat the exercise for this purpose. It is, however, useful to
carry out similar reconstructions for the present material for two other reasons.
The first is that 1t provides a means of checking the environmental indications
that have been assigned to the species. The second is that it may thus be
possible to assess whether there has been a change since the end of the subfossil
record, about 1 500-2 000 B.P. in the present samples, and the present. In fact,
the introduction of agriculture about 300 years ago has caused major, although
quite artificial, changes in the vegetation and it may well not be possible to
compare the present situation with that pertaining in the past. A further
restriction on interpretation is caused by the fact that the precise area hunted
by the owl or owls is not known. For this reason it is not possible to make
quantitative comparisons between proportions of small mammals and those of
the various vegetation types. Such a lack of precision applies equally to the
subfossil material. Some indication of general proportions may, however, be
warranted.

Byneskranskop 2 (BNK2)

Basically, the area around BNK2 can be divided into hills, plains and
riverside. The hills tend to support fynbos, although this may vary from more
restioid to more proteoid depending on the substrate. At the base of the
limestone ridges and along the river, coastal scrub is found. The difference is
that along the river there is a dense under-layer of vegetation, whereas on the
lower slopes of the limestone ridges there tends to be little vegetation beneath

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 247

TABLE 11

Generalized correlation between distribution of various species and rainfall (based on distri-
bution in the southern and south-western Cape).

Up to 400 mm Up to 600 mm 200 mm upwards 400 mm upwards
G. paeba C. asiatica M. varius C. duthiae
O. unisulcatus E. edwardi M. minutoides A. hottentotus

C. c. infumata (500 mm)
C. flavescens

G. capensis

A. subspinosus
D. incomtus

M. albicaudatus
S. krebsi

O. laminatus

O. trroratus

O. saundersae
G. ocularis

P. verreauxi
D. melanotis
D. mesomelas

C. c. cyanea (500 mm)
A. namaquensis
S. campestris (S00 mm)

Taawe 12

Main environmental factors correlated with various small mammal species in the

southern Cape.

Species Vegetation Location Rainfall

C. asiatica fairly open grass or flats up to 600 mm
semi-arid scrub

C. duthiae probably bush or forest flats 400+ mm

A. hottentotus open grassland flats 400+ mm

E. edwardi sparse semi-arid scrub rocky slopes up to 600 mm

M. varius dense vegetation hillsides or flats 200-+- mm

C. c. cyanea grass or fynbos rocky slopes up to 500 mm

C. c. infumata dense vegetation 500+ mm

C. flavescens dense vegetation rocky ground 400+ mm

G. capensis geophytes (fynbos) near pans on flats 400+ mm

C. hottentotus sparse vegetation flats

A, subspinosus dense scrub rocky slopes 400+ mm

A, namaquensis sparse (under 75 percent cover) rocky slopes up to 600 mm
fynbos or semi-arid scrub

D. incomtus dense reeds or grass near water 400+ mm

P. verreauxi fynbos rocky slopes 200+ mm

R. pumilio dense vegetation hillsides or flats

M. albicaudatus grassland flats 400+ mm

T. afra open grasslands flats

G. paeba sparse grassland flats or sandy slopes up to 200 mm

D. melanotis dense grassland flats 200+ mm

D. mesomelas grassland with trees or bushes flats 200+ mm

S. krebsi grassland flats 400+ mm

S. campestris open woodland with grass flats up to 500 mm

O. laminatus grassland rocky slopes 400+ mm

O. saundersae grass or fynbos higher rocky slopes 400+ mm

O. irroratus dense (75+ per cent cover) hillsides flats 400+ mm
grass, reeds or fynbos

O. unisulcatus sparse semi-arid scrub rocky or sandy flats up to 400 mm

lower hillsides
G. ocularis sparse fynbos or semi-arid scrub _ rocky slopes 400+ mm

248 ANNALS OF THE SOUTH AFRICAN MUSEUM

the trees. On the flats, vegetation generally tends to be fairly open; the alien
Acacia species tend to form dense thickets but with little vegetation beneath
them. There are also marshy areas near the rivers (Schweitzer & Wilson 1978).

The micromammalian data (Table 7) can be divided in similar fashion. On
the hillsides Praomys verreauxi and Otomys saundersae indicate scrub (here
fynbos) on the upper slopes. If, as is thought likely, the subspecies of Crocidura
cyanea represented is C. c. cyanea, this would indicate fynbos on the hills.
Otomys laminatus is said to occur in grassland (Davis 1974) but possibly in this
case it is the restioid element that is suggested. Equally, O. saundersae could
represent the restioid element. Aethomys namaquensis indicates that the lower
rocky slopes support relatively sparse scrub vegetation, which Elephantulus
edwardi would tend to confirm. Myosorex varius and Otomys irroratus would
both indicate dense vegetation, possibly proteoid in the latter species if they
occurred on the hillsides. Crocidura flavescens indicates dense vegetation on
rocky ground which is here most likely to be found on the hillsides.

The prevalence of burrowing forms on the plains is entirely consistent with
the loose, sandy nature of the substrate. Georychus capensis suggests the
possibility of one or more pans in the area, possibly the marshy areas near the
rivers. The majority of species occurring on flat ground is indicative of grass-
land and it would seem that the coastal fynbos is largely unrepresented, except
in so far as this includes a fairly high proportion of grasses. Chrysochloris
asiatica, Georychus capensis and Tatera afra also occur in cultivated ground
and their presence is consistent with the fact that much of the area either is or
has been under cultivation. The relatively high proportion of Rhabdomys
pumilio (see Table 7) suggests that this species is, in fact, very well represented
in the area since it could be expected to be under-represented in Tyto alba
pellets. Work by Jarvis & David (n.d.) indicates that R. pumilio has adapted to
the alien Acacia habitat; it is also known to occur in grassland and dense
streamside vegetation. In conjunction, these could help to explain its apparent
abundance near Byneskranskop.

Myosorex varius, Dasymys incomtus and Otomys irroratus are all indicative
of the dense grass or reeds along the banks of the river. Dendromus mesomelas
further suggests the presence of trees in or near the damp vegetation.

It is unfortunate that the habitat preferences of some of the more numer-
ously represented species are either unknown or catholic. Suncus varilla may
inhabit dense vegetation, as do other soricids, but its preferences are not
known. Mus minutoides is very adaptable but there is some indication (D. M.
Avery unpublished data) that it occurs in greater numbers in owl-pellet samples
from areas where agriculture is fairly extensive. It has been reported from fields
and sometimes houses (Sclater 1900-1: 51), and Vesey-Fitzgerald (1966: 117)
notes its occurrence in secondary and declining vegetation. It may, in fact, be
that the presence of relatively large numbers of this species, and perhaps of
Suncus varilla, indicates the occurrence of expanses of more open ground, in
this case possibly fallow fields. This is suggested by the fact that individuals of

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS

TABLE 13

Proportions of species in different habitats near Byneskranskop.

Hillside

E. edwardi sparse vegetation

M. varius* dense vegetation .

C. flavescens dense vegetation .

C. c. cyanea fynbos

A, namaquensis sparse scrub .

P. verreauxi fynbos

O. laminatus ?restioid element

O. saundersae ?restioid element

O. irroratus* dense ?proteoid element
Plains

C. asiatica cultivated, grass .
meeponewiowis grass ... ..
G. capensis pans, cultivated, grass
R. pumilio* grass, Acacia .

M. albicaudatus grass ;

T. afra cultivated, grass .

D. melanotis dense grass

S. krebsi grass

Riverside

M. varius* grass, fallen trees

D. incomtus reeds, grass

R. pumilio* grass, Acacia .
D.mesomelas near trees .

O. irroratus* grass, reeds

Sundry

S. varilla

A, subspinosus
M. minutoides
R. rattus

Nie

* Score divided arbitrarily between two categories.

Number

249

Percentage

0,13
4,78
3,32
1,86
VA he
5,18
0,66
0,66
8,37

27,76

Doe
0,13
6,64
3,85
0,13
e525
pee
4,25

28,69

——--

4,91
0,13
3,85
2,82
8,50

19,52

10,09
0,80
12,08
1,06

24,04

250 ANNALS OF THE SOUTH AFRICAN MUSEUM

both species are small and are thus unlikely to be hunted preferentially in dense
vegetation. This would be a similar situation to that found by Southern (1954)
for Strix aluco (tawny owl) in England. Rattus rattus is a purely commensal
species whose presence is attributable to human settlement in the area. Acomys
subspinosus suggests dense vegetation, but whether on the edges of the scrub or
on the hillsides is not known since it apparently behaves differently in coastal
and inland situations.

Table 13 shows that the hillsides and the plains are approximately equally
represented, with just under 30 per cent each of the total sample. The dense
riverside vegetation is rather less well represented, with nearly 20 per cent of
the total. The plains category would thus appear to be under-represented, but
is may be that the majority of species included under the heading of ‘Sundry’
should be placed with the plains species, which would increase the score
considerably. It may be that at Byneskranskop the majority of Otomys irroratus
and Myosorex varius come from the riverside habitat, but it is not possible to
assess this at the moment. The fact that the proportions of the different
topographic units may not be very accurately represented in the micromamma-
lian evidence need not be important. Except along the modern coastline
topographic features have not changed during the period under discussion so
that there is little merit in attempting to analyse the evidence for the physical
setting. It is the vegetation that is of principal interest and this is accurately
represented within the various topographic units. Examination of contemporary
changes of vegetation within the different units will provide a fairly detailed
overall picture of the situation in the area adjacent to the site being studied.

Cango Valley (BPB-C and NGA-B)

The Cango valley may be divided basically between the valley floor and the
sides of the valley. In some places where the valley is broader, the proportion
of floor to sides is clearly greater than in others. Thus, at Boomplaas, approxi-
mately one third of the area probably hunted by Tyro alba constitutes valley
floor. At Nooitgedacht, on the other hand, no more than about a quarter of the
area probably hunted is valley floor. Natural and alien vegetation occur in the
vicinity of both sites. At Boomplaas extensive farming has opened up most of
the alluvial valley floor, except along the Grobbelaars River and the water-
furrows down each side of the valley, where larger trees and bushes occur.
Directly below the cave several fields of grass with scattered walnut trees
provide the equivalent of savanna vegetation. On the hillsides Dense Asteroid
Shrubland, Limestone Vegetation and Mixed Bush (Moffett & Deacon 1977)
succeed each other at different altitudes. Of these categories only the Lime-
stone Vegetation has an appreciable grass element. Other vegetation types
occur at higher altitudes but these are thought not to be relevant to the present
study. At Nooitgedacht the succession on the hillsides also includes Dense
Asteroid Shrubland and Limestone Vegetation, but with Closed Woodland in
the bottom of the narrow-sided valley directly below the site. In the wider De

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS Ps il

Hoek valley, into which the Nooitgedacht valley leads, a certain amount of
agriculture has been practised on the alluvial soil. This comprises mostly the
growing of fruit trees, but also involves the provision of pasture for grazing
cattle. Large trees and bushes also occur along the banks of the river.

Table 14 shows that the small-mammal data reflect the greater relative
importance of the hillside at Nooitgedacht compared with Boomplaas. In both

TABLE 14

Proportions of species in different habitats near Boomplaas and Nooitgedacht
in the Cango valley.

Boomplaas B—C Nooitgedacht A-B

No he No. yA
Valley floor
C. duthiae OOUSH, FQNCESE.. = 2 «we we le i 1,20 8 1,00
M. varius* @emse vegetation. . . . . . =. 12121 9,66 98 227
C. hottentotus openvegetation . . kha segs «9 LZ =) 0,63
D. incomtus dense waterside vegetation a — ps 0,25
P. natalensis Grass. Gullivated . . « «§ « «.3 bo 17,80 45 5,63
R. pumilio Gemse vemetatiom. . =: « . »« » 36 4,47 19 2,38
Merearaais ptasS. =. 3. kl lk lt 2 0,16 i! 0,13
D. melanotis dense grass... ET lee ee 3,19 11 1,38
D. mesomelas — grass + trees/bushes oe ees ga dt 1,36 13 1,63
S. krebsi Grass. |. ‘ee hee Se 2 0,16 — —
S. campestris open woodland is grass sd Fe OY se 3%, 8 1,44 5 0,63
O. irroratus* Memeemeceralion. . :. «1 5 « «. T4 5,91 78 9,76

Hillside
E. edwardi Open semi-arid scrub . . . .. 6 0,48 2 0,25
M. varius* Memsevereation. . . . .«. « « Iai 9,66 98 1227
C. c. cyanea Peaeeyebes. «1 el lk Cl! U8 4,63 18 2,23
C. flavescens Gemsewerctation:.. . .) 5 ss 5 0,40 4 0,50
pueseesmimosus’. Gensesctub . .*. : .. . . 23 1,84 43 5,38
momameguensis Open Scrub . . . . ... . . 146 11,65 109 13,64
P. verreauxi PoeOseGense. 5. fs s sow « 2 1,84 69 8,64
O. laminatus TOPE S “a ee or 8 0,64 7 0,88
O. saundersae PRPCINEIOOS .-9t « wk ks ew’ 8 5,43 33 4,13
O. irroratus* Beusoveretmlion.. . . . «= +» « IS 5,83 Ci; 9,64
G. ocularis OOemisense. Ge. «9 6 oe ww 1 0,08 3 0,38
532 42,46 463 57,95
Sundry
S. varilla Mead). cut he Ly es 8 1,84 16 2,00

M. minutoides MEER wile) eet yee ia Ss. 2 al 8,86 55 4,38

ee ee Eten on eyes oni LOS 799

* Score divided arbitrarily between two categories.

-)
nN
i.)

ANNALS OF THE SOUTH AFRICAN MUSEUM

cases, however, the valley floor would appear to be heavily over-represented.
This suggests either that 7. alba has a marked preference for hunting on the
valley floor or that the small-mammal biomass is actually much higher here
than on the valley sides. However, as was pointed out above, disproportionate
representation of the different topographic units need not inhibit accurate
interpretation of the vegetation of a region. Assessment of change will, in any
case, be within, rather than between, such units.

On the valley floor the species reflect the combination of both dense and
more open vegetation. The former is indicated by Myosorex varius, Dasymys
incomtus, Rhabdomys pumilio, Dendromus melanotis and Otomys irroratus.
The majority of the remaining species in this category in Table 6 suggests more
open vegetation. Bush or forest may be indicated by Chlorotalpa duthiae while
more open woodland with grass, perhaps the orchards in this case, is suggested
by Dendromus mesomelas and Saccostomus campestris. Mystromys albicauda-
tus, D. melanotis and Steatomys krebsi all indicate grass, possibly in wheat or
fallow fields. The presence of Praomys natalensis is thought particularly to be
an artefact of cultivation (Avery 1977). It is noticeable that this species is
approximately three times as common at Boomplaas as it is at Nooitgedacht,
which agrees with the greater amount of agricultural activity in the former area.
It is also noticeable that most of the species which inhabit the valley floor away
from the river banks are less well represented at Nooitgedacht. This is presum-
ably an accurate reflection of the relative situation in the two areas.

On the hillsides the main indication is of scrub, which is entirely to be
expected. In some cases this scrub is open (Elephantulus edwardi, Aethomys
namaquensis, Graphiurus ocularis) and in others denser (Acomys subspinosus,
Otomys irroratus). A considerable variety of bush and scrub types is described
by Moffett & Deacon (1977) and this is reflected in the variety of small
mammal species occurring on the hillsides. A certain amount of grass is
suggested by Otomys laminatus and probably also by Crocidura c. cyanea and
Otomys saundersae, and grass is, in fact, a component of the Limestone
Vegetation. In general, the proportions of individual species are similar at both
sites. The evidence would suggest, however, that there is a higher proportion of
dense vegetation at Nooitgedacht than there is at Boomplaas. This is, in fact,
the case, since there is no vegetation at Nooitgedacht comparable to the more
open overgrazed (Moffett & Deacon 1977: 127) scrub on the western side of
the main valley opposite Boomplaas. The very much higher proportions of
Praomys verreauxi at Nooitgedacht may suggest that this species prefers either
denser vegetation or perhaps the rather wetter conditions prevailing on the
eastern side of the Cango valley. The position regarding Otomys irroratus and
Myosorex varius is difficult. In view of the findings of Bond et al. (1980), it has
been considered necessary to divide the scores for the two species arbitrarily
between the hillside and the valley floor. This will almost certainly prove to be
an inaccurate reflection of the situation, but it is unavoidable until further data
become available.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 23

Mus minutoides and Suncus varilla are classed as ‘Sundry’ for the reasons
given above for Byneskranskop 2. It is noticeable that M. minutoides occurs in
approximately double the proportions at Boomplaas that it does at Nooit-
gedacht. This may be taken as further proof that this species is at an advantage
in cultivated areas and more at risk on open ground.

It would appear that the micromammalian evidence provides an accurate
indication of the vegetation of the Cango valley. Moreover, proportions of
vegetation types within the topographic units seem to be approximately sug-
gested by the proportions of small mammals in the samples. Greater precision
and detail will only be possible when more data are available concerning the
habits and ecological distribution of the small mammals.

MICROMAMMALIAN EVIDENCE FOR PALAEOENVIRONMENTAL
CHANGE

The micromammalian material has been examined in three ways in an
attempt to extract information concerning vegetational and climatic change.
Kowalski (1971: 473) notes that in the Late Pleistocene in Europe ‘the rodent-
spectrum method enables better recording of even slight fluctuations in the
vegetation on the basis of mammalian remains than does any other method’.
This is confirmed in the present study where analysis of change in community
composition has proved by far the most productive method at present. It has,
therefore, received the most attention and has been considered in the greatest
detail. For palaeoclimatic interpretation Kowalski (1971: 466) lists three areas
of investigation. One of these concerns morphological change in certain species
in response to changes in climate. Experiment has shown this to be a poten-
tially useful line of investigation in the southern Cape as well. However, it
requires a great deal more basic information regarding the reaction of modern
populations than is currently available before the full potential can be realized.
A second method concerns the fact that there tend to be more mammalian
species represented in mild climates than in harsh ones. Investigation of
changes in species diversity and community structure, which constitute exten-
sions of the basic fact of numbers of species, suggests that information thus
derived is likely to remain more general even if more data on modern
communities are acquired. The third line of investigation, physical adaptations
reflected in the structure of the teeth and skeleton, has been reserved for future
attention.

EVIDENCE FROM COMMUNITY COMPOSITION

The various types of analysis performed and the basic procedures involved
in interpretation have been described above. At this stage it is necessary only to
outline the format of the present section. Each analysis is treated separately,
beginning with the analyses of the individual sites and continuing with the
multiple analyses. Because the species analyses may be ancillary to the levels

254 ANNALS OF THE SOUTH AFRICAN MUSEUM

analyses, they have been placed first. In each case tables list variables that load
highly on factors in the different analyses, together with the actual loadings. In
the case of the Oblique matrices the variables are listed in the order of their
loadings, with the highest first.

Boomplaas A

Species analysis

In the species analysis, species loading highly on Factor 1 (Table 15) are
those that occur in higher proportions during the Holocene or minimally
throughout the sequence (Fig. 8). One group of these species represents hillside
vegetation. Praomys verreauxi and Crocidura cyanea (if C. c. cyanea) indicate
scrub vegetation, probably on the higher slopes: Aethomys namaquensis and
Elephantulus edwardi suggest sparse, possibly semi-arid vegetation, probably
on the lower slopes. Otomys laminatus may indicate grass or ‘grass’ sensu Bond
et al. (1980: 38). The valley floor is represented by another group of species,
Saccostomus campestris, Steatomys krebsi and Mystromys albicaudatus, which

TABLE 15

Species loading highly on factors in the Varimax rotated factor matrix (A) and the Oblique
rotated factor pattern matrix (B) for Boomplaas A.

Factor 1 Factor 2 Factor 3
A:

E. edwardi 0,92836 C. duthiae 0,89387 O. unisulcatus 0,86738
C. flavescens 0,87271 M. varius 0,96018

C. cyanea 0,81764 S. varilla 0,93976

C. hottentotus 0,78570 D. melanotis 0,97245

S. campestris 0,78925 O. saundersae 0,89522

S. krebsi 0,87093 A, subspinosus 0,75281

M. albicaudatus 0,92848 M. minutoides 0,83571

O. laminatus 0,75185

O. irroratus 0,76388

A. namaquensis 0,82348

P. verreauxi 0,76681

R. pumilio 0,75536

B:

E. edwardi 0,93864 D. melanotis —1,00653 O. unisulcatus 0,74233
M. albicaudatus 0,92943 M. varius —0,97088

S. krebsi 0,90613 S. varilla —0,93473

C. flavescens 0,86056 C. duthiae —0,87845

C. cyanea 0,81309 O. saundersae —0,87140

S. campestris 0,80834 M. minutoides —0,80355

A. namaquensis 0,79290
C. hottentotus 0,74969

O. laminatus 0,73416
P. verreauxi 0,72780
O. irroratus 0,72192
R. pumilio 0,71261

88,4% of total variance accounted for. Factor correlations all low.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 255

suggests that grass, possibly with some trees, occurred in the vicinity. Crypto-
mys hottentotus endorses the suggestion of fairly open grass on the alluvial soil
along the floor of the valley. Otomys irroratus and Rhabdomys pumilio may
occur in either place but both exhibit a preference for dense vegetation,
particularly the former. It is perhaps likely in the present case that the majority
of Otomys irroratus represents dense grass and reeds along the banks of the
river. Equally Rhabdomys pumilio may occur here or elsewhere on the valley
floor, depending perhaps in part on numbers of O. irroratus. O. irroratus may
also have come from dense microphyllous vegetation on wetter hillslopes.
Rainfall appears to have been in the region of 400-600 mm per annum.

Species loading highly on Factor 2 (Table 15) are those that occur in
greater proportions during the Upper Pleistocene (Fig. 8). Otomys saundersae
and Acomys subspinosus refer to the hillsides; the former indicates relatively
open restioid or ‘grassy’ vegetation and the latter dense vegetation, probably on
the higher hillsides. Dendromus melanotis suggests grass on the valley floor.
Chlorotalpa duthiae is also a valley floor species that may indicate trees.
Myosorex varius normally occurs in dense vegetation and sometimes under
fallen trees and probably here represents the streamside habitat. It has,
however, obviously wide tolerance and may occur in a variety of habitats at
different altitudes. In general, therefore, this factor seems to indicate a more
Open vegetation with less scrub on the hillsides. The generally smaller size of
the species may tend to confirm the more open nature of the vegetation. It
would seem also that generally colder conditions must be represented.

It is relevant at this point to consider the possible relationship between
Otomys irroratus and Myosorex varius since this could have a major bearing on
interpretation of the data. These species apparently occur in similar habitats
and, yet, shifts in their proportional representation are largely complementary.
Such a seeming contradiction may indicate dependent reaction as discussed
above. To some extent this could be true. M. varius is apparently a broad-niche
species and O. irroratus a narrow-niche species. Thus, the latter would have the
advantage within the niche to which it is adapted (see Brooks 1974), but
M. varius would have the advantage under all other circumstances. In particu-
lar, it would be better able to adapt to environmental change. This greater
adaptability may be connected with a lesser dependence on dense vegetation. It
appears, for instance, that O. irroratus is dependent upon dense vegetation,
whereas M. varius may be principally sensitive to atmospheric moisture levels.
If this is the case, dense vegetation, frequent mists or adequate rainfall may
satisfy its requirements. It has also to be borne in mind that as well as providing
O. irroratus with protection from predators, the vegetation also constitutes the
source of food. M. varius, on the other hand, may be nocturnal and therefore
less in need of protection; it is also insectivorous and thus only indirectly
dependent on the vegetation for food. Thus it seems that the relationship
between the two species is complex, being partly interdependent and partly
independent.

256 ANNALS OF THE SOUTH AFRICAN MUSEUM

Only Otomys unisulcatus loads highly on Factor 3 (Table 15). It is
distinguished by occurring in its highest proportions mainly during the later
Upper Pleistocene. Its occurrence suggests very open vegetation, probably
semi-arid scrub. In particular, it is likely to have been found on the valley floor
and lower slopes of the hills.

Levels analysis

In the levels analysis the sequence is divided into three main periods, one
Holocene and two Upper Pleistocene. The upper levels load highly on Factor 3
in the Varimax solution (Table 16A). These levels, which have been dated
radiometrically to the Holocene, are characterized by relatively high propor-
tions of the species that load highly on Factor | in the species analysis (Fig. 9).
This indicates that the vegetation during the Holocene comprised scrub on the
hillsides, possibly sparse and semi-arid in places and elsewhere dense and
microphyllous. Fairly open grass, perhaps with some trees, on the valley floor
would have been replaced by denser grass and reeds on the river-banks.

The levels loading highly on Varimax Factor 2 (Table 16A) are mainly
those dated radiometrically to the late Upper Pleistocene and including the last
glacial maximum. Level BOLS alone is earlier (Fig. 9). These levels are
characterized by relatively high proportions of Otomys unisulcatus (Fig. 9).
High proportions of Otomys saundersae are also an important feature, as the
high loading of level YOL in the Oblique solution (Table 16B; Fig. 9) indicates.

TABLE 16

Levels loading highly on factors in the Varimax rotated factor matrix (A) and Oblique rotated
factor pattern matrix (B) for Boomplaas A.

Factor 1 Factor 2 Factor 3

A:

BP4 0,84357 GWA 0,75605 DGL 0,95019
OLP 0,77611 LPC 0,79799 BLD3 0,90311
OLP1 0,88619 YOL 0,83436 BL 0,94740
OLP2 0,91826 BP2 0,75382 BLA 0,86908
OLP3 0,85462 BP3 0,73674 BRL 0,81538
BOL 0,84045 BOLS 0,77767

B: :

YOL 1,01588 BL 0,97979 OLP2 1,02529
BOLS5 0,91898 DGL 0,96491 OLP1 0,94273
LPC 0,91728 BLD3 0,86622 BP4 0,87656
BP2 0,83946 BLA 0,81764 BOL 0,82354
GWA 0,83182 BRL 0,75035 OLP3 0,82152
BP3 0,82136

LP 0,71981

92,2% of total variance accounted for. Factor correlation moderate between Factors 1 and 2,
high between Factors 1 and 3.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS Zo?

Crocidura cyanea is absent and Myosorex varius generally occurs in moderate
proportions. The indication is that vegetation was generally fairly open, poss-
ibly semi-arid on the valley floor and restioid or ‘grassy’ on the hillsides. There
must have been a certain amount of dense vegetation, especially in view of the
fact that the climate was apparently fairly dry. This dense vegetation, probably
grass and reeds, is most likely to have been situated along the river-banks. It is
perhaps most likely to have been cold rather than aridity which drove out C.
cyanea. If it had been the latter, it might be expected that proportions of M.
varius and O. irroratus would have been lower.

The levels loading highly on Varimax Factor 1 (Table 16A) represent a
period of time preceding that discussed above, that is from about 60 000 B.P.
to perhaps 32 400 B.P. These levels are characterized in particular by high
proportions of Myosorex varius and slightly lower proportions of Otomys
saundersae (Fig. 9). This pattern is shown by level OLP2 (Fig. 9) which loads
most highly in the Oblique solution. Otomys irroratus also occurs in fairly high
proportions. The indication is that conditions were less severe than those
pertaining in glacial maximum levels. An extension of dense cover is notice-
able. Generally low proportions of Cryptomys hottentotus suggest that much of
the expansion in dense vegetation occurred on the valley floor, although there
may have been rather more dense scrub on the hillsides as well. The general
indication is that conditions were approximately intermediate between glacial
maximum and interglacial (Holocene), but that they were closer to the former
than to the latter. This is suggested partly by the fact that the same two species
are dominant and partly by the main division of species into a Holocene-
dominant group and an Upper Pleistocene-dominant group. There is,
moreover, a high correlation between the two Upper Pleistocene factors in the
Oblique solution, which would seem to confirm the suggestion.

A supplementary analysis, aimed at examining the Holocene sequence in
greater detail, confirmed the basic pattern already established. It further
showed that conditions during the Holocene were sufficiently stable for any
changes not to be detectable by present methods.

Byneskranskop I

Species analysis

In the species analysis, species that load highly on Factor 1 (Table 17) are
those that tend to occur in higher proportions in the upper half of the sequence
(Fig. 10). The indication is of fairly extensive grass on the plains (Steatomys
krebsi), dense in places (Dendromus melanotis), especially along the river-
banks (Myosorex varius). The generally smaller size of the species loading on
this factor may indicate relatively open vegetation as was discussed above.
These species are all prominent in the Upper Pleistocene at Boomplaas A and
it may be that their ascendency at Byneskranskop 1 could indicate relatively
cold and dry conditions.

Species loading highly on factors in the Varimax rotated factor matrix (A) and

ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 17

rotated factor pattern matrix (B) for Byneskranskop 1.

Factor 3

O. laminatus
O. saundersae
O. irroratus

Factor | Factor 2

Re

M. varius 0,85083 C. flavescens 0,70743
S. varilla 0,88919 G. capensis 0,90159
D. melanotis 0,83358 R. pumilio 0,74821
S. krebsi 0,96512

M. minutoides 0,72959

B:

S. krebsi 1,00507 G. capensis 0,95846
S. varilla 0,91596
M. varius 0,84172

D. melanotis 0,78344

O. laminatus

81,6°% of total variance accounted for. Factor correlations low or very low.

FACTOR |
%o

M.varius

20

fe)

5 M.minutoides
(@)

5 _D.melanotis

30 S.krebsi

20

FACTOR 2
%
20 C.flavescens

10

O

G

‘Landes

R

capensis
. pumilio

5
0
f2e5o78 Sanit i
ABOI 2345679

FACTOR 3

%o

5 _Q.laminotus

O

O.saundersae

20

10

O.irroratus

1256789

ABO|2

Oblique

0,92387
0,72222
0,73970

0,87055

WLLL A
345679

Fig. 10. Variation in percentage representation of species loading highly in the analysis of
species from Byneskranskop | (. = less than 0,5 %).

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 259

The species that load highly on Factor 2 (Table 17) show a tendency to
vary inversely in proportion with the species loading highly on Factor 1| (Fig.
10). Tatera afra shows a similar though less distinct tendency. These species
again represent the flat ground with the possibility of pans in the area (Geory-
chus capensis). Low scrub, possibly coastal fynbos (G. capensis), together with
patches of dense vegetation (Rhabdomys pumilio) are indicated. Tatera afra
would probably indicate more open grass. Dense vegetation, probably grass on
the lower hillsides, is suggested by Crocidura flavescens. It is possible that
conditions indicated by this factor are rather wetter than those indicated by
Factor 1. The alternative is that greater seasonality is suggested by Factor 2
than by Factor 1. The Oblique solution shows Georychus capensis to load
highly on Factor 2 and Steatomys krebsi most highly on Factor 1, and fynbos
would require a more seasonal rainfall than would grass.

The species that load highly on Factor 3 (Table 17) tend to show a general
decline in proportional representation from the bottom to the top of the
sequence, but with a certain amount of recovery in level 2-4 (Fig. 10).
However, whereas the first two factors referred to the plain, the third factor
refers to the hillsides as well as the river-banks. The data suggest a restioid or
‘grassy’ vegetation on the upper and lower hillsides (Otomys saundersae,
Otomys laminatus). This vegetation may perhaps have been on the sandstone
hills as it is at present. Otomys irroratus probably indicates dense waterside
vegetation but it may also suggest a dense microphyllous proteoid element
which is today found on the limestone ridges. The general decline in O.
saundersae may suggest a gradual increase in temperature on the grounds that
this species was clearly at an advantage during the last glacial maximum at
Boomplaas A but declined rapidly thereafter.

Levels analysis

In the levels analysis the lower levels and level 2 load highly on Factor 1
(Table 18). Radiometric dates indicate that the period represented is up to
6500 B.P. and from approximately 3 900 to 3 400 B.P. These levels are
generally characterized by high proportions of Otomys saundersae and, to a
slightly lesser extent, Otomys irroratus (Fig. 11). Tatera afra and Crocidura
flavescens also tend to be fairly well represented. Level 13 loads most highly in
the Oblique solution (Table 18B), which suggests that a high proportion of
Otomys saundersae is the most important element. The indication is that there
was extensive dense vegetation on both the river banks and the lower hillsides
and that there was grass elsewhere on the flats; a restioid or ‘grassy’ element
and possibly a proteoid element occurred on the hills. On the basis of
connotations suggested for Boomplaas A conditions would appear to have been
relatively cold and wet.

Levels comprising much of the upper half of the sequence load highly on
Factor 2, with level 9A loading most highly in the Oblique solution (Table
18B). This level is distinguished by high proportions of Steatomys krebsi and

260 ANNALS OF THE SOUTH AFRICAN MUSEUM

FACTOR | FACTOR 2 FACTOR 2 (cont.)
% %o

10

-

nm
io)

.

i)

re)

ro) = ro
o Oo
- a
re) = nm
i

oO

FACTOR | (cont.)

14 26). "VF

1S 20.19
is that

i)
oO

-
E

ine)
oO

° re)
O

O
abcdefghijkimno abcdef ghijkimno abcdefghi jk!imno
a C.asiatica e G.capensis i M.albicaudatus m O.laminatus
b Mvarius f M.minutoides j T.afra n Q.saundersae
c¢ Svarilla g Pverreauxi k D.melanotis o O.irroratus
d Cflavescens h R.pumilio | S.krebsi

Fig. 11. Proportions of species from levels loading highly in the analysis of levels at
Byneskranskop 1 (. = less than 0,5 %).

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 261

TABLE 18

Levels loading highly on factors in the Varimax
rotated factor matrix (A) and the Oblique rotated
factor pattern matrix (B) for Byneskranskop 1.

Factor 1 Factor 2
A:

2 0,79895 5 0,83539
11 0,71621 6 0,84614
12 0,88697 7 0,81348
13 0,89755 8 0,92488
14 0,89078 9A 0,94780
We 0,70501 9B 0,84664
16 0,88258 10 0,71036
iy 0,86701
19 0,88958
B:
be 0,98908 9A 1,08319
19 0,96082 8 1,02657
16 0,96055 9B 0,85978
14 0,96000 6 0,82629
12 0,95225 5 0,81876
17 0,92879 ‘, 0,77142

2 0,79323

88,4% of the total variance accounted for.
Factor correlations moderate.

secondarily by fairly high proportions of Myosorex varius. The indication is that
conditions were rather drier and probably also slightly warmer, there being
fewer O. saundersae. There would appear to have been extensive, more or less
open, grass on the flats. The reason for the replacement of Tatera afra by
Steatomys krebsi as the most plentiful plains animal is not clear. It may,
however, be that S. krebsi prefers more open grass or it could be a question of
climate, with S. krebsi preferring drier conditions if it is similar to S. pratensis,
which is said not to favour marshy conditions in the Namib Desert (Coetzee
1969: 32). That S. krebsi prefers warmer conditions is suggested by the fact that
it apparently did not enter the southern Cape until after the end of the Last
Glacial.

Die Kelders 1 Middle Stone Age samples

Species analysis

In the Varimax solution of the amended species analysis, species that load
highly on Factor 1 (Table 19A) are those occurring in higher proportions in the
upper and lower levels than in the central levels (Fig. 12). There is particular
emphasis on the top two levels, except in Acomys subspinosus which occurs in
very small numbers throughout. Tatera afra indicates grassy plains which may

262 ANNALS OF THE SOUTH AFRICAN MUSEUM

FACTOR |
%
S _A-subspinosus FACTOR 2
%
10 _Pverreauxi 40. M.varius
fe) ' ory 30.
30 __ T.afra 20
10 FACTOR 3
O 5 _C.flavescens
(@) e@eee#e#
S.varilla
me) lO M.albicaudatus
Q.irroratus
30 O O eoeepveeete ese
20 S _M.minutoides 20 O.saundersae
10 e 10
S _D.melanotis
O O (@)
{ 325-7. Sarvs (-& 3.7" Sill BS | 35s 7S ie

Fig. 12. Variation in percentage representation of species loading highly in the analysis of
species from Die Kelders 1 M.S.A. levels (¢ = less than 0,5%).

have been above or below the cave, depending upon the height of the sea at the
time. Otomys irroratus probably indicates dense waterside vegetation, perhaps
near a presently submerged marsh or lake, but it may also suggest dense scrub
on hillsides. Such dense scrub is also suggested by Acomys subspinosus, while
Praomys verreauxi may indicate more open scrub. The high loading of Otomys
irroratus in the Oblique solution (Table 19B) confirms the fact that dense
vegetation is a major feature of this factor. Otomys irroratus and Praomys
verreauxi form part of the Holocene pattern at Boomplaas A, and it was
suggested that Tatera afra was dominant at a relatively wet time at Byneskrans-
kop 1. It seems likely, therefore, that relatively warm, wet conditions are.
indicated by this factor.

The species loading highly on Varimax Factor 2 (Table 19A) constitute the
small cold element previously recognized at Boomplaas A and Byneskranskop

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 263

1. More specifically, they form the intermediate element at Boomplaas A. At
Die Kelders 1 these species occur in higher proportions in the central levels,
their proportions being basically the converse of those exhibited by Factor 1
species (Fig. 12). The climate was probably cooler and drier with more open
vegetation containing a greater proportion of grass.

The species loading highly on Varimax Factor 3 (Table 19A) occur in
higher proportions in the lowest three levels. The indications are of fairly
extensive open grassland on the flats (Mystromys albicaudatus) with denser
vegetation at the base of the hills or cliffs (Crocidura flavescens) and restioid or
‘grassy’ vegetation on the hillsides (Otomys saundersae). The climate was
probably relatively cold.

Dendromus melanotis, which loads highly on Varimax Factor 4 (Table
19A), is distinguished by occurring in its highest proportions in level 2 (Fig.
12). This species suggests an increase in bush and perhaps even some trees. The
implication would be a more moderately warm, wet climate which is not
inconsistent with findings for Factor 1.

TABLE 19

Species loading highly on factors in the Varimax rotated factor matrix (A) and Oblique
rotated factor pattern matrix (B) for Die Kelders 1 M.S.A. levels (amended data).

Factor 1 Factor 2 Factor 3 Factor 4

A:

T. afra 0,92560 M. varius 0,76203 C. flavescens 0,96852 D. mesomelas 0,81893
O. irroratus 0,93543 S. varilla 0,90260 M. albicaudatus 0,83947

. subspinosus 0,89375 D. melanotis 0,85728 O. saundersae 0,81984 °
. verreauxi 0,89097 M. minutoides 0,92113

. afra 0,93805 M. albicaudatus 0,78883 S. varilla 0,92614
. verreauxi 0,91248 O. saundersae 0,74763 D. melanotis 0,84101

A
P
B:
o. irroratus 0,96112 C. flavescens 0,98743 M. minutoides 0,94282
P
A. subspinosus 0,86600

90,3 % of the total variance accounted for. Factor correlations low in all cases.

Levels analysis

In the levels analysis the central levels, 11 up to 3 or 4, load highly on
Factor 1 with level 2 added in the analysis of amended data (Table 21). The
distinguishing features are high proportions of Myosorex varius and rather
lower proportions of Suncus varilla, Tatera afra, and Otomys irroratus (Fig.
13). The effect of amending the data is to raise the proportions of the first two
species in level 2 (Fig. 13) which presumably explains why this level loads
highly on Factor 1 when amended data are used. Levels 6 and 8 load most
highly in the Oblique solutions (Tables 20-21) whether or not the amended
data are used. It would appear that conditions were moderately cool and dry
with a reduced amount of open grass on the flats but a moderate amount of
restioid or ‘grassy’ vegetation on the hillsides.

The lowest three levels, with or without the top two levels, load highly on
Factor 2. Using the original data, level 1 loads most highly on this factor in the

264 ANNALS OF THE SOUTH AFRICAN MUSEUM

Oblique solution (Table 20). Here again Myosorex varius, Tatera afra, and
Otomys irroratus are most numerous, but the latter two species have overtaken
Myosorex varius. The fact that basically only proportions and not the species
themselves have changed, may explain why the two factors are highly or
moderately correlated in the Oblique solutions. It also suggests that the
differences between the factors are not great. Apparently wetter conditions
with much more extensive grassland are indicated by this factor. It may be that
when the original data are used there is a colder and a warmer element within
this factor. It is possible that Praomys verreauxi indicates scrub under warmer
conditions in the upper levels and Otomys saundersae suggests restioid or
‘grassy’ vegetation under colder conditions in the lower levels. In this context it
is notable that the two species load on different factors both at Die Kelders 1
and Boomplaas A. Moreover, Praomys verreauxi is barely present during
glacial maximum periods at the latter site. The suggestion of milder conditions
with more closed vegetation in the upper levels is endorsed by high proportions
of Otomys irroratus. The lower levels are further distinguished by the presence
of Mystromys albicaudatus. This is particularly noticeable when the amended
data are used; the order in which the levels load in the Oblique solution (Table
21) agrees with the relative importance of M. albicaudatus. Proportions of

TABLE 20

Levels loading highly on factors in the Varimax

rotated factor matrix (A) and Oblique rotated

factor pattern matrix (B) for Die Kelders 1
M.S.A. levels (original data).

Factor 1 Factor 2

A:
3 0,73229 1 0,86177
4 0,89936 Z 0,80720
5 0,86842 12 0,79641
6 0,91322 13 0,81997
7 0,91296 14 0,74322
8 0,91306
9 0,86275

10 0,87094

11 0,88203

EB: :
6 1,04775 1 0,99601
8 1,04580 13 0,87854
7 1,04276 2 0,82514
4 0,99940 12 0,83845

11 0,94879 14 0,70535

10 0,90559
5 0,89793
9 0,88881

92,3% of the total variance accounted for.
Factor correlation high.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 265

TABLE 21

Levels loading highly on factors in the Varimax

rotated factor matrix (A) and Oblique rotated

factor pattern matrix (B) for Die Kelders |
M.S.A. levels (amended data).

Factor 1 Factor 2

>

RK OOMANAMAHWDN !,
i=)
\O
—
foe]
\©
—

0,78890 12 0,79377
0,75778 ‘3 0,85395
0,89531 14 0,90062

ph peek

to

NONAOBRNK DO...
So
\O
ON
Q
|
ee

1,03659 14 0,84118
1,02994 13 0,81722
0,98597 12 0,78936

—

—

89,9% of the total variance accounted for.
Factor correlation moderate.

Tatera afra are also high and there would appear to have been a great deal of
grass in the area at that time. The precise differences in the habitat preferences
of these two species are not known. Possibly, however, Mystromys albicaudatus
occurred on lower ground which subsequently became submerged. It might
then have been unable to compete with Tatera afra on the higher ground, due
perhaps to unfavourable conditions such as reduced grass cover.

Coastal samples

Species analysis

In the species analysis the solutions are virtually identical and the Varimax
solution, including Steatomys krebsi, is taken as representative (Table 22).
Factor 1 is more comprehensive in the picture afforded by the greater number
of species loading highly on this factor. Dense waterside vegetation is indicated
by Myosorex varius, Otomys irroratus, and possibly Rhabdomys pumilio. Else-
where on the flats, grass (Dendromus melanotis) with scattered trees or bushes
(Tatera afra, Dendromus mesomelas) and possibly some open scrub (Chryso-

266

aoqo9g&a

ANNALS OF THE SOUTH AFRICAN MUSEUM

FACTOR |

“/o

abc defghijkIimno

C.asiatica e
M.varius f
S.varilla g
C.flavescens h

FACTOR

o

G.capensis
A.subspinosus
M.minutoides
P.verreauxi

| (cont.)

abcde fghijkIimno

i
j
k
|

FACTOR 2
“o

30

|
20
10
0

12
20
10
O
13
20
10
14
20
“10
oO
abcdef ghi jk Ilmno
R.pumilio m O.mesomelas
M.albicaudatus n OQO.saundersae
T.afra o O.irroratus

D.melonotis

Fig. 13. Proportions of species from levels loading highly in the analysis of M.S.A.
levels at Die Kelders 1 (. = less than 0,5 %).

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 267

TABLE 22

Species loading highly on factors in the Varimax rotated factor
matrix (A) and Oblique rotated factor pattern matrix (B) for
coastal samples (Steatomys krebsi included).

_

Factor | Factor 2

A:

C. asiatica 0,96655 C. flavescens 0,96267
M. varius 0,96153 G. capensis 0,86370
S. varilla 0,93595 S. krebsi 0,84076
D. melanotis 0,94581 M. albicaudatus 0,73592
D. mesomelas 0,90984 O. laminatus 0,76755
T. afra 0,88450

O. irroratus 0,94844

A, subspinosus 0,93390
M. minutoides 0,95332

P. verreauxi 0,90731
R. pumilio 0,93716
B:
C. asiatica 0,96698 C. flavescens 0,99900
M. varius 0,96251 G. capensis 0,82996
M. minutoides 0,95298 S. krebsi 0,82315
O. irroratus 0,94370
4 D. melanotis 0,94333
S. varilla 0,93407
A. subspinosus 0,93054
R. pumilio 0,93053
D. mesomelas 0,89963
P. verreauxi 0,89579
T. afra 0,86892

83,4% of the total variance accounted for. Factor correlation
very low.

chloris asiatica) is indicated. On the hillsides Acomys subspinosus and Praomys
verreauxi suggest scrub. The climate would appear to have been at least
moderately warm with a rainfall in excess of 400 mm per annum. The picture
indicated by Factor 2 is dense vegetation (Crocidura flavescens), and probably
grass (Otomys laminatus) on the lower hillslopes. Grass (Mystromys albicauda-
tus, Steatomys krebsi) is also indicated on the flats, together with pans and
perhaps some scrub (Georychus capensis).

In the Oblique solution, Chrysochloris asiatica and Myosorex varius load
most highly on Factor 1 whether or not Steatomys krebsi is included (Tables
22-23). Crocidura flavescens is central to the pattern represented by Factor 2.
The indication is that central aspects of Factor 1 are sparser vegetation and a
drier climate. For Factor 2 denser vegetation and rather wetter conditions are
indicated.

In general, the species loading highly on Factor 1 are those which occur
either in higher proportions in the DK1 M.S.A. levels or in approximately

268 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 23

Species loading highly on factors in the Varimax rotated factor
matrix (A) and Oblique rotated factor pattern matrix (B) for
coastal samples (Steatomys krebsi excluded).

Factor 1 Factor 2

A:

C. asiatica 0,96553 C. flavescens 0,96345
M. varius 0,95874 G. capensis 0,86511
S. varilla 0,93186 M. albicaudatus 0,76555
D. melanotis 0,94620 O. laminatus 0,76059
D. mesomelas 0,91990

T. afra 0,88604

O. irroratus 0,94861

A. subspinosus 0,94549
M. minutoides 0,95184

P. verreauxi 0,91106

R. pumilio 0,93963

B:

C. asiatica 0,96461 C. flavescens 0,98828
M. varius 0,95723 G. capensis 0,83015

M. minutoides 0,94863
A. subspinosus 0,94815

O. irroratus 0,94521
D. melanotis 0,94185
R. pumilio 0,93417
S. varilla 0,92605
D. mesomelas 0,91540
P. verreauxi 0,90101
T. afra 0,87401

83,7% of the total variance accounted for. Factor correlation
very low.

equal proportions throughout. The species that load highly on Factor 2 are
those which occur in higher proportions at BNK1.

Levels analysis

In the levels analysis, most of the BNK1 levels load highly on Factor 1,
whether or not Steatomys krebsi is included (Tables 24-25). Because BNK1 is
dated radiometrically to the postglacial period, it may be assumed that this
factor represents relatively warm conditions. In the Oblique solution, level 6
loads most highly when S. krebsi is included, whereas level 11 loads most highly
when it is omitted. Since level 11 is the second highest loading level in the first
analysis, it may be considered the most generally representative. This is
probably explainable in terms of the fact that in the BNKI1 analysis, level 11
tends to be intermediate. Level 6 is distinguished by equally high proportions of
Myosorex varius and Steatomys krebsi and slightly lower proportions of Croct-
dura flavescens and Otomys saundersae. Level 11, without Steatomys krebsi, is
distinguished by high proportions of Otomys saundersae and slightly lower

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 269

proportions of Myosorex varius and Crocidura flavescens. The omission of
Steatomys krebsi would appear, therefore, only to promote a slight change in
emphasis. The indication is of relatively cold, dry conditions with reduced
dense streamside vegetation, but fairly extensive dense vegetation on the lower
hillsides. Extensive grass on the plains and upper hillsides is indicated.

The majority of the DK1 M.S.A. levels loads highly on Factor 2 with levels
7 and 8 loading most highly in the Oblique solution whether or not Steatomys
krebsi is included (Tables 24-25). Since the DK1 M.S.A. levels are thought to
be Upper Pleistocene in age, this factor should represent rather colder condi-

TABLE 24

Levels loading highly on factors in the Varimax rotated factor matrix (A) and Oblique rotated
factor pattern matrix (B) for coastal samples (Steatomys krebsi included).

Factor 1 Factor 2 Factor 3 Factor 4

A:
BNKI1: 1 0,92329 DKI M.S.A: 3 0,93394
2 0,86238 4 0,98497
5 0,90716 5 0,98596
6 0,93519 6 0,97390
7 0,89096 7 0,98565
8 0,86791 8 0,98093
: 9A 0,87869 9 0,97217
9B 0,78084 10 0,97645
10 0,95231 11 0,96473
Tt 0,97358 14 0,83609
12 0,90922 13 0,71645
13 0,86676
14 0,91204
15 0,90712
16 0,82248
17 0,79429
19 0,86448
B:
BNKI: 6 0,98901 DK1 M.S.A: 7 1,02513
11 0,98452 8 1,01752
10 0,96754 4 1,01722
9A 0,96334 5 1,00761
8 0,94502 6 0,99812
5 0,94372 10 0,98780
Z 0,92715 9 0,97808
1 0,92101 11 0,96408
15 0,88448 3 0,87517
14 0,85260 12 0,73803
12 0,84869
9B 0,83782
z 0,81650
13 0,79294
19 0,78708
16 0,74114

91,6% of the total variance accounted for. Factor correlations low in all cases.

270 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 25

Levels loading highly on factors in the Varimax rotated factor matrix (A) and Oblique
rotated factor pattern matrix (B) for coastal samples (Steatomys krebsi excluded).

Factor | Factor 2 Factor 3 Factor 4

A:

BNKI: 1 0,91154 DKI1 M.S.A: 3 0,88514
y 0,87867 4 0,97039
5 0,86818 5 0,96722
6 0,90341 6 0,95426
7 0,83742 7 0,97045
8 0,80173 8 0,97866
9A 0,81366 9 0,96167
10 0,92246 10 0,95671
11 0,96692 11 0,95212
12 0,92068 12 0,78523
13 0,91092
14 0,95965
15 0,87502
16 0,85971
17 0,82840
19 0,89241

B:

BNKI: 11 1,00342 DK1 M.S.A: 8 1,03686
14 0,99307 7 1,01440
2 0,96243 4 1,01118
1 0,95995 5 0,99676
2 0,94240 9 0,98918
13 0,94231 6 0,98714
6 0,93010 10 0,97629
10 0,92427 11 0,97431
19 0,92199 3 0,85559
5 0,87770 12 0,72414
16 0,85570
15 0,84938
dG 0,81348
7 0,81472
9A 0,78855
8 0,78172

91,5% of the total variance accounted for. Moderate correlation between Factors 1 and 2,
otherwise low.

tions than those represented by Factor 1. Levels 7 and 8 are distinguished by
high proportions of Myosorex varius and rather lower proportions of Suncus
varilla, Tatera afra, and Otomys irroratus. The high proportion of Myosorex
varius in levels loading on Factors 1 and 2 explains the moderate correlation
between these two factors. The fact that Otomys irroratus also occurs in a fairly
high proportion possibly suggests that Factor 2 represents a situation where
dense waterside vegetation makes up a relatively large part of the overall
vegetation. Tatera afra indicates grass elsewhere on the flats but the significance
of Suncus varilla is unfortunately not known. It seems that basically the

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS os

difference between the two factors is one of degree as far as the vegetation is
concerned. The only clear difference in kind appears to lie in the proportions of
Crocidura flavescens which is virtually absent from the DK1 M.S.A. levels.
There is, therefore, no connotation of dense vegetation on the lower hillsides
for Factor 2.

Holocene and modern samples

Species analysis

In the species analysis the grouping of the species is similar to that
encountered in other analyses (Table 26). Factor 1 indicates a moderate
amount of dense waterside vegetation (Myosorex varius) with grass elsewhere
on the flats (Dendromus spp.) and lower hillsides (Otomys laminatus). Also on
the hillsides, scrub is suggested by Praomys verreauxi. A moderate climate is
indicated. Factor 2 suggests a colder climate with grass or open scrub on the
upper hillsides (Otomys saundersae), dense vegetation on the lower hillsides
(Crocidura flavescens), and open grassland on the flat ground (Mystromys
albicaudatus). Factor 3 indicates more or less open grassland with some bushes
(Steatomys krebsi), although it is not clear how this factor differs from the
others climatically.

- In the Oblique solution, the species central to the pattern represented by
Factor 1 are Mus minutoides, Dendromus melanotis and, to a lesser extent,
Myosorex varius (Table 26B). These species are those that are interpreted as

TABLE 26

Species loading highly on factors in the Varimax rotated factor matrix (A) and Oblique
rotated factor pattern matrix (B) for Holocene and modern samples.

Factor 1 Factor 2 Factor 3

A:

M. varius 0,90679 C. flavescens 0,89048 S. krebsi 0,93227
S. varilla 0,82715 M. albicaudatus 0,90012

D. melanotis 0,92403 O. saundersae 0,77251

D. mesomelas 0,79299

O. laminatus 0,72755

M. minutoides 0,92981

P. verreauxi 0,74619

R. pumilio 0,85238

B:

M. minutoides 0,94258 M. albicaudatus 0,88385 S. krebsi 0,92996
D. melanotis 0,93562 C. flavescens 0,86102

M. varius 0,90510

S. varilla 0,83443

D. mesomelas 0,81630

R. pumilio 0,81603

82,6% of the total variance accounted for. Factor correlations very low.

272 ANNALS OF THE SOUTH AFRICAN MUSEUM

representing interstadial conditions in the Boomplaas A analysis. Mystromys
albicaudatus and Crocidura flavescens load highly on Factor 2. The distinction
between these two factors is basically the same as that occurring in the previous
analysis of coastal samples. Factor 1 appears to represent cooler drier condi-
tions with sparser vegetation than does Factor 2. Factor 3 is distinguished by a
high loading of Steatomys krebsi. The reason for this is almost certainly
topographic and, as such, of little use in the present work.

Levels analysis

In the levels analysis the BPA levels load highly on Factor 1, together with
BNK1 level 19 in the Varimax solution (Table 27A). In the Oblique solution
BPA level BRL4UA loads most highly (Table 27B). The BPA levels are

TABLE 27

Levels loading highly on factors in the Varimax rotated factor matrix (A) and Oblique rotated
factor pattern matrix (B) for Holocene and modern samples.

Factor 1 Factor 2 Factor 3 Factor 4

A:

BPA: BLD3A 0,80190 BNKI: 1 0,79037 BNK2 0,84484
BL 0,86097 5 0,91847 BPB-C 0,90962
BLA 0,84449 6 0,90805 NGA-B 0,88355
BRL 0,84450 7 0,92357 GLEN 0,86964
BRL4UA _ 0,91946 8 0,92301
BRL4 0,92291 9A 0,95951
BRLS5 0,90189 9B 0,95343
BRL6 0,92471 10 0,86978
BRL7 0,92105 11 0,82139
CEI 0,88865 12 0,73449

BNK 1-19 0,71169 15 0,81116

B:

BPA: BRL4UA __ 0,97970 BPB-C 0,89277 BNKI1: 9B —1,11151
BL 0,97658 NGA-B __ 0,86808 9A —1,00763
BRL4 0,95835 BNK2 0,86490 7 —0,96127
BRL 0,92726 GLEN 0,82436 6 —0,86357
BRL7 0,90905 5 —0,84630
BRL6 0,86040 11 —0,83457
BLD3A 0,85334 15 —0,83247
BLA 0,84857 10 —0,78009
BRLS5 0,80609
CLI 0,71308

92,1 % of the total variance accounted for. Moderate correlations between Factors 1 and 4, otherwise very low.

characterized by an extremely high proportion of Otomys irroratus, but in level
BRL4UA this is particularly noticeable. Although this is probably due to the
small size of the sample, the same pattern is discernible in level BL, which
loads next most highly. Generally Crocidura flavescens is of secondary impor-
tance and the implication of this factor is extensive dense vegetation both on
the river-banks and along the lower hillsides.

The majority of the BNK1 levels loads highly on Factor 2 (Varimax).
(Table 27A). Level 9B loads most highly, followed by level 9A, in the Oblique
solution (Table 27B). The sample from level 9B is again very small but the
picture provided by level 9A is very similar. The distinguishing features in this
case are high proportions of Steatomys krebsi together with rather lesser
proportions of Myosorex varius. Otomys saundersae is the only other species of

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 293

M.varius

20 1

an -- Pa |
o—_—e-

30, S.krebsi A

O. saundersae

2 3 4 3) 6 7 8 5) 10 \ 4 ie)
x l|OOO yrs BP

Fig. 14. Correlation of Boomplaas A levels BLD3A to CL1 with the Byneskranskop |
sequence. (Solid line = Boomplaas A; dashed line = Byneskranskop 1.)

274 ANNALS OF THE SOUTH AFRICAN MUSEUM

any importance. The moderate correlation between these two factors is prob-
ably due to the fact that O. saundersae occurs in approximately equal propor-
tions at both sites (Fig. 14). It would appear that this factor indicates drier
conditions than does the previous factor. The basic distinction is probably
geographic or topographic in origin. That is to say, large proportions of
Steatomys krebsi are made possible by the presence of open plains, whereas the
high proportions of Otomys irroratus are probably due to the relatively small
amount of flat ground on the valley floor. However, the fact that Myosorex
varius 1s present in higher proportions than is Otomys irroratus in one area and
not in the other does indicate a difference between the two. Factor 2, then,
represents a relatively reduced streamside habitat with more or less open grass
elsewhere on the flat. It probably also represents drier conditions than Factor 1.

The modern samples all load highly on Factor 3, with BPB-C loading most
highly in the Oblique solution (Table 27). The distinguishing features here are a
high proportion of Myosorex varius and a fairly high proportion of Otomys
irroratus. Mus minutoides also occurs in much higher proportions than it does
in the Holocene samples. This is thought to be an artefact of the practice of
agriculture but, in any case, this species is too unspecific in its habitat
requirements to be of use for interpretation. The indication is that there is a
great concentration on the streamside habitat in this factor. Since there are no
levels loading highly on the fourth factor, it was not possible to determine the
environmental connotations of this factor.

Upper Pleistocene and modern samples

Species analysis

In the analysis of species the division of species is very similar to that
observed in the coastal analysis described above (Table 28). Myosorex varius,
Otomys irroratus and, possibly, Rhabdomys pumilio attest dense waterside
vegetation. Acomys subspinosus and Praomys verreauxi indicate scrub on the
hillsides, while Dendromus spp. suggest grass on the lower ground, with some
trees or bushes in the case of D. mesomelas. Factor 2 indicates restioid or
‘grassy’ vegetation on the upper hillsides (Otomys saundersae), dense vegeta-
tion on the lower hillsides (Crocidura flavescens) and open grassland on the
flats (Mystromys albicaudatus). On the basis of the interpretation of the
Boomplaas A analysis, it would seem likely that Factor 1 represents rather
warmer interstadial conditions while Factor 2 represents full glacial conditions.

In the Oblique solution Dendromus mesomelas, followed by Rhabdomys
pumilio, Mus minutoides, and Praomys verreauxi, is central to the pattern
represented by Factor 1 (Tables 3-4). Only Otomys saundersae loads highly on
Factor 2. Since Rhabdomys pumilio and Praomys verreauxi both appear to be
part of the Holocene element at Boomplaas A, this would tend to confirm the
suggestion that Factor 1 represents milder conditions than does Factor 2. It may
also be that wetter conditions are indicated.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS iS

TABLE 28

Species loading highly on factors in the Varimax rotated factor
matrix (A) and Oblique rotated factor pattern matrix (B) for
Upper Pleistocene and modern samples.

Factor 1 Factor 2

A:

M. varius 0,71515 C. flavescens 0,71424
S. varilla 0,86903 M. albicaudatus 0,77563
D. melanotis 0,81438 O. saundersae 0,89907
D. mesomelas 0,90757

O. irroratus 0,84780

A. subspinosus 0,84536
M. minutoides 0,93138

P. verreauxi 0,91645

R. pumilio 0,92568

B: joa

D. mesomelas 0,97707 O. saundersae 0,95466
R. pumilio 0,94764

M. minutoides 0,93837

P. verreauxi 0,93625

S. varilla 0,83498

O. irroratus 0,81435

A, subspinosus 0,79736
D. melanotis 0,75197

81,9% of the total variance accounted for. Factor correlation low.

In general, the species that ioad most highly on Factor 1 occur in approxi-
mately equal proportions at both sites or are rather better represented at DK1.
The species that load highly on Factor 2 are those that occur in greater
proportions at BPA.

Levels analysis

In the levels analysis the two solutions are slightly different (Table 29).
The reason for this is not known, except that it may have been caused by the
moderate correlation between Factor 1 and Factors 3 and 4. Because of this
correlation it was considered that the Oblique solution was more likely to be
correct and it was therefore utilized for interpretation. Apart from BPB and C,
the levels here loading highly on Factor 1 are those that, in the analyses of
individual sites, loaded highly on factors considered to represent interstadial
conditions. Level OLP3 loads most highly on this factor with a very high
proportion of Myosorex varius and with Otomys saundersae of secondary
importance. As has been suggested above, this probably represents a mod-
erately dry, cold climate.

NBC level YSL loads highly on Factor 2 and this level is dated radio-
metrically to the last maximum of the Last Glacial (Table 29). This factor may
thus be considered to represent glacial conditions. The very high proportions of

276 ANNALS OF THE SOUTH AFRICAN MUSEUM

Otomys saundersae tend to support this suggestion. The only other species to
occur in any numbers is Mystromys albicaudatus, so that the general impression
is of extensive grassland on the flats and restioid or ‘grassy’ vegetation on the
hillsides.

DK1 M.S.A. level 2, which loads highly on Factor 3, is distinguished by
very high proportions of Otomys irroratus, with Praomys verreauxi and Myo-
sorex varius the only other species to occur in even moderate proportions.
Extensive dense waterside vegetation and scrub on the hillsides are indicated.
By comparison with the Boomplaas A analysis, it would appear that this factor
most nearly represents Holocene or interglacial conditions. It is probable,
therefore, that conditions were relatively warm as well as relatively wet.

TABLE 29

Levels loading highly on factors in the Varimax rotated factor matrix (A) and Oblique rotated
factor pattern matrix (B) for Upper Pleistocene and modern samples.

Factor 1 Factor 2 Factor 3 Factor 4

A:

BPA: YOL 0,86001 BPA: OLP3 0,71777 DK1 M.S.A:2 0,77182
BP2 0,82808 DK1M.S.A: 3 0,80435
BP3 0,89820 4 0,95147
BP4 0,82391 5 0,91220
OLP 0,76818 6 0,94493
OLP1 0,73770 7 0,94014
BOL 0,76889 8 0,95415
BOLI 0,96818 9 0,91429
BOL4 0,93851 10 0,88617
BOLS 0,88003 11 0,94384
OCH 0,91062 BNK2 0,77737
LOH 0,90113 BPB-C 0,89406

KRMIA: 15 0,71570 NGA-B 0,82756

DKI1 M.S.A:13 0,85308

NBC: YSL 0,87318
YGL 0,88311

B:

BPA: OLP3 0,98289 NBC: YSL 0,71206 DK1 M.S.A:2 0,91355 KRM1A:15_ 0,88213
BPI 0,97444 32 0,87497
OLP2 0,96506 BPA: BOL4 0,86883
OLPI 0,88768 BOLI 0,80520

DKI M.S.A: 4 0,87412 BP3 0,71704

5 0,85567
7 0,84874
8 0,84319
6 0,83297
11 0,81468
9 0,80619
10 0,79499

BPA: OLP 0,79118
BOL 0,74424

BPB-C 0,74415

BPA: BP4 0,73993

92,8% of the total variance accounted for. Moderate correlation between Factor 1 and Factors 3 and 4, other-

wise low. ’

KRMIA level 15 loads most highly on Factor 4 (Table 29). High propor-
tions of Otomys irroratus and moderately high proportions of Otomys saunder-
sae distinguish this level. The only other species to occur in some quantity in
these levels are Crocidura flavescens and Myosorex varius. It would seem that
wet and fairly cold conditions are indicated, with extensive dense vegetation at
the waterside and a certain amount on the lower hillsides. The BPA levels that
load highly on this factor are some that load highly on the glacial maximum

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS pa |

factor in the Boomplaas A analysis. It is possible that the difference between
this factor and Factor 2 is largely topographic. It is probable, however, that
rather warmer, wetter conditions are represented by Factor 4 compared with
those represented by Factor 2. Certainly Crocidura flavescens would not nor-
mally be expected in high proportions in very cold conditions. It is also
probable that the apparent anomalies in the KRMIA data are due to the
incompleteness of the available samples. The moderate correlation between
Factor 1 and Factors 3 and 4 is probably due to the fact that variations are in
proportion rather than in species.

EVIDENCE FROM MEAN SIZE VARIATION

That populations of certain species tend to vary in mean size of individual
members is an observed fact. It is particularly suggested that these changes
represent responses to differing temperatures. Bergmann’s Rule, as has been
noted above, states that within a particular species those individuals living in
warmer climates will tend to have a smaller body mass than those individuals
living in colder climates. The converse of this should be that if there is variation
in mean body mass of populations living in the same place but at different times
in the past, this must indicate warmer or cooler climates at the relevant times.
In other words, if body mass can be correlated with temperature it should be
possible to use this method to provide direct evidence of past temperature
fluctuations. In fact, Kowalski (1971) quotes a study by Sych (1965) in which
increase in the body size of a fossil hare from Poland is shown to indicate a
gradual decrease in mean temperature of about 10 °C during the Pliocene and
early Pleistocene.

In order to achieve such results it is clear that basic data must be available
for modern representatives of the species concerned. Initially it must be
established whether or not the species does, in fact, behave according to
Bergmann’s Rule, since not all species do. If the species exhibits a consistent
negative or positive response, it is necessary then to ascertain the scale of
changes involved. Such an adequate data base is not currently available for the
species being examined in the present study, so that a quantitative assessment
cannot yet be made. The subfossil records show, however, that real changes
have taken place which can be correlated in a broad way with known climatic
change. In particular, the shrews Crocidura flavescens and Myosorex varius
provide good evidence. Limited data for Tatera afra suggests that this species
may also be useful but the trend in Aethomys namaquensis does not appear to
be correlated with known climatic fluctuations.

The nature of the evidence forthcoming from Cryptomys hottentotus 1s
more problematic. The mean size of individual in this species varies quite
considerably in samples of different ages. It would appear, therefore, that once
it has been established what causes the changes, C. hottentotus should be of
considerable use in palaeoenvironmental interpretation. Climatic change may
normally be reflected in temperature change, but Tchernov (1968) has shown

278 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 30

Size variation in mandibles of Crocidura flavescens from Boomplaas A and B.

A: height of ascending ramus B: depth of body mandible ,
N _ Range X S 95% N Range x S 95%
BPA: DGL . « 70 6,9-8,7 7,74 0,36 0,09 106 2,4-3,5 2,80 ° "0.23 0,04.
BLD3 . . 70 7,2-8,5 7,80 0,31 0,07 131 2,5-3,5 2,90 0,21 0,04-
BG Se 6S Sho Ta 7,88. “O38 .-0;11 69 2,5-3,4 2,95 0,21 005m
BLA «2 6 36-74-90 8.12. .0,37. .-0:13 54 2,5—3,4 3,02 0,19 0,05—
BRL. . . 72 _ 7,4-9,2 8,25 0,38 0,09 96 2,6-3,8 3,10 0,23 0,055
Cl a aes DOP ea 8,34 0,39 0,10 56 2,7-3,7 3,16 0,23 0,068
GWA . :. @& &2-9,2 8,43 22 3,0-3,5 3,25 0,17 OGM
LP 2. «679289 8,41 0,35 0,19 69 2,9-3,7 3,24 0,22 0,059
YOU. «=. «#7690 8,60 8 2,9-3,6 3,25 0,29 (Game
BPA. .-:. 8 8,0-9,0 8,48 0,44 0,37 29 2,7-3,5 3,05 0,17 Om@
OLP*. 0. -« “G6! "80292 8,52 0,45 0,47 12 2,/-3,3 3,01 0,19 0,12
OLP1 2 S64 59-92 8,63 0,46 0,48 S 3,0-3,4 3,22 0,18 O23
OLP2. .. «. 26- F795 8,44 0,44 0,18 40 2,7-3,6 3,19 0,19 0,06
OLP3 > = WS? 82-88 8,47 0,23 0,14 42 2,8—3,6 Sut 0,23 0,0;
BOL se & EL. 8732590 8,17 0,50 0,33 31 2,6-3,5 3,03 0,21 0,08
OCH aw ty S12 - - FOES 8,38 0,30 0,19 19 2,9-3,6 3,13 0,18 0,09
Pe ke me OO e ul 8 7,10 -0,40 ~ 041 1 2,5—3,2 2,80 0,23 Off
Measurements in mm. 95% = 95% confidence interval for the mean.

TABLE 31

Size variation in mandibles of Crocidura flavescens from Boomplaas A upper levels.

A: height of ascending ramus B: depth of body mandible

N Range xX S 957 N Range x Ss 95%
Unit DGL
PRGL TS . . ks eee 215 7,6-8,5 S01 0:26" ~ 0:15 14 2,7—-3,4 2,92 0,20 . Om
"Sy Best heen a ewe | 6,9-8,7 7,66 0,39 # 0,14 44 2,6-3,5 2,88 0,20 0,06
BER SS 6 ees 9 7,2-8,3 7,66 0,33" 0235 16 2,0-3,2 2.73 0,30 Of
BE pA xo 8 7,3-8,5 ids OAO 023 12 2,4-3,3 2,94 0,25 0,16
BED A oh). ees 7 7,3-7,8 7.63. O,16, —“O.95 20 2,6-3,1 2,81 0,15 O@
Unit BLD3
BR4ie c- bie see 39 7,3-8,5 Ta? =6 O34 sO ,10 (6 2,4-3,4 2,85 ‘- 0,20 O06
BEDSA*. ©. 82. oe 1 3! 7,2-8,5 7,84 0,32 0,12 56 2,5—3,) 2,96 0,23 0,06
Unit BRL
BRL fi et eek oles 7,5-8,8 8.23 “0,38: 0,190 2 22 2,6-3,3 3,07 0,19 Ofte
BRE se, « 7,6-8,9 8,18 0,34 0,18 25 2,7-3,7 3,04 0,20 0,08 |
BREAMA.. & «24% 6 7,7-8,4 8:15. 0,27. . 0,29 14 2,7-3,3 3,01. 0,19 Off
Pe te a ae ek Sb 7,7-8,7 8,12 038 “025 11 2,8—3,4 3,03 0,18 Of@
BELG) ees 5:.5 2, 2 7,4-9,2 8,41 0,42 0,19 24 2,6-3,8 3,22 0,28 Om
Unit CL
Petr 5 ke Sy ce 7,9-9,0 8,46 0,39 0,19 20 2,8-3,6 3,25. 0,21 Gam
Cer ae le ers 7,9-8,7 8,18 0,30 0,18 16 2,7-3,6 3,16 0,20 0,11
Ce Sy i vie 9 8,1-8,9 8,50 0,31 0,24 8 2,8-3,7 3,20 0,31 Of
CURB eg os 9 7,7-8,8 §,13.. 0,43 0.33 12 2,8—3,1 2,98 0,11 0,07

Measurements in mm. 95% = 95% confidence interval for the mean.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 279

~

C: length of M,-3; D: length of lower tooth row E: length of mandible plus incisor
N_ Range 4 s 95% N__ Range x s 95% N_ Range 4 s 95%

30 5,6-6,3 5,91 0,18 0,07 6 11,4-12,6 12,08 0,40 0,42 9 16,8-19,2 18,32 0,85 0,28

51 5,7-6,5 6,04 0,19 0,06 &. P15-13,1- 42,26: -0,53.-°0,45 © 17,4-19,3° 17,97 0,72 0,75

39 5,7-6,3 6,04 0,17 0,06 13 11,8-13,1 12,27 0,38 0,23 25 16,5-20,0 18,40 0,77 0,32

. 5,7-6,7 6,20 0,26 0,10 6 11,9-12,4 12,00 0,31 0,33 9 17,1-19,1 18,12 0,66 0,51

65 5,8-6,8 6,27 0,22 0,05 19 12,0-13,6 12,71 0,43 0,21 20 17,9-20,5 19,19 0,73 0,34

30 6,0-6,8 6,43 0,21 0,08 11 13,0-13,9 13,30 0,27 0,18 11 19,0-20,9 20,06 0,72 0,48
8 6,3-6,9 6,54 0,21 0,18

m 6,2-7,.0 6,55 0,25 0,13 f Cs) 3 19,0-20,4 19,70
4 6,2-6,7 6,53 2 12,2-13,6 12,90 3 18,3-20,6 19,40
6 6,0-6,6 6,35 0,25 0,26 P88)
3 6,3-6,5 6,47 1. “3,1)) 2 19,9-20,4 20,15
4 6,3-6,7 6,45 t. (18;7)
22 6,2-7,0 6,56 0,19 0,09 6 13,1-14,2 13,47 0,50 0,52 6 19,5-21,3 20,45 0,67 0,70
18 6,1-7,0 6,48 0,28 0,14 8 12,6-13,9 13,35 0,47 0,40 8 19,0-21,2 20,29 0,88 0,74
8 5,9-6,5 6,24 0,25 0,21 2 AD H13.2)~ 12.95 3 18,4-19,6 19,10
9 6,0-6,8 6,27 0,23 0,18 3 123-13,0' 12,67 3 19,2-19,7 19,53
5 5,7-6,2 5,88 0,19 0,24 = 15-120 11,75 4 17,4-18,3 17,83
C: length of M,-; D: length of lower tooth row E: length of mandible plus incisor

-N Range X S 95% N_ Range 4 s 95% N_ Range % 6 - 25

= 35,8-6,1 5,95 4 (12,3) 4 17,3-19,0 18,33
ome os 599 0,18 0,10 3 11,4-12,1 11,87 3 16,8-19,2 17,93

4 5,5-6,2 5,85

4 5,8-6,0 5,93 2 12,0-12,6 12,30 2 18,6-19,2 18,90

5 5,8-6,0 5,88 0,11 0,14

go 5,7-6,4 6,03 0,19 0,07 6 11,5-13,1 12,22 0,57 0,60
=m 3,/-0,5 6,05 0,21 0,11 2 12,0-12,8 12,40

17,5-18,0 17,67
17,4-19,3 18,27

Ww Ww

14 5,8-6,4 6,19 0,17 0,10 12,4-12,9 12,70
m 5,9-6,8 6,21 0,22 0,11 12,2-13,5 12,68 0,44 0,18

3 18,6-19,5 19,08
6
12 6,0-6,7 6,28 0,26 0,16 5 12,0-13,0 12,42 0,45 0,49
2
3

18,2-19,7 18,90 0,53 0,56
17,9-20,2 18,77
19,2-19,5 19,35

T 6,1-6,5 6,31 0,18 0,16 12,7-13,0 12,85 ’
18,8-20,5 19,82 0,68 0,84

m 5,9-6,6 6,37 0,22 0,12 $2,9-13,6, 13,17

UANWAHA

mete—6.6 642 0,21 0,16 2 13,2-13,3 13,25 5 19,0-20,9 19,94 0,83 1,03
1) 6,0-6,7 6,39 0,23 0,17 5 13,0-13,6 13,24 0,26 0,32 2 19,0-20,7 19,85

= 6,2-6,7 6,50 0,25 0,30 3 13,1-13,9 13,40 3 19,8-20,9 20,30

6 6,3-6,7 645 0,14 0,15 1 (3,4) 1* (20,3)

280 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 32

Size variation in mandibles of Crocidura flavescens from coastal sites

A: height of ascending ramus B: depth of body of mandible

N Range xX S 95% N Range x S
eG Bae i 6,9-8,2 7,54 0,36: 0,14 31 2,5-3,2
| ere 16 7,2-8,3 7,75 0,41 0,22 32 2,5-3,2
fe A 7,7-8,0 7,87 8 2,7-3,1
M.S.A.4 ... 6 6,9-7,6 7,33 0,24 0,25 V1 2,5-2,8
ee 10 7,3-8,4 ido O30 OI 15 2,6-3,1
10 3 7,3-8, 1 767 © O35 ~O43 8 2,6-3,2
ERMIA? 6 <i> 2 (8,5) | 3,1-3,5
| ae 6 8,0-8,9 §.37 039. 0,41 Ey 2,9-3,6
ty ae 4 7,2-8,4 7,80 14 2,7-3,4
Sve pet 3 7,3-8, 1 7,70 8 2,7-3,4
NBG {Yok : ot 4 13 8,1-8,7 8,41 0,19 0,11 47 2,5—351
MmeEBas ee 17 7,4-9,1 §,21— -0;46- 0,23 37 2,8-3,5
BNE ve. S40 72> F525 6,6-7,8 1,29 10:32". O}13 38 2,1-3,2

Measurements in mm. 95 = 95 % confidence interval for the mean.

that in the case of Spalax ehrenbergi (Palestine mole-rat) in Israel the correla-
tion is with rainfall. Demonstrable changes in Cryptomys hottentotus in the
present study may perhaps also be attributable to rainfall fluctuation. Data on
such a possible correlation have not yet been accumulated (J. U. M. Jarvis 1979
pers. comm.) but De Graaff (1981) notes a tendency for this species to be
geographically variable in size.

Crocidura flavescens

Five parameters were measured in Crocidura flavescens as was described
above. Tables 30-32 give the resultant data for different populations and
Figures 16-21 present the same data in graphic form. Figure 15 illustrates the
variation in the means of the various parameters throughout the sequence at
Boomplaas A. It is clear that there is considerable variation in the mean sizes
of the various populations and, in particular, the good sequence at Boomplaas
A shows clear trends.

Figure 15 shows that the pattern of size change in the mandible of C.
flavescens at Boomplaas A is very similar in all the parameters measured. This
would suggest that, in fact, the mandible was changing in overall size rather
than in one aspect alone. Until such time as the nature of the relationship of
the size of the mandible to overall body mass is established for this species, it
will be assumed that a larger mandible indicates greater body mass. In general
terms, on this basis, C. flavescens is shown to be larger during the Upper
Pleistocene than during the Holocene. The only exception is level BOL where
the means are smaller than those for the earliest part of the Holocene
sequence. Larger samples for level BOL may clarify this situation and it will be
important to discover whether the lower BOL levels follow the same trend or

RB | Za

-—* i

ae Dee Os
IB UWON KENNA

8

: length of M,-;
Range

5,6-6,0
5,4-5,7
5,8-6,0
(6,0)

6,2-6,6
5,9-6,4
6,1-6,4
6,0-6,4
6,2-6,8
5,7-6,7

5,1-6,1

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 281

D: length of lower tooth row E: length of mandible plus incisor
4 s 95%: N ~ Range X 5. “25%, N_ Range XK gs’ 95%

cet? 008 23 10,9-12,2 11,70 0,39 0,17 31 16,4-18,7 17,70 0,63 0,23
3,87 0,19 0,08 10 11,3-12,7 11,89 0,47 0,34 15 16,5-19,2 18,11 0,79 0,44

5,86 0,16 0,15 > 10;6-12,1 11,62 0,61 0,76 3 18,0-18,4 18,27
Pb) 3 16,8-17,4 17,07
5,85 1 (11,5) a 17,5-11,6 17AT
1 (12,0) 2 47,9-18,3 18,10
6,40
eeu 0,17 0,13 4 12,1-12,8 12,43 6 17,9-20,6 19,28 0,88 0,93
6,23
6,14 0,17 0,21 3 12,3-12,5 12,40 3 18,4-18,9 18,67
6,44 0,18 0,10 4 12,7-13,8 13,40 0,48 0,77 5 -18,6-21,0 -20;26 0,95 -1,18
6,40 0,29 0,15 3 13,3-14,0 13,60 7 18,7-20,8 19,89 0,66 0,61

3,05 0,25 0,10 19 10,4-12,2 11,50 0,50 0,24 24 15,9-18,3 17,16 0,62 0,26

whether specimens from these levels will be larger. In the length of M,_3 and
depth of mandible means are highest during the period from about 25 000 B.P.
to about 14 000 B.P., that is the last glacial maximum (Figs 16-17). In the
height of the ascending ramus the mean is higher at the beginning of this
period. In all three parameters an earlier high occurs in levels OLP1 or OLP2
when means are highest for length of lower tooth row (Fig. 18) and length of
mandible plus incisor (Fig. 19). The data for level OLP2 are reliable, being
based on a good sample, but those from level OLP1 require improvement. In
view of the correlation between large size and glacial maximum, it seems safe
to assume tentatively that the species is behaving in accordance with Berg-
mann’s Rule. This being the case, it would appear that there is an indication
during the Upper Pleistocene of two main cold periods separated by a milder
period, and that the Holocene was considerably warmer than almost all of the
preceding period represented at Boomplaas A.

From about 14 000 B.P. onwards there is a continuous trend towards
smaller means in the various parameters up to and including the modern
sample from Boomplaas B. That this trend is far from smooth is shown by the
means for the upper levels at Boomplaas A (Figs. 20-21). There are perhaps
two main periods of fluctuation. The first is in the lowest postglacial levels,
CL3BG to BRLS5S, and the second is in the uppermost levels, BLD2A to DGL1,
and Boomplaas B. In the first it is the height of the ascending ramus that
fluctuates most noticeably. This parameter displays moderate coefficients of
variation (Table 33), which suggests that real fluctuation rather than changes in
population composition are being monitored. Such fluctuations are perhaps to
be expected during a period of climatic change. During the later period of
fluctuation, on the other hand, the greatest amplitude is found in the depth of

282 ANNALS OF THE SOUTH AFRICAN MUSEUM

M.varius

C.flavescens

abcde fghj |!mnopqs a b cde f abcdefghijkIimnopaqrst
1234512 123451234

a DGL hy EP o OLP2

b BEDS i J JEBG p OLP3 al DGLI b2 BLD3A e4 BRLS

¢ BL j YOL q BOL a2 DGL2 e BL e5 BRL6

d BLA k BPI r BOL4/5 a3 BLD d BLA fl BRET

e BRL | BP4 s OCH a4 BLD2 el BRL f2 Ck

f. 6E m OLP t LOH a5 BLD2A e2 BRL2 ts CE2

g GWA n OLPI bl BLD3 e3 BRL4/4UA f4 CL3BG

Fig. 15. Variation in the means of parameters measured in Crocidura flavescens and Myosorex
varius from Boomplaas A. See Tables 30 and 36 for details.

283

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS

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284 ANNALS OF THE SOUTH AFRICAN MUSEUM

NBC -YSL ee eee

YGL |}; 17
KRMIA-6 +—_}— 2
15 |—__—_() 9
22 +—_}+—— 3
32 5
BNK2 eee)
DKILSA- I2 | 7
BNKI-5 E423
19 nj —— 24

DKIMSA-4 +—_}|— 2
6 ae 4
fe) Dat.

BPB a 5
BPA- DGL So —— 30
BLD3 al
BL a1 39
BLA a
BRL ($$$ aa 65
CL a 30
GWA I ——i 8
LP a
YOL W—--.-f- 4
BP4 a =
OLP r—h} 3
OLPI +—_}—_—_ 4
OLP2 (+ Soo 22
OLP3 —— So 8
BOL 6
OCH a
mm 5| 53 55 57 59 6! <63 65 ‘G7) scam

Fig. 17. Variation in parameter C, length of M,;, in Crocidura flavescens from various sites.

the mandible. This parameter has a much higher coefficient of variation
(Table 33), presumably because it is affected by the age of the individual. It is
therefore much more likely that differences between populations are due, at
least in part, to changes in the composition of the population. In other words, if
there were a higher proportion of young individuals the mean would be lower
than if there were a higher proportion of older individuals. The fact that there
is a tendency for the percentage of C. flavescens in the sample to be higher
when mean depth of mandible is less (Fig. 21) may suggest a population
explosion when the proportion of younger individuals was unusually high. On
the other hand, the fact that changes are mirrored, albeit less obviously, in the

285

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS

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286 ANNALS OF THE SOUTH AFRICAN MUSEUM

E
NBC-YSL | $s
YGL nS
KRMIA - 6
15 =
22
32 1 3
BNK2 Sa
DKILSA-12 r—H 3
BNKI-5 (eo 5
19 + a5
DKIMSA- 4 +—}— 3
6 3
10 tH 2
BPB +——_}|—__ 4
BPA-DGL - —— aa, 9
BLD3 SAR
BL Sos os
BLA | Sa 9
BRL | —_____——— 29
CL a ee
GWA
Pe -———_}———_4 3
YOL — } a
BP4 =
OLP +—{+—; 2
OLPI [st
OLP2 | i 6
OLPS es =
BOL —— | 2
OCH nnd

mm 157 16! 165 169 173 17,7 18,1 18,5 18,9 193 197 20! 205 209 213

Fig. 19. Variation in parameter E, length of mandible plus incisor, in Crocidura flavescens
from various sites.

other parameters, indicates real change must also have been taking place. In
particular, the length of M,_3, which has low coefficients of variation (Table
33), shows this to be the case. Of some interest, within the fluctuations, is the
fact that levels DGL1 and DGL2 indicate a reverse trend which suggests a
cooler period at the top of the sequence.

In general, the differences between means of one Spleens and those of
preceding or succeeding populations are not statistically significant. This would
suggest that the rate of change was normally relatively slow. On at least two
occasions, however, there would appear to have been an increase in the rate of
change. Given the one-sided hypothesis that successively younger levels in the
postglacial period should contain populations of smaller individuals, the differ-
ences between levels BRL6 and BRLS, and levels BLD3 and BLD2A are
significant (Table 34). In both cases this represents quite rapid change over

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 287

periods of a few hundred years. In the first case, means for level BRL5 are
significantly smaller than those for level BRL6 in height of ascending ramus and
depth of mandible but not in length of M,_3. The mean for this parameter is,
however, significantly smaller in level BRL2 than in level BRL7, which also
constitutes quite rapid change. The means for height of ascending ramus and
length of M,_3 in level BLD2A are significantly smaller than those in level
BRL3, but not for depth of mandible. Similar periods of accelerated change
could well occur in the lower levels but their detection must await analysis of
more detailed samples.

The data for other sites are meagre but they do tend to confirm the picture
provided by Boomplaas A. At Byneskranskop 1, for instance, the means for
level 5 are smaller than those for level 19, except in the depth of the mandible.
In this parameter the mean for level 5 is very slightly larger than that for level
19 but this is not considered of any great importance because of the relatively
high coefficient of variation, as was discussed above. The means for all
parameters in the modern sample from Byneskranskop 2 are considerably
smaller than in either of the subfossil samples. The samples from Die Kelders 1
are generally very small and the impression is that the M.S.A. levels are more
nearly comparable to the BNK2 sample than the BNK1 samples. The means for

DGLI DGL | aS A
DGL2 (SSS

BLD ee
BLD2 ee ee
BLD2A a _
BLD3 BLD3 (—SSSSE EE) 39
BLD3A |__| 6
BLD3AM ES - ——_. 17
BLD3AM2 CE ¢
BL eG 51
BLA b|{$- rr 4 36
BRL BRL |+—________ se ig
BRL2 |} ___——— 17
BRL4-4A | $i
BRL5 | > 2S
BRL6 F ee 20
BRL7 CL | —__ e g
_ = 13
| | _ i 9

CL3BG “es

mm Cris 7s 75 7%) FO Bf 8S ° 85.- 67 89: oi 93
Fig. 20. Variation in parameter A, height of ascending ramus, in Crocidura flavescens from

postglacial levels at Boomplaas A.

ANNALS OF THE SOUTH AFRICAN MUSEUM

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MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 289

TABLE 33

Mean coefficients of variation in parameters measured in Crocidura flavescens, Myosorex
varius, Aethomys namaquensis, Tatera afra, and Cryptomys hottentotus.

A B C D E
emavescemS =. 5 ne wks 4,51 6,61 3,44 3,54 3,90
ee ls 3.53 2,61 3.27 2,87
mmere@guensis . . 5. fk es 3,24 3,58 8,67
T. afra es a eee cer 3,08 Si)
CGeNGeMPOWIS 5 kk ll le 4,90 4,65 8,06 11,84 10,91

See Tables 30 and 36-38 for details of parameters and basic data.

TABLE 34

Selected ‘t’ tests for Crocidura flavescens.

id d.f. P(1-sided)
BPA: BLD3/BLD2A
A Sell aN a a5 44 0,005—0,0005
B I a we ae 0,83 93 N.S.
c TINS Wr ey ce ik ce we 1,71 38 0,05
BPA: BRL7/BRL2
c. Se ee 235 24 0,025
BPA: BRL6/BRL5
A NS 1,90 29 0,025
B I ee ee Oe as 2,06 33 0,025
C ey he aS SS ie 0,63 20 N.S.

* See Table 30 for explanation of parameters and data on which the ‘t’ tests were based.

the L.S.A. level, on the other hand, are higher than any for Byneskranskop or
other DK1 levels.It would appear that the M.S.A. levels were deposited during
a relatively warm period, but that about 2 000 B.P. (L.S.A. level 12) the area
was colder. The samples from Klasies River Mouth 1A show a gradual increase
in the means for the various parameters from earlier to later. This is consistent
with the apparent pre-Last Glacial age of these deposits. Moreover, levels 9
and 6 have provided means for the non-dental parameters which are even
larger than those from Nelson Bay Cave. This is surprising in view of the fact
that the two NBC levels were deposited during the last glacial maximum and
the means for the parameters are comparable to the greatest at BPA. This
would suggest that conditions were already cold in the upper part of the
KRMIA sequence, although larger samples are needed before the situation can
be verified.

Since the original hypothesis was that geographicaily separated contempor-
ary populations may vary in mean size of individuals, it follows that a similar
situation is likely to have pertained in the past. It also draws attention to the
fact that evidence concerning temperature is restricted in its application. It may

290 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 35

Percentage difference between selected populations of Crocidura flavescens and Myosorex
varius at Boomplaas, Byneskranskop, and Nelson Bay Cave.

C. flavescens

A* 2 é 0,24
B 7,28 4,24 12,46 2,86 1,25
Cc 4,07 4,50 8,86 2,34 1,55
D 2,00 4,87 M35
E 3,90 3,95 9,61
xX 4,57 4,41 9,57 255 1,01
BPB/BPA: CLI BNK2/BNK1: 19 BPB/BPA: BL BNK2/BNK1: 5
A 6,23 6,31 2,34 543
B 12,86 7,66 5,36 8,43
© 8,67 3,89 QZ 2,30
D 12,87 3,89 4,60 1,74
E ies 5,54 3,20 — 3,05
xX 10,39 5,36 3,64 PENG |
M. varius
BPB/BNK2 BPA:BL/BNK1:5
A 4,67 ese.
B 6,63 3,10
C 4,19 1,69
xX 4,50 4,04
*

See Tables 30 and 36 for details of parameters and basic data.

be possible to extrapolate on the basis of known present-day differences. The
evidence tends to suggest, however, that the relationship has not been static in
the past, which would make such extrapolation potentially misleading. Com-
parison between approximately contemporary levels at Boomplaas A and
Byneskranskop 1 and at Boomplaas A and Nelson Bay Cave illustrates the point
(Table 35). It is noticeable that on average the means for Boomplaas A are
9,57 per cent higher than those for BNK1 in the early postglacial period. By
about 4 000 B.P. the BPA means are only 4,41 per cent higher. Thereafter, the
position appears to have stabilized because the average for the modern BPB
sample is virtually the same at 4,57 per cent higher than that for BNK2. With
BPA and NBC the situation is not so clear, possibly because of small samples
or because the levels cannot be closely correlated or because both were inland
sites at the time. However, although the difference between BPA and NBC is
much smaller than that between BPA and BNK1, the same point is evident; the
percentage difference is not consistent throughout. The present evidence sug-
gests that the climate was relatively either much cooler or more extreme in‘the
Cango valley than around Byneskranskop about 12 000 years ago than it is
today. This is indicated not only by the greater difference between the two
older levels, but also by the fact that in the Cango valley the BPA level CL1
population is on average 10,39 per cent larger than the BPB population (Table
35). This contrasts with Byneskranskop where the BNK1 level 19 population is
on average only 5,36 per cent larger than the BNK2 population. Even such
initial exercises illustrate the potential of such work for providing detailed
regional climatic information.

Myosorex varius
Four parameters were measured in Myosorex varius as was described
above. Table 36 gives the data for the different populations and these are

291

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS

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294 ANNALS OF THE SOUTH AFRICAN MUSEUM

shown graphically in Figures 22-23. In Figure 15 variation in the means for the
different parameters in the Boomplaas A sequence is plotted. As with Croci-
dura flavescens, there is considerable variation, with the pattern being clearest
in the sequence from Boomplaas A.

The pattern of change is very similar in all parameters. This is not
unexpected since three of the parameters were, in effect, alternatives. For this
reason the best represented, length of P, to M3, will be taken as representative
of the three length measurements in the following discussion. The coefficients
of variation for this parameter are low and are only moderate for height of
ascending ramus (Table 33). It would seem, therefore, on arguments put
forward above, that real change in the mean size of individuals is being
monitored. This change would appear to have been generally gradual; certainly
there are as yet no demonstrable periods of accelerated change, although these
may be found in the lower levels later.

There are several major trends noticeable in the Boomplaas A sequence.
From the base there is a general decline in mean size for the various parameters
until level BP4. Thereafter, a moderate increase in levels BP1 and YOL leads
eventually to a decline culminating in level CL. A general increase in the early
postglacial period is replaced in level BLA or BL by a slight decrease. As a
generality most of the Holocene levels contain populations whose mean man-
dible size is larger than those in most of the Upper Pleistocene populations.
This would suggest that Myosorex varius displays a negative response to
Bergmann’s Rule. If this were the case, the evidence would suggest that
temperatures in the lowermost levels were comparable to those of the
Holocene. By the same token, the coldest periods in the sequence would be in
levels OLP-BP4 and again in level CL. This picture is not entirely in accor-
dance with that suggested by other lines of evidence and it may be that there is
some other explanation for the size variation. Possibly rainfall or effective
precipitation may play a part in determining the size of individuals. It is,
however, also possible that the apparent anomalies may be due to lack of
information and that when more complete data are available the situation will
be clarified. The limited data from other sites indicate that this is perhaps most
likely since results vary inversely to those obtained for Crocidura flavescens.
Table 35 shows that the percentage difference in mean size of individuals in
contemporary samples from Boomplaas and Byneskranskop is virtually identi-
cal to that found in C. flavescens. However, Myosorex varius is smaller inland
than on the coast, whereas the reverse is true for Crocidura flavescens. The Die
Kelders 1 M.S.A. levels show a reduction from earlier to later mean individual
size, with no change in the L.S.A. sample. This is not the complete reverse of
the situation in C. flavescens, although it tends to hold good in the M.S.A.
sample. Larger samples of C. flavescens would almost certainly have clarified
the situation. It seems in general, therefore, that the size of the Myosorex
varius will be shown to vary in response to changing temperature, but in
contradiction to Bergmann’s Rule.

Oe, SO ee ee ee eS ee Oe 7? Oe

Aethomys namaquensis and Tatera afra

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS

295

Aethomys namaquensis was measured in three parameters and Tatera afra
in two, as was described above. Tables 37 and 38 give the data on size variation
in these two species and Figures 24 and 26 show these data graphically. Figure
25 illustrates variation in the means of different parameters for A. namaquensis
at Boomplaas A. The mean coefficients of variation for dental parameters are
fairly low in both species, but depth of mandible in A. namaquensis is very
much higher (Table 33). This is probably due to the fact that this parameter is
likely to be influenced by the age of the individual and possibly the sex as well.
The size of the teeth, on the other hand, is apparently little affected by either
of these factors.

Size variation in teeth and depth of mandible in Aethomys namaquensis from the Cango valley.

A: length of M?-*

N

BPA: DGL . 23
BLD3 . 26

BL 19

BLA 14

BRL 136

CL 30

LP. 5

BP4 6
OLP1 =
OLP2 36
OLP3 34

BOL =

NGB «17

Measurements in mm.

BNK2_.
DKIMSA:

i ae
NBC: YSL A 3-5.
YGL B 1-4.

Measurements in mm.

Range

5,1-5,8
5,2-5,9
5,4-6,0
5,3-5,9
5,1-6,0
5,2-6,1
5,5-5,9
5,270.9
5,6-6,1
5,5—-6,4
5,6-6,4
5,8-6,2
3,1-5,9

X so" 957,

TABLE 37

B: length of My_;

N

Range

5,1-5,7
5,0-6,0
5,1-5,9
5,3-5,7
5,1-6,0
5,3-5,8
(5,7)

5,1-5,8
5,4-5,8
5,4-6,0
5,4-6,0
5,5-6,1
4,7-5,8

Xx

C: depth of body of mandible
s 957,

S/, 957, .N
0,20 0,08 43
0,17 006 34
0,17 0,06 31
0,18 0,12 8
0,18 0,04 83
0,16 0,07 28

5
0,30 0,38 10
O13 10 12
0,16 0,06 35
0.17 0:07 30
0:26: 0,27 10
0,30 0,15 18

Range

95% = 95% confidence interval for the mean.

TABLE 38

Size variation in teeth of Tatera afra.

A: length of M,-2

N Range xX Ss
9 5,0-5,2 5,09 0,09
a = 90-55. 3,28. 022

18 4,7-5,3 4,99 0,16
3 *(5;2)

18 4,8-5,2 5,04 015

22 4,8-5,4 5,13 0,16

2 48-53. - 5,13... O12

10 5,0-5,4 5,22 0,15
9 4,8-5,4 5,09 0,16
1 (4,9)

38 §65,3-6;0 5,63 0,19

Sie 3°63 85,76. 0,22

95.7%

0,07
0,27
0,08

0,08
0,07
0,05
0,11
0,12

0,06
0,08

B: length of M!~-?
N__ Range x
3 4,7-4,8 4,77
4 5,1-5,6 5,30
7 4,6-5,2 4,96
6 4,8-5,2 5,00
19 4,8-5,3 4,95
38 §©64,7-5,3 4,93
34 4,6-5,3 4,91
14 4,5-5,1 4,88
18 4,7-5,1 4,92
3. ~ 49-533) > S40
48 5,1-5,7 5,43
48 4,9-5,9 5,46

95% = 95% confidence interval for the mean.

Xx

B04
3,19
3,90
4,11
3,88
3,94
3,40

0,12

0,14
0,19

296 ANNALS OF THE SOUTH AFRICAN MUSEUM

It is unfortunate that samples of A. namaquensis are available only from
the Cango valley and of 7. afra only from the coastal sites. For this reason it
has not been possible to check possible trends in contemporary sites. It would
appear, however, that in A. namaquensis there is a general reduction in the size
of the teeth from the earliest levels at Boomplaas A to the latest; the mean size
of the teeth in the Nooitgedacht B sample is also low (Figs 24-25). This trend
does not seem to be correlated with temperature change and, as such, is of little
value in the present context, although it is almost certainly of intrinsic interest.
The depth of the mandible in A. namaquensis provides a very clear pattern
(Figs. 24-25) which suggests that, like M. varius, this species may respond
negatively to Bergmann’s Rule. In this parameter the mean size in postglacial
populations is noticeably larger in most cases than it is in the glacial popula-
tions.

NGB ES —— 18 Cc
BPA-DGL SS eee

BLD3 [_— rr ——— _ 34

BL [}_—— _ a

BLA | i —< g

BRL (ee Fg

SL (> +28

LP 2 -

BP4 + 0

OLPI eee

OLP2 (SSS 355

OLP3 (SS 30

BOL a | 0

- B
NGB RS 7 Ss -— 18
BPA- DGL [—_ a 23 SS 27
BLD3 |+—__i—)— 26 p< 34
BL [S19 ———. a =
BLA [+ 14 a
BRL 5-136 (> 88
Ck SE — 30 + 23
LP Te = i
BP4 | CR = | Ts — 5
OLPI a = EE 9
OLP2 -_ a —_—_— 36 —_ ES 33
OLP3 ————_ — —— -—a=—— 28
BOL +—_}— 4 a =
5| 53 55 57 59 6! 63 mm4r 49 5,0 5,5 55 " Saeeaumees

Fig. 24. Variation in parameter A, length of M'“, parameter B, length of M,;, and parameter
C, depth of mandible, in Aethomys namaquensis from the Cango Valley.

DGL
BLD3
BL
BLA
BRL
CL
GWA
LP
YOL
BP4
OLP
OLPI
OLP2
OLPS

=a “oe & Oo 18

ago =

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 297

fC)
Pp
al
a2
a3
a4
el
e2
e3
ea
e5
fl
f2
tS

BOL
OCH
DGLI
DGL2
BLD2
BLD2A
BRL
BRL2
BRL4
BRLS
BRL6
BRL?
CLI
CL3BG

A.namaquensis

A

mm

6,0

5,8

5,6

3,5

3,3

abcdefhj lmno

C.hottentotus

abcdefghijkImnop

a
1234

bede f
12345123

Fig. 25. Variation in means of parameters measured in Cryptomys hottentotus and Aethomys
namaquensis from Boomplaas A. See Tables 37, 39-40 for details.

298 ANNALS OF THE SOUTH AFRICAN MUSEUM

2 @
¢ 8

RM

DAY OID
DIY OW
BIB GW

>
nN
oO
fs
o
o

QIU
DPD
BEY
AHO)
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45
35
OLP3 BPB NGA-B

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15

B
5
25 234016 123456 123456 123456
BRL LP BP4 OLPI
15
: a. i.
123456 123456 123456 123456

Fig. 26. A. Tooth-wear categories in Aethomys namaquensis. B. Numbers of individuals in
each category in selected samples from the Cango Valley.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 299

An interesting fact is that the coefficients of variation vary inversely with
mean depth of mandible (Fig. 25). Thus, when the depth of mandible is great
the coefficient of variation is low, and vice versa. This suggests that there is
some connection between the two. One possibility was that there were changes
in the age composition of the populations at different times as was suggested
for Crocidura flavescens. It may be, for instance, that in Boomplaas A level
BLA the population sample consisted of mostly mature or older individuals
with few young individuals, which would give a relatively high mean depth of
mandible and a relatively low coefficient of variation. In level LP, on the other
hand, the age groups may be much more evenly represented but with the
emphasis on younger individuals. A pilot study was undertaken in which the
samples were divided into six age categories based on the degree of tooth wear
(Fig. 26) such as was done by Davis (1959) and Dean (1973) for Praomys
natalensis. This showed that there were apparently no clear-cut differences in
the composition of the samples (Fig. 26). The general proportions were simi-
lar in all the samples examined and the extent of the observed range in indi-
vidual size variation was also similar for all levels (Table 37). This latter fact
would suggest that neither the young nor the old were missing from the
samples. It may be that with more data a pattern would emerge, but present
evidence does not suggest that there exist differences in proportions of age
categories such as could explain the observed pattern. It will, however, prob-
ably be worth examining this problem in greater detail in order to establish
what mechanisms may be operating as well as what environmental information
is forthcoming.

For Tatera afra the evidence from Die Kelders 1 is not internally consis-
tent. That is to say, the data from the upper teeth do not exhibit the same
trends as do the lower teeth (Fig. 27). It is difficult to know how this may be
interpreted but is seems unlikely that it can have any environmental signi-
ficance. It also means that it is not possible, on present evidence, to decide
which set of data may be more reliable for palaeoenvironmental interpretation.
The fact that the general relationship between samples from Die Kelders 1,
Byneskranskop 1 and Nelson Bay Cave is similar to that found in Crocidura
flavescens, would suggest that there may be some potential information to be
gained from T. afra. It looks as if T. afra is conforming to Bergmann’s Rule,
but more data are needed before this can be confirmed or refuted.

Cryptomys hottentotus

Mandibles of Cryptomys hottentotus from Boomplaas A and B-C were
measured in five parameters as was described above. The resulting data are
given in Tables 39 and 40 and these results are shown graphically in Figures
28-30. Figure 25 illustrates variation in the mean for the various parameters.
The mean coefficients of variation indicate that alveolar length and distance
between posterior border of M, and posterior border of symphysis are
apparently little affected by age or sex (Table 33). There is much more

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300 ANNALS OF THE SOUTH AFRICAN MUSEUM MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 301
TABLE 39
Size variation in mandibles of Cryptomys hottentotus from Boomplaas.
A: alveolar length B: posterior border M;-symphysis » C: depth attachment ascending ramus D: foramen mentale-alveolar margin E: depth of mandible
N Range x s 95% N Range x s 95%\N Range X s 95% N Range X s 95% N Range X s 95%
BPA: DGL .. . . 60 4,5-6,1 5,25 0,31 0,08 60 7,3-9,1 811 0,41 O16) 3,448 4,16 0,35 0,10 60 1,5-2,5 1,93 0,23 0,06 21 4,0-6,0 4,89 0,70 0,32
BLD3 .. . . 30 4,9-5,8 5,24 0,25 0,09 30 = 7,3-9,3 8,00 0,44 01630 3,64,9 4,16 0,33 0,12 30 1,5-2,9 1,97 0,30 0,11 10 3,6-5,8 4,89 0,70 0,55
BL. .. - . 30 457-5,6 5,19) 0;22 (0,08 30 7,5-8,8 7,98 0,31 031230 3,4-5,5 4,13 0,44 0,16 30 1,5-2,4 1,92 0,25 0,09 7 4,4-6,1 5,07 0,66 0,61
BLA... . 30 4,7-5,8 5,37 0,29 0,11 30 7,5-8,7 8,08 0,36 0,123 3,5-4,9 4,11 0,34 0,13 30 1,5-2,3 1,97 0,22 0,08 1 (5,5)
BRL «. . ... 121 4,9-6,1 5,43 0,26 0,05 121 ~=—-7,3-9,1 8,25 0,44 0,08)2i 3,2-5,3 4,17 0,36 0,06 121 1,6-2,7 2,08 0,23 0,04 20 4,2-6,4 5,04 0,50 0,23
CL . . . . 90 48-62 5,43 0,26 0,05 90 7,3-9,3 8,18 0,40 010890 3,5-5,2 4,24 0,35 0,07 90 1,7-2,7 2,11 0,21 0,05 31 42-67 5,15 0,69 0,25
GWA .. . . 30 5;0-5;8 5,49 0,27 0,10 30 3=7,5-8,7 8,18 0,35 01230 3,5-5,2 4,30 0,42 0,16 30 1,7-2,6 2,09 0,22 0,08 3 4,6-6,8 5,93
LP. ee oe (ee 30) “489=6:2 5,55 0,32 0,12 30. = 7,3-8,8 8,21 0,36 =0;12:30 3,9-5,1 4,41 0,30 011 30 1,7-2,6 2,13 0,24 0,09 3 5,4-6,1 5,67
YOL . =. . = 10) ~5/3=6:0 5,74 0,19 0,14 10 7,9-8,9 8,31 0,36 0/2610 40-47 4,34 0,23 0,16 10 1,7-2,3 2,05 0,21 0,15 1 6,0)
BP4 eee 230)- 531=6;3 5,67 0,29 O11 30. = 7,9-9,3 8,44 0,37 031430 3,9-5,2 4,50 0,31 0,11 30 1,7-2,8 2,23 0,25 0,09 5 5,2-6,2 5,60 0,39 0,49
OLP ... . 14 5,464 5,87 0,32 0,19 14 80-94 8,58 0,40 0;22!4 3,7-4,9 4,51 0,34 0,20 14 1,8-2,7 2,24 0,24 0,14
OLP1 ... . 30 5,2-6,2 5.78 0,29 0,11 30 = 7,5-9,7 8,69 0,53 03203) 3,8-5,8 4,52 0,43 0,16 30 1;6-2,8 2,19 0,30 0,11 _11 4,8-6,3 5,50 0,52 0,35
OLP2 .. . . 30 5,3-6,4 5,82 0,26 0,10 30 = 7,8-9,2 8,60 0,40 03143) 3,9-5,5 4,64 0,40 0,15 30 1,8-3,1 2,36 0,33 0,12 30 4,6-7,5 5,71 0,67 0,25
@BP3) 5. . = 4 30) 332-61 5,80 0,24 0,09 30 © ©8,3-9,4 8,83 0,28 01103) 4,0-5,5 4,68 0,31 0,11 30 1,6-3,1 2,32 0,32 0,12 20 4,5-7,2 5,73 0,64 0,30
BOL, =. .: ws . 30) 5,3-6,;3 5,67 0,28 0,10 30. ~3=—-'7,8-9,4 8,56 0,40 03193) 3,5-5,0 4,41 0,31 0,12 30 1,7-2,8 2,17 0,25 0,09 14 4,9-6,0 5,49 0,39 0,22
OGH:- =. « « « 30) 5;2=6:3 5,79 0,31 0,12 30 = 7,8-9,2 8,49 0,40 03183) 4,1-60 466 0,41 0,15 . 30 1,9-3,0 2,36 0,24 0,09 8 5,0-6,7 5,79 0,60 0,50
BPB-C . soe © 66 4 8=5;4 5521 0,18 0,10 16 ~=—7,8-8,8 8,29 0,33 031815 3,3-4,6 4,06 0,34 0,18 16 1,7-2,4 2,03 0,18 0,10 16 4,1-6,8 5,12 0,65 0,35
Measurements in mm. 95% = 95% confidence interval for the mean. |
|
TABLE 40 |
Size variation in mandibles of Cryptomys hottentotus from Boomplaas A upper levels.
A: Alveolar length B: posterior border M;-symphysis ij depth attachment ascending ramus D: foramen mentale-alvyeolar margin E; depth of mandible
N Range x s 95% N Range Xx s 95%N Range X s 95% N Range X s 95% N Range X s 95%
Unit DGL :
DGL1 Sue. On! See 10 4,8-5,7 5,25 0,28 0,20 10 = 7,9-8,8 8,24 0,34 0/2410 3,445 4,04 0,39 0,28 10 1,6-2,5 1,93 0,29 0,21 4 40-5,6 4,90
DGL2 we ee on 20 -AS=5.9 5,22 0,30 0,14 20 ~=7,5-9,0 8,19 0,38 01820 3,64,8 4,23 0,34 0,16 20 1,6-2,3 1,93 0,17 0,08 10 4,0-6,0 4,79 0,79 0,56
BED2 a 6h ey ee 13. 4,8-6,1 5,36 0,39 0,23 13. = 7,3-9,1 8,03 0,50 0,3013 3,5-4,8 4,28 0,38 0,23 13. 1,5-2,4 1,97 0,25 0,15 5 4,3-6,0 5,10 0,72 0,89
BLD2A by" cee 17 4,9-5,7 5,21 0,26 860,13 17 —‘7,3-8,8 8,01 0,41 0,2117 3,6-4,6 4,04 0,29 0,15 17 1,5-2,4 1,88 0,25 0,13 2 4,6-5,1 4,85
Unit BRL Z
WN 5 6 5 2 39 9 IA See Sev Wp ays 12 7,3-8,5 8,03 0,40 0.2512 35-49 4,10 0,39 0,25 12 1,7-2,6 2,04 0,23 0,14 1 (6,4)
BRE ae ge eo 1G) 512651 5,55 0,29 0,14 19 7,5-9,0 8,35 0,39 00,1919 3,6-4,8 4,13 0,33 0,16 19 1,7-2,4 2,07 0,24 O11 4 4,2-5,3 4,85
BRI4..... . . 30 5060 5,45 0,26 0,10 30 7,5-9,0 8,32 0,42 0)1630 32-50 415 0,35 0,13 30 1,6-2,5 2,07 0,24 0,09 8 4,3-5,6 5,03 0,41 0,34
BRES . .-. = +. . 30 4(9-6,1 5,42 0,28 0,10 30 = 7,3-9,1 8,24 0,47 0)1830 3,6-5,3 4,22 0,35 0,13 30 1,6-2,5 2,09 0,21 0,08 5 4,3-5,2 4,88 0,41 0,51
BREG6G . .:. 5 «... 30) 4:9-5:8 5,39 0,24 0,09 30 ~=—7,4-9,0 8,23 0,46 01730 35-51 4,21 0,39 0,15 30 1,7-2,7 2,12 0,23 0,09 2) Syl—552 os LD
Unit CL
BRL7 .... . . 30 4860 5,49 0,27 0,10 30 7,5-9,3 8,25 0,45 01730 35-50 4,29 0,33 0,12 30 1,7-2,6 2,11 0,21 0,08 5 4,5-6,3 5,12 0,71 0,88
Oul 5 6 so o 6 5 8) SEE Go WG On 30 7,8-9,2 8,26 0,37 0;1430 3,6-4,9 4,23 0,30 0,11 30 1,9-2,5 2,13 0,19 0,07 8 4,3-5,5 4,94 0,50 0,42
CL3BG ... . . 30 49-56 5,30 0,19 0,07 30 7;3-8,6 8,03 0,33 01730 35-52 4,19 0,43 0,16 30 1,7-2,7 2,07 0,25 0,09 18 4,2-6,7 5,26 0,76 0,38

Measurements in mm. 95% =95% confidence interval for the mean.

ANNALS OF THE SOUTH AFRICAN MUSEUM

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304 ANNALS OF THE SOUTH AFRICAN MUSEUM

variation in depth of attachment of ascending ramus and, particularly, in the
two parameters related to depth of mandible.

There are very noticeable differences in the various parameters in different
levels at Boomplaas A. In all the parameters except depth of mandible (Figs.
28-30) there is a tendency for a general decline in size in progressively younger
samples. There appears to be some indication of a slight reverse in the trend in
the uppermost levels, although this is not entirely borne out by the more
detailed analysis of these levels, which will be discussed below. The depth of
the mandible shows quite a different pattern, with the mean for this parameter
being very much lower in the postglacial levels than in the glacial levels (Fig.
30). Such a pattern is very similar to that for Crocidura flavescens, which would
tend to suggest a correlation with temperature. It is almost certain, however,
that the pattern is the spurious result of incomplete data (Tables 39-40) and it
is quite possible that, given more complete data, the pattern would be rather
more like those provided by the other parameters. Under the circumstances,
the main pattern is to be preferred as being based on sounder evidence.

The pattern exhibited by C. hottentotus is different from those found in the
other species examined, except for the teeth of Aethomys namaquensis. As was
suggested for the latter, it is unlikely that this pattern can be correlated with
temperature change. If this is the case, it is possible that C. hottentotus 1s,
indeed, responding to changes in rainfall, or perhaps effective precipitation. C.
hottentotus may, aS was suggested above, be showing a reaction analogous to
that of Spalax ehrenbergi in Israel, which increases in mass with increasing
rainfall (Tchernov 1968: 39). If this is so the present data would suggest a
general decline in rainfall at Boomplaas during approximately the last 80 000
years. It is, however, also possible that C. hottentotus exhibits a reverse
reaction and that decreases in mass coincide with increases in rainfall. This
may, in fact, be suggested by evidence (D. M. Avery, unpublished data) that
this species is smaller in the south-western Cape than it is further north. On the
other hand, the fact that C. hottentotus is relatively small in the glacial
maximum samples from Boomplaas A (Figs. 28-30), would suggest that
decreases in mass and rainfall are coincident. This is because the glacial
maximum is thought on other evidence to have been dry. Detailed studies will,
of course, be needed to determine more precisely what correlation, if any,
there is between body mass and rainfall.

The patterns exhibited by the detailed analysis of the upper levels show
general consensus, except in depth of mandible which, again, is different from
the others (Fig. 25). The indication is that the mean size of individual in the
populations sampled in the lower half of the sequence, approximately levels
CL1 to BRL2, was smaller than the mean size in the upper half. The smallest
mean size apparently occurred in levels BL to BLD2A approximately. There is
some partly inconsistent evidence for an increase in mean size thereafter which
is most obvious in length of M; to symphysis. In this case the modern BPB—C
sample continues the trend but in the other parameters this is not the case.

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MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 307

Although similar to the means for the material from the latest levels at BPA,
the means from BPB-C tend not to conform to the trend (Fig. 25).

There is no conclusive evidence of rapid change in this species. Where ‘t’
tests show there is significant difference in the means of successive levels (Table
41), the radiometric dating tends to indicate that a relatively long period of
time is involved. Only in the case of levels OLP3 and BOL is the difference

TABLE 41
Selected ‘t’ tests for Cryptomys hottentotus.
‘ff d.f. P

A*

i Aw 271 58 0,01
SE oe ee a at we Dove 29 0,05
ee Rh 1,93 58 0,05

B

es Oe Sl et 20 29 0,05-0,02
ge 2,54 58 0,02-0,01
Ue 2,58 58 0,02-0,01
a ee 3,03 58 0,01-0,001
e

SE ek 3537 58 0,001
MOE eG 2,66 58 0,01

D

UM lk 2,04 38 0,05
LS ee 2,09 58 0,05
Ee Re 2,02 58 0,05
ee 3,00 58 0,01-0,001

* See Tables 39-40 for explanation of parameters and basic data.

significant in all four parameters tested. Depth of mandible was not tested
because it did not appear to show any real pattern of change. The null
hypothesis was two-sided in tests for this species because no assumptions could
be made about the direction of size change; that is to say, it was not possible to
hypothesize that the trend should be towards larger or smaller during a given
period of time.

EVIDENCE FROM SPECIES DIVERSITY

It has been observed that, in general, animal life in tropical climates tends
to be more abundant and varied than that in temperate climates (Krebs 1972:
500). This is, in effect, the same point as that made by Kowalski (1971: 466)
and mentioned above, that mammalian communities in harsh climates contain
fewer species than those in milder climates. If different climatic conditions can
affect the structure of contemporary communities, it should follow that changes
in community structure or species diversity in one place at different times in the
past would also be due to climatic differences. Considerable differences are, in
fact, noticeable in the structure of micromammalian communities of different

308

ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 42

Variation in aspects of species diversity in the Cango Valley and at Klasies River Mouth 1A,

Nelson Bay Cave, and Glentyre.

N
BPA: DGL. 848
BLD3 1055
BE: 280
BLA . 604.
BRL . 1761
a) ae 1449
GWA 1805
EPs. 1839
LPG 140
YOR. 315
BPI 353
BP? 110
BP3 93
BP4 . 1708
OLP . 355
OLP1 1189
OLP2 9500
OLP3 4197
BOL. «. 1802
BOLI-3 . 110
BOL4 106
BOLS 96
OCH 529
LOH. 120
BPB . 747
BPG@:. 504
NGA 532
NGB ..«: 261
KRMIA: 6 47
15 103
16 43
22 42
32 107
NBC: BSL . 59
Yo... 2882
YGL 770
GLEN 542
N = number of individuals
S = number of species
c = dominance
d = species richness
€ = evenness
H = general diversity

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 309

ages both at Boomplaas A and at coastal sites. Analysis of the differences
involved suggests that this line of investigation could prove useful for palaeo-
environmental interpretation. In particular the general diversity index (H)
appears to conform well to known general climatic trends. Thus, H is much
higher in postglacial than in glacial communities. If glacial is taken to mean
harsh conditions and postglacial, or interglacial, to mean mild conditions, this
suggests that the basic hypothesis given above is true and that climatic variation
is being reflected in differences in species diversity.

It is, however, important to point out here that the equation of mild
conditions with interglacials, and vice versa, refers specifically to the sites
discussed below and generally to any sites currently experiencing mild climates.
The modern samples from the Cango valley and Byneskranskop 2 exhibit high
indices and present conditions are known to be mild. Similar high indices in the
Holocene may be expected to indicate conditions similar to those of the
present, always assuming that the faunal list is not completely different. In
areas where it can be shown that modern conditions are reflected in lower
indices of diversity, interpretation would obviously have to be adapted accord-
ingly. The index may be artificially low, as in the case of Glentyre (Table 42),
the situation apparently being caused by the predator’s reliance on one major
prey item. In these circumstances it would be complicated to compare this
sample with another. Naturally low indices would be expected in a desert or
other harsh environment, so that changes in the past may conceivably include
increases in diversity; in other words, the pattern may be the opposite of that
observed in the southern Cape.

The index of species richness or variety (d) was calculated in order to
discover whether there was any correlation between this factor and general
known climatic conditions. If the situation noted by Kowalski (1971) were to
find any application in the present study, the index of species richness might be
expected to show this. In fact, at Boomplaas A the index is low for samples
from levels LP and GWA which are known to have been accumulated during
the last glacial maximum and high in the Holocene (Fig. 31). The indices for
the modern samples are also high. Likewise, the indices for the Die Kelders 1
M.S.A. samples are lower than those for the L.S.A. sample and the Bynes-
kranskop | samples. The BNK2 sample also has a high index. This would seem
to indicate that generally this index constitutes a reliable expression of climatic
conditions. It would appear, however, that minor fluctuations are more likely
to be due to differences in the size of the sample than to changes in conditions.
More specifically, large samples tend to have low indices of species richness,
but this is almost certainly a function of the size of the sample. There would
seem to be an optimum sample size which is governed by the maximum
possible number of species; once all possible species are represented, any
subsequent increase in the sample will merely have the effect of reducing the
index of species richness. For this reason it is suspected that the indices for
BPA levels OLP3 and OLP2 may have been artificially lowered. The situation

310 ANNALS OF THE SOUTH AFRICAN MUSEUM

H Boomplaas A H Byneskranskop | H Die Kelders |

abcdefghijkImnoparstuvwx

t SF Sas AAS

DGL d BLA g GWA j «“¥OL m BP3 Pp OER s BOL v BOLS
b BLD3 e BRL ft SEP k BPI n BP4 qr (OERZ t BOLI-3 w OCH
¢ ‘BE f Ce r ERE BR2 6, OEP to JOLRS u BOL4 x LOH
Pig, 31

Aspects of community structure in samples from Boomplaas A, Byneskranskop | and
Die Kelders 1. (H = general diversity; c = dominance; d = species richness;
e = evenness. )

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 311

at DK1 illustrates the process well. Here it would seem that the maximum
possible number of species is seventeen and that an optimum sample size was
approximately 1 000 since no new species were added in samples larger than
that. Therefore, in levels 7 up to 3 the index of species richness is determined
solely by the size of the sample. Again the index for samples from these levels
will be artificially low. It would seem, therefore, that this is not an ideal index
to use in the present context because of these complicating factors.

Evenness (equitability) of species representation is another aspect of
diversity which required assessment. The index (e) used to express this seems
to be virtually independent of sample size and, as such, is intrinsically of
greater use for subfossil samples. The same general pattern emerges as for the
species richness, that is, postglacial samples have a higher index than do glacial
samples (Fig. 31). There are some exceptions and it is of interest to note high
values for this index in samples from BPA levels BOL1 to BOLS and again
from levels BP1 and BP2. This may suggest more equable conditions during
these periods. By the same token indices for samples from the lowest M.S.A.
levels at DK1 fall within the range expected for interglacial conditions. Indices
for BNK1 samples are generally higher than those for postglacial BPA samples,
which could reflect the fact that maritime climates are normally less extreme
than continental climates, especially in mountain regions. It also emphasizes

~the fact that comparisons can be made between different areas provided it is
known that the samples are contemporary. There may thus be some potential
for investigating spatial variation as well as temporal variation.

Dominance is, in effect, the converse of evenness and this is shown very
clearly by the indices. Figure 31 illustrates that the index of dominance (c)
varies inversely to the index of evenness. Thus, at Boomplaas A the highest
index of dominance is recorded for samples from the last glacial maximum,
levels LPC to GWA. Very high indices also occur for the samples from Nelson
Bay Cave levels of a similar age. Holocene indices of dominance are very much
lower, with a tendency to be rather lower at Byneskranksop 1 than at Boom-
plaas A. Indices for Die Kelders 1 are relatively low in the lower half of the
M.S.A. sequence. Indeed, the index for many samples from these levels is
similar to or even lower than that for the sample from Die Kelders 1 L.S.A.
level 12. It would appear that either this index or the index of evenness can be
used with advantage for palaeoenvironmental interpretation, but that the
pattern exhibited by dominance is rather clearer.

The Shannon index of general diversity (H), which takes into consideration
both species richness and evenness, is probably the most suitable for present
purposes. It is relatively unaffected by sample size, as was mentioned above, it
takes the two main aspects of diversity into account and it produces a clearer
pattern than any of the other indices (Fig. 31). At Boomplaas A there is a
striking difference between the high indices for the postglacial samples and the
low indices for the glacial samples with no overlap between the two (Fig. 31,
Table 42). Even in the case of Byneskranskop 1 and the Die Kelders 1 M.S.A.

S12 ANNALS OF THE SOUTH AFRICAN MUSEUM
samples, where the distinction tended to be less clear in the individual indices,
there is little overlap. The dividing-line between what may be termed the glacial
and interglacial values of the index would appear to fall somewhere between
2,10 and 2,20. It is also noticeable that coastal and inland samples seem to be
directly comparable, with very similar values.

At Boomplaas A the lowest diversity occurs in samples from levels BOL4
and BOLS and again in levels LPC to GWA (Fig. 31). Since the latter are
known to have been deposited during the last glacial maximum, it would
appear that the earlier samples were accumulated during the previous glacial
maximum. The period between the two maxima seems to have been rather
more moderate, with peaks of improvement in levels BOLI and BP2. After
BP2 there is a general reduction in diversity until level GWA. Thereafter, a
major amelioration is indicated which is maintained, with minor fluctuations,
throughout the Holocene.

In the Die Kelders 1 M.S.A. samples, diversity shows a general reduction
in progressively younger samples (Fig. 31). This is particularly noticeable in the
five upper levels where the youngest sample has the lowest index. This suggests
that the onset of glacial maximum conditions in the area occurred at this time.
The lowest level, on the other hand, has a much higher index than the others,
well within the range for Byneskranskop 1 samples, which indicates a mild
climate when the site was first occupied. Indices for samples from levels 13 to 6
fluctuate but tend to be relatively high, suggestive of intermediate conditions.
The L.S.A. level has a low index for a Holocene sample. Whether this is due to
climatic conditions or to a biased sample is not known, but is perhaps likely to
be the latter. This is partly because the sample was derived from a shell midden
where conditions are unlikely to have been ideal for the preservation of small
mammal bones but mainly because the index for the youngest sample at
Byneskranskop 1 is high. Being part of a sequence it seems reliable and
suggests that the Die Kelders 1 index is too low.

The indices for samples from Byneskranskop 1 indicate that there were
two periods of harsher climate, one in level 18 and a second in levels 9A and
9B. The former may have been about 10 000 B.P. and the latter is about 6 200
B.P. Between these two there was apparently a peak in level 12 perhaps about
8 000 B.P. The upper levels indicate a general improvement in the climate after
about 6 200 B.P. until the top of the sequence about 1 850 B.P. Byneskranskop
2 has a much higher index than any of the subfossil samples which is difficult to
explain but may be due to the much higher number of species present
(Table 43).

Nelson Bay Cave samples have low indices as might be expected for glacial
samples. The indices for Klasies River Mouth 1A rise from the oldest to the
youngest but are still low throughout. It is, however, almost certain that the
results for this site are affected by the fact that not all the microfauna was
collected. If, as was apparently the case, only larger species were collected this
would tend to reduce species richness and could introduce artificially high

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS

Bent? la.

FO.
1(a+b)
ee 2.1:
3

4 .
2-4

9A.
9B.
10
1]
12
13
14
15
16
Ls
18
let

BNK2
DK1: LSA 12 .

TicnanZz

Hel AP aie

MSA 1.

numbers of individuals
numbers of species
dominance

species richness
evenness

general diversity

N

TABLE 43

Variation in aspects of species diversity at Byneskranskop and Die Kelders 1.

Ss

als

314 ANNALS OF THE SOUTH AFRICAN MUSEUM

dominance of certain species. It is possible that the general trend is accurately
reflected if the same biases are present in all samples, although it is interesting
that this trend is the opposite of what might be expected on other evidence. On
balance, because of the unknown extent of the bias, it was considered that
results for this site should probably be discounted until more reliable samples
can be obtained.

PALAEOENVIRONMENTAL RECONSTRUCTION BASED ON MICRO-
MAMMALIAN EVIDENCE

Reconstruction of past environments on the basis of the micromammalian
evidence takes into account all the data from the different lines of investigation.
Generally these are not at variance with each other, but where they are an
explanation must be sought which could account for the apparent anomaly. In
the present exercise it would appear logical to work from the particular to the
general. For this reason the data for the three main individual sites are
examined first. All the evidence for the coastal area is then considered as a
unit. Finally, the complete coastal sequence is compared with that from the
Cango valley in order to assess the degree of difference or similarity between
the sequences of the two areas, and to make some general interpretation for
the whole region if possible.

BOOMPLAAS A

Variation in all aspects of the small-mammal community examined indi-
cates extensive change in the environment of the site during the period of its
occupation. In general the various lines of evidence are in agreement. The basic
pattern, with vegetational interpretation, is provided by the factor analysis of
community composition. Variation in mean size of selected species has been
taken as indicating changes in climate which can also be referred to the basic
framework. Variation in species diversity or community structure has been used
to provide general confirmation of the relative harshness or mildness of
conditions, effectively climate, at various times during the period under discus-
sion.

The BPA sequence has been divided into five palaeoenvironmental units
(Table 44) on the basis of the factor analysis (Table 16). Of these the oldest,
unit 5, refers to the period represented by levels LOH up to BOLI. This unit
is, in fact, not homogeneous and should perhaps be considered in terms of
three sub-units. Samples from levels LOH and OCH probably represent the
early glacial period, as was mentioned above. The factor analysis suggests that
this was a period of intermediate conditions with a slight indication of deterio-
ration towards glacial conditions. The general indication from Crocidura
flavescens and from the general diversity index is also of moderate conditions,
that is, those approximately intermediate between full glacial and interglacial.
Myosorex varius suggests that conditions may even have been nearer those

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 315

TABLE 44

Sequence of levels at Boomplaas A with radiocarbon dates, cultural units and palaeo-
environmental interpretation.

i

Holocene nit 1
1 630+50 Herder scrub on hillsides, sparse fluctuating
?semi-arid in places,
1955+65 elsewhere ?dense microphyllous;
fairly open grass, ?trees on valley
floor, extensive dense riverside reeds
or grass

rapid
temperature
rise

Wilton
mildest
6 400+75 in BL
9 100+135

10 425+125

fluctuating
rapid

temperature
rise

Albany

Upper
Pleistocene
12 425+ 130

14 200+ 240

UNIT 2
transitional between Unit 1 and
Unit 3

Robberg becoming mild

warmer

UNIT 3
generally fairly open,

?semi-arid on valley

floor; reduced dense reeds or grass
along river; restioid or ‘grassy’ vege-
tation on hillsides; BP1 conditions
rather more moderate than general

coldest harsh

Undefined
LSA

21 100+420 industry

UNIT 4
fairly extensive dense vegetation on
valley floor; extensive restioid or
‘grassy’ element on hillsides plus
some ?semi-arid vegetation
(conditions harsher than general in
OLP)

32 400+ 420

moderatt
MSA

UNIT 5
general increase in scrub

on hillsides, grass on valley floor
and dense riverside vegetation:
BOLS glacial maximum as in Unit 3

period of change, ?becoming drier,

more open vegetation warm moderate

pertaining in the Holocene. This is not entirely inconsistent with the evidence
from Crocidura flavescens and it could perhaps be the case. There remains,
however, the possibility that Myosorex varius is responding to more than one
climatic factor.

The sample from level BOLS indicates full glacial conditions, on the
evidence of the factor analysis, with the samples from levels BOL4 and BOLI
suggesting progressive amelioration thereafter. At the time of level BOLS the
vegetation was probably fairly open but with a moderate amount of dense grass
or reeds along the river. Elsewhere grass and some semi-arid scrub are
indicated on the valley floor and a restioid or ‘grassy’ element apparently

316 ANNALS OF THE SOUTH AFRICAN MUSEUM

dominated the hillsides. (It should be noted that here and elsewhere the term
restioid or ‘grassy’ is employed to indicate a broad vegetation category; the
specific type of vegetation will clearly vary from one place to another but
cannot necessarily be identified from the small-mammal data.) An increase in
the Holocene element in level BOL4 and, particularly, in level BOL1 would
suggest rather more scrub on the hillsides, a reduction in semi-arid scrub and an
increase in denser vegetation generally. The diversity indices confirm the
suggestion that at the time of level BOLS the climate was harsh. They also
indicate that conditions were as harsh during the time that level BOL4 was
accumulated but that they were considerably milder during the time of level
BOLI. The general indication is that although levels BOLS and BOL4 were
deposited during a glacial maximum, this was rather less severe than the
subsequent one which is discussed below.

Palaeoenvironmental unit 4 refers to the period represented by samples
from levels BOL up to BP4. These levels appear to have been deposited during
an interstadial in the Last Glacial when conditions were approximately inter-
mediate between full glacial and interglacial. On the valley floor there would
appear to have been fairly extensive dense vegetation but on the hillsides the
open restioid or ‘grassy’ element would seem to have predominated with,
possibly, a smaller semi-arid scrub element on the lower hillslopes. It is possible
that the restioid or ‘grassy’ element is that which is today called Dense
Restioid-Proteoid Shrubland (Moffett & Deacon 1977) and confined to higher
altitudes, especially where conditions are relatively dry. At the time of level
OLP conditions may have deteriorated somewhat. The evidence from Croci-
dura flavescens indicates that the period of level BOL was relatively warm.
There is an apparent inconsistency in the temperature data. The C. flavescens
data suggest that the period covered by levels OLP and BP4 may have been
warmer than that represented by levels BOL to OLP1; Myosorex varius
suggests the opposite. The diversity indices suggest that the samples from all
these levels were accumulated under moderate conditions and it is probably
preferable to accept this generalization until it becomes possible to clarify the
situation.

Palaeoenvironmental unit 3 refers to conditions pertaining during the
period represented by levels BP3 up to GWA. Radiometric dating indicates
that this period lasted from approximately 30 000 B.P. to about 15 000 B.P. As
has been pointed out previously, this is known to include the last glacial
maximum and there is no difficulty in interpreting the micromammalian
evidence in the light of this fact. The factor analysis suggests that conditions at
the time of level BP1 were rather milder than general for this period. The
overall pattern indicates fairly open vegetation with a reduced amount of dense
grass or reeds along the riverside. Grass and probably semi-arid scrub occurred
elsewhere on the valley floor. Indeed, the relatively high proportions of Otomys
unisulcatus during this period suggest that semi-arid scrub was at its maximum
extent and that the climate must have been drier than at any other time.

q

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS hy

Moreover, A. Scholtz (1979 pers. comm.) has pollen evidence to suggest dry
hillside scrub during this period. The micromammalian evidence suggests that
the predominant vegetation type on the hillsides was restioid or ‘grassy’ and
open. The absence of Crocidura cyanea from the fauna during this period is
thought to be indicative of a cold climate, whether in the form of generally
depressed temperatures or of very cold winters. At the same time the presence
of semi-arid scrub in the valley would suggest dry conditions. Mean size of
Crocidura flavescens indicates that the climate was coldest during the time that
levels YOL to GWA were deposited, perhaps from 25 000 B.P. to 15 000 B.P.,
and the diversity indices confirm this suggestion. These indices further suggest
that conditions were mild when levels BP3 and BP2 were deposited and
moderate during the deposition of levels BP1 and YOL. The situation regard-
ing level BP1 has already been discussed and the factor analysis, particularly
the Oblique solution, may be interpreted as supporting the suggestion concern-
ing levels BP3 and BP2. For level YOL, however, the evidence is conflicting.
The factor analysis and C. flavescens indicate that, far from being moderate,
conditions may have been at their most extreme during this period. It is difficult
to explain this, although it is possible that the difference between moderate and
extreme may not be as great as might appear to be the case. It is also possible
that an increase in the C. flavescens sample may help to clarify the situation.
~ Following the last glacial maximum, a transitional period represented by
samples from the CL levels forms palaeoenvironmental unit 2. The period
involved is approximately 15 000 B.P. to 11 000 B.P. Unit CL appears to
represent a combination of Holocene and interstadial elements, according to
the factor analysis, while the individual levels show a clear trend from the
glacial to the Holocene in each succeeding level. A large increase in dense
vegetation, together with a decrease in the restioid or ‘grassy’ element on the
hillsides, is indicated. There was also an increase in sparse, possibly semi-arid,
scrub on the hillsides, but a disappearance of this vegetation type on the valley
floor. As may be expected a general increase in temperature is indicated. The
diversity indices show the climate to have been mild and much closer to that of
the Holocene than to that of the remainder of the Upper Pleistocene.
Palaeoenvironmental unit 1 refers to the Holocene postglacial period
represented by levels BRL up to DGL. The samples from these levels are
consistently shown by the factor analysis to indicate a quite different set of
conditions from those indicated by the samples from the Upper Pleistocene
glacial levels. In general there appears to have been scrub on the hillsides,
sparse and possibly semi-arid in places but elsewhere perhaps dense and
microphyllous. On the valley floor fairly open grass with possibly some trees
probably grew. Relatively extensive dense riverside grass or reeds are also
suggested and it is possible that the trees were, in fact, mostly confined to the
area along the edges of the river. While the main trend was apparently towards
higher temperatures there is evidence of two periods of fluctuation; one is in
unit BRL, between about 10 500 B.P. and 9 000 B.P., and the other is in unit

318 ANNALS OF THE SOUTH AFRICAN MUSEUM

DGL, about | 600 B.P. (Fig. 15). There is also an indication of two periods of
accelerated increase in temperature; one of these is between levels BRL6 and
BRLS5S, about 10 000 B.P., and the other is between levels BLD3 and BLD2A,
about 2 000 B.P. During the time of unit BRL, the climate apparently contin-
ued to be moderately dry, as is suggested by the relatively high proportion of
sparse possibly semi-arid scrub on the hillsides and even a relative increase in
semi-arid scrub on the valley floor.

It should be noted at this stage that there may be a restriction on
interpretation. This is related to the inequality of the time periods represented
by the different units in the archaeological sequence. At Boomplaas A, for
instance, some levels such as YOL may represent up to 10 000 years, whereas
others such as BP2 may represent only a few hundred years (H. J. Deacon 1979
pers. comm.). This could have a bearing on the amplitude of the trends being
reflected in the micromammalian evidence. The results of the factor analysis
suggest, in fact, that there exists an overall pattern of long-term trends. Within
these trends the evidence from individual species may indicate lesser
fluctuations where shorter time periods can be recognized. It would seem,
therefore, that where there is a long sequence it should generally be possible to
identify major trends, but only where that sequence is divided into short-term
units will it be possible to identify lesser fluctuations.

BYNESKRANSKOP 1

The series of micromammalian samples from Byneskranskop 1 provides
evidence for the last 13 000 years (Table 45). This includes the late glacial
period at the end of the Upper Pleistocene and the postglacial period or
Present Interglacial in the Holocene. The series has been divided into three
palaeoenvironmental units on the basis of the factor analysis. Climatic data for
individual species are sparse but species diversity provides some indication of
changing conditions.

Palaeoenvironmental unit 3 encompasses the earlier half of the period,
from about 13 000 B.P. to 6 500 B.P., and involves the samples from levels 19
up to 11 inclusive. The indication is that extensive, possibly fairly closed, grass
occupied the flats. Pans, which may suggest a relatively or seasonally wet
climate, are also indicated on the flats, as is a certain amount of scrub.
Considerable dense vegetation apparently existed along the river banks and on
the lower hillsides; it is possible, in fact, that the entire area between the
Uilenkraal River and Byneskranskop was so covered. On the hills themselves
there was a prominent restioid or ‘grassy’ element, especially at the time when
level 13 was deposited. There may also have been a proteoid element on some
of the hills as there is today (Schweitzer & Wilson 1978). The diversity indices
suggest that conditions were relatively harsh at the time that level 18 was
deposited and less mild than usual at the time of level 13. This latter agrees
with the suggestion, made above, that the restioid or ‘grassy’ element was most
prominent at this time, using the analogy indicated for Boomplaas A. The

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 319

TABLE 45

Sequence of levels at Byneskranskop 1 with radiocarbon dates, cultural units and palaeo-
environmental interpretation.

Dates B.P. General

1 880+50 ee
intermediate i
3 220+45 as unit 3 but more scrub and yee
3 400+55 less restioid vegetation on hills mild
UNIT 2
3 900+55 extensive ?more open grass on flats; warmer
rather less restioid or “grassy’ vegetation
on hills and more scrub; less dense
. vegetation near river and lower slopes
Wilton (?relatively dry) moderate
D
6 370+90
6 100+ 140
6 540+55
changing
vegetation
UNIT 3
; ; ; mild
extensive dense vegetation near river
and lower hillslopes; extensive ?more 12 mildest
closed grass on flats with pans and ene
some low scrub; restioid or ‘grassy’ 13 rather
?and proteoid element on hillsides general harsher
9 760+85 ; (peak restioid element in level 13) increase in
Pre-Wilton (?relatively wet) temperature
2 moderate
18 harsh
12 730+185 cooler 19 moderate

period represented by levels 12 and 11, perhaps from 8 000 B.P. to 6 500 B.P.,
apparently enjoyed a milder than general climate, especially during the earlier
part. In this context it should be pointed out that in this section references to
mild or harsh conditions are to be understood as being within the general
Holocene range; the amplitude is not comparable to that found at Boomplaas
A (see Fig. 31 and Tables 42-43).

In palaeoenvironmental unit 2, the period from about 6 500 B.P. to
approximately 3500 B.P. is covered by samples from levels 10 up) to. 3
inclusive. The factor analysis shows that the time when levels 11 and 10 were
being deposited was a period when the vegetation was changing from that of
unit 3 to that of unit 2. In unit 2 there appears still to have been extensive grass
on the flats but it is possible that it had become more open and that ground
conditions were drier. The apparent reduction in the number of pans would
suggest the same. There seems to have been a reduction in the amount of dense
vegetation on the river banks and on the lower hillsides. Grass would perhaps
have migrated into the area between the river and Byneskranskop. On the
hillsides there is an indication that to a certain extent scrub increased at the
expense of the restioid or ‘grassy’ element. Diversity indices suggest that, after
an initially mild period from about 6 500 B.P. to 6 100 B.P., there was a short
period perhaps about 6 000 B.P. when conditions were relatively harsh. There-

320 ANNALS OF THE SOUTH AFRICAN MUSEUM

after, in the last part, perhaps from 3 900 B.P. to 3 500 B.P., after a period of
moderate conditions, they again became mild. The evidence from Crocidura
flavescens indicates that at this time temperatures were considerably higher
than they were at about 12 700 B.P. This suggests a general warming trend, as
might have been expected, but it would have been interesting to know whether
there were fluctuations in the period between the two extremes.

Palaeoenvironmental unit 1 covers the latest period, from about 3 500 B.P.
to 1800 B.P., represented by samples from the top four levels. The factor
analysis indicates that the sample from combined level 2—4 is very similar to those
comprising unit 3. The level 2-4 sample suggests, however, that while the
vegetation was generally similar, there was probably more scrub and less restioid
or ‘grassy’ vegetation on the hills in the later period (about 3 500 B.P. to 3 200
B.P.). The diversity index suggests that conditions were milder than during the
earlier period, which agrees with the vegetational evidence. The evidence from
the sample from level 1 suggests intermediate conditions, apparently indicative
of a return to more open vegetation. The evidence could, of course, be
influenced by the fact that a long period of time is involved. It is of interest to
note, however, that the general diversity index is the same for the upper and
lower parts of level 1 (Table 43). This suggests that conditions cannot have been
greatly different at 1 800 B.P. from those pertaining at 3 200 B.P. although there
could still have been changes at some time between these two dates. In general,
however, the indication is that conditions during this later period were milder
than at any time since those indicated by the sample from level 12.

DIE KELDERS 1 MIDDLE STONE AGE SAMPLES

The series of samples from the M.S.A. levels at Die Kelders | provides
evidence of conditions during a period of possibly some 40 000 years during the
Upper Pleistocene. The sequence has been divided into three palaeoenviron-
mental units on the basis of the results of the factor analysis (Table 19). There
are few climatic data available from individual species but the evidence from
species diversity or community structure provides a general indication of
prevailing conditions. Because Die Kelders 1 is situated on the present coast,
there is also the possibility that the data provide some indication of sea-level
changes, although this is tenuous.

The samples from the lowest three levels, 14 up to 12, comprise palaeoen-
vironmental unit 3 (Table 46). The evidence suggests that during this period
there was extensive grassland on the flat ground which may also have existed
below the cave as well as above it. Extensive dense vegetation may perhaps
have been situated near a marsh or lake which is currently submerged. Both
these lines of evidence may suggest that the sea-level was relatively low during
this period. On the hillsides the dominant vegetation was apparently of an open
restioid or ‘grassy’ type. The general indication is that conditions were rela-
tively cold and rather wet. In level 12 there is some slight evidence for changing
conditions.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS a2

Palaeoenvironmental unit 2 comprises the samples from levels 11 up to 3,
although the sample from the latter level shows that conditions were intermedi-
ate at that time (Table 46). In general there is evidence for reduced grass and
dense vegetation on the flats. Because there is no evidence for a compensatory
increase in another type of vegetation, it is suggested that the sea-level was
higher during this period and that there was actually less level ground available.
It may also be that an expansion of neighbouring sand-dunes reduced the
available habitat. The dominant hillside vegetation remained restioid or ‘grassy’
but the proportion was reduced, again without compensatory increase in
another type of vegetation. The tendency for small species to predominate
suggests a generally fairly open vegetation. There is also some indication that
the climate was rather warmer and drier than it was during the previous period.

The samples from levels 1 and 2 comprise palaeoenvironmental unit 1
(Table 46). The indication is that conditions during this period were very
similar to those pertaining during the earliest period. Thus there was again an
increase in grass and dense vegetation on the flats with the possibility of a
renewed lowering of the sea-level. On the hillsides, however, scrub appears to

TABLE 46

Sequence of Middle Stone Age levels at Die Kelders 1 with palaeoenvironmental interpre-
x tation and approximate dates (after Tankard & Schweitzer 1974).

Approx.
dates B.P. Level Vegetation Climate General Sea-level

UNIT 1

35 000 i extensive grass, ? lower
dense vegetation on
2 flats; scrub on hills
UNIT 2
3 3 intermediate ? higher
4
5, reduced grass, ? general
dense vegetation on increase in
6 flats; moderate temperature
restioid or ‘grassy’
vegetation on hills generally
7 mild conditions
8 perhaps
deteriorating
9 gradually
10
1]
UNIT 3
12 12 slight change; ? lower
extensive grass,
13 dense vegetation on
flats; restioid or
80 000 14 ‘grassy’ vegetation

on hills

322 ANNALS OF THE SOUTH AFRICAN MUSEUM

have replaced the restioid or ‘grassy’ element and the general indication is that
the climate may have been rather warmer and wetter than it was during the
preceding period.

As was mentioned above, data from individual species are few so that any
interpretation is necessarily uncertain. However, the evidence from Myosorex
varius (Table 36) suggests that there might have been an increase in tempera-
ture at the time of level 8 compared with that pertaining when levels 4 and 11
were deposited. The evidence from Crocidura flavescens (Tables 30-32) indi-
cates a general increase in temperature from the time that level 10 was
deposited to that of level 6 and then level 4. These data may be less reliable
because they are based on fewer observations. Both sets of data suggest
temperatures might have been higher even than they were during the Holocene
and thus confirm the suggestion that conditions were mild during this period.
As has been pointed out above, the data for Tatera afra are not yet adequate
for palaeoenvironmental interpretation. Information for this species has not,
therefore, been considered here.

The information available from species diversity does not appear to agree
entirely with the evidence from other lines of investigation. There is a general
decline in the index of general diversity from the beginning of the period to the
end (Fig. 31, Table 43). This would suggest a trend towards harsher conditions
which are not otherwise apparently indicated. In fact, this is relative and for the
earlier samples, from levels 14 up to 5, the index is within the range exhibited
by Holocene samples from Byneskranskop 1. This would then confirm that
conditions were comparatively mild during that period. The low index for the
later samples is largely due to an increase in dominance rather than to other
factors (Fig. 31, Table 43). There are two reasons for suggesting that this
increase in dominance may be due to factors other than climatic conditions. In
the first place, the two dominant species are not the same in all five samples.
This would suggest that the vegetation—and presumably therefore the cli-
mate—must, in fact, have changed during this period. Secondly, Otomys
irroratus, which is one of the dominants in the samples from the upper levels, is
normally associated with mild conditions. On the other hand, it could perhaps
be that extremes in available moisture are being monitored. This might indicate
dry conditions during the time that levels 5 up to 3 were deposited and wet
conditions during the time when levels 2 and 1 were deposited. It is, however,
perhaps unlikely that conditions could have been sufficiently extreme to affect
the diversity in this way, especially since the same three species are most
numerous throughout the sequence, but in differing proportions.

COASTAL SAMPLES

Conditions pertaining at Die Kelders 1 during the time when the M.S.A.
levels were being deposited are shown to have been different from those at
Byneskranskop 1. This could be expected in view of the fact that the latter is a
Holocene site and the former is thought to be an Upper Pleistocene site. It is

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS ay

necessary, however, to attempt to establish how much of the difference is due
to climate and vegetation and how much to topography. It is clear, for instance,
that the species that occur in higher quantities at BNK1 are plains and lower
hillside animals. The inference is that these niches were reduced in the DK1
area. In fact, it has been suggested above that the area of flat ground below the
cave at DK1 fluctuated but was perhaps unlikely to have been extensive at any
time when the site was occupied. It is also possible that the nature of the cliffs
is not such as to provide a suitable habitat for some of the species which
occurred at BNK1.

Variation within the site may, however, provide some insight into the
relative situation. At DK1 in the majority of the levels Myosorex varius is
present in much higher proportions than Otomys irroratus, which suggests a
relatively dry climate. At BNK1 the situation tends to be similar in the upper
half of the sequence but not in the lower half. In general, therefore, it would
appear that the Holocene was rather wetter than the Upper Pleistocene on the
coast. This accords with the interpretation of the sequence at Boomplaas A. At
both sites it would appear that the restioid or ‘grassy’ vegetation generally
predominated on the hillsides. The only exceptions to this were in the latest
part of the DK1 M.S.A. sequence and in the DK1 L.S.A. level 12 and BNK2
samples. The evidence from Crocidura flavescens and Myosorex varius is not
really adequate for sound interpretation. It appears to be conflicting, at least in
part, but this may be due to the fact that complete sequences are not available.
As was mentioned above, the general diversity for the lower half of the DK1
M.S.A. sequence falls within the range for the BNK1 sequence. This would
suggest that conditions were not dissimilar during these periods at the two sites.

The modern (BNK2) and late Holocene (DK1 L.S.A. level 12) samples
suggest that conditions were different in this period from either of the two
earlier periods. The DK1 L.S.A. sample indicates that possibly conditions were
similar to those pertaining during the central part of the M.S.A. period but that
there was replacement of grass by scrub on the hillsides. Steatomys krebsi
replaced Tatera afra as the dominant plains animal as it did in apparently drier
times at Byneskranskop 1. Dense waterside vegetation was apparently an
important vegetational feature as it had been during the M.S.A. It was,
however, presumably located on the present coastal plain since the sea-level
would have risen to its present level before about 2 000 B.P. when the site was
reoccupied. Perhaps rather wetter conditions are suggested by the BNK2
sample, with relatively extensive dense waterside vegetation. The replacement
of grass by scrub on the hillsides is confirmed.

HOLOCENE AND MODERN SAMPLES

The factor analysis shows that there is a basic difference between the
Boomplaas A, Byneskranskop 1 and modern samples. In the case of the two
Holocene samples this is possibly due to differences in topography, as was
suggested in the previous analysis. Certainly variation in the proportions of the

324 ANNALS OF THE SOUTH AFRICAN MUSEUM

individual species shows a certain amount of agreement between the two sites,
even though the actual proportions are different (Fig. 14). There is, for
instance, a trend in both areas towards a reduction in the restioid or ‘grassy’
element on the hillsides and a slight reduction in the amount of dense waterside
vegetation. There is also some indication of an increase in open grassland on
the flat ground. In addition, from about 8 000 B.P. there appears to have been
an increase in dense vegetation on the lower hillsides in both areas.

It is interesting to note that in many cases the proportions of different
species vary considerably in the modern samples from the Holocene samples.
In general it is the species from the flat ground which are affected and it would
thus appear that the differences are most probably due to the effects of
agriculture. It has already been suggested that Mus minutoides is at an advan-
tage in cultivated land, and it is here noticeable that relatively high proportions
of this species distinguish all the modern samples. Reductions in the propor-
tions of Mystromys albicaudatus, Steatomys krebsi and Crocidura flavescens
could also be due to the destruction of their natural habitat by ploughing.
Equally, the relatively high proportions of Myosorex varius and low propor-
tions of Otomys irroratus could be due to the clearing of the valley floor at
Boomplaas. At Byneskranksop, on the other hand, the habitat of these two
species was not apparently altered. Also at Byneskranskop, the apparent
replacement of the restioid or ‘grassy’ vegetation by scrub within the last 2 000
years is presumed to have been natural and, indeed, appears to have been a
continuation of a trend. At Boomplaas, however, the situation seems to have
remained stable during this period.

In this respect it is of interest to note that the evidence from Crocidura
flavescens suggests that during approximately the last 4 000 years there has
been a similar rise in temperature in both areas. The mean percentage differ-
ence between modern populations and those about 4000 B.P. is 3,64 for
Boomplaas and 2,57 for Byneskranskop (Table 35). It is possible, however,
that the changes took place earlier at Boomplaas than at Byneskranskop,
although there are insufficient data from BNK1 for this period to prove the
point. It has, however, to be borne in mind that the mean difference of about
4,5 per cent between Byneskranksop and Boomplaas samples suggests a differ-
ence in temperature which could affect the time at which vegetational change
took place in the two areas. This would, in fact, tend to suggest that changes at
Byneskranskop were likely to have preceded those at Boomplaas because the
temperature was higher in the former area. More data are clearly needed
before the problem can be solved. This is especially the case since the diversity
indices for Byneskranskop and Boomplaas are virtually identical, and it is clear
that very similar conditions are indicated.

UPPER PLEISTOCENE AND MODERN SAMPLES

The factor analysis indicates that conditions during the central part of the
M.S.A. occupation at Die Kelders 1 were very similar to those obtaining at

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS S23

Boomplaas A during the period which has been interpreted as experiencing
interstadial conditions. It is of some interest to note that the modern Cango valley
samples also appear to represent similar conditions. The common denominator is
a high proportion of Myosorex varius. Since it has already been suggested above
that in modern samples this may be due to the effects of agriculture, it is thought
that the correlation in the present case is probably largely spurious.

Conditions at Nelson Bay Cave at the time when levels YSL and YGL
were deposited were quite different. This is to be expected because it is known
that the last glacial maximum occurred at that time. This provides confirmation
of the suggestion that there was a considerable difference between glacial
maximum and interstadial conditions. At NBC there is evidence for open
grassland on the flats, much as there is at the time of the lower M.S.A. levels at
DK1, which may suggest relatively cold conditions at the latter site. It may also
provide evidence for the known lower sea-level at NBC, as was suggested
above for DK1. At NBC there is an indication of extensive restioid or ‘grassy’
vegetation which may perhaps have been situated on the land above the cave
on what is now the coastal plain.

The evidence from Crocidura flavescens (Figs. 16-21, Tables 30-32) sug-
gests that temperatures were very similar at NBC and at BPA during the last
glacial maximum. This is of some interest in view of the fact that the later
evidence from BNK1 suggests that there is a difference between the coast and
the Cango valley. Also between DK1 and BPA, during what is thought to be an
interstadial in the Last Glacial, there is evidence that temperatures were higher
at DK1, which was presumably not far from the sea at the time. It may,
therefore, be that at about 18 000 B.P., when the sea was about 80 km east of
the site, NBC experienced a continental climate which was similar to that
affecting the Cango valley. Subsequent raising of the sea-level would mean that
a maritime climatic régime would have been reintroduced to the area of the
present coast near Nelson Bay Cave. The fact that this would have the effect of
moderating the climate could explain why there was apparently a differential
rate of increase in temperature on the present coast and in the Cango valley.
The mean percentage difference between the BPA sample and the BNK1
sample for the period about 13 000 B.P. is approximately double (9,57 per
cent) that between the two samples for the period about 4 000 B.P. (4,41 per
cent) (Table 35). This would suggest a relatively accelerated rate of tempera-
ture increase in the Cango valley during the time between the two dates,
possibly in compensation for postulated earlier increases along the coast.

The general diversity indices for Nelson Bay Cave and the glacial maxi-
mum levels at Boomplaas A are very similar, although somewhat lower for the
former. This again suggests that conditions were much the same in both areas.
The diversity indices for Die Kelders 1 M.S.A. samples are also similar to those
for samples thought to be approximately contemporary at Boomplaas A. The
slightly higher results for DK1 would tend to confirm the suggestion that
conditions were rather milder near this site than in the Cango valley.

326 ANNALS OF THE SOUTH AFRICAN MUSEUM

EXISTING EVIDENCE FOR LATE QUATERNARY ENVIRONMENTS

In the present study the word ‘environment’ is used in the restricted sense
of vegetation and climate. The latter, in turn, is represented by temperature
and rainfall or, possibly, effective precipitation. This is because micro-
mammalian evidence can provide information concerning the considerable
changes which are known to have occurred in these particular aspects during
the Upper Pleistocene and Holocene. Through assessment of the degree of
difference from the present, taken as the norm, of these variables it is possible
to establish a broad picture of past environments. However, what effect one
parameter will have had on others is extremely difficult, if not impossible, to
judge. This reflects both the complexity of the interrelationships of various
aspects of the environment and the present state of knowledge on the subject.
The result is that extrapolation from one aspect to another is a doubtful
process, although necessary at times. However, by examining all the available
evidence from different lines of research, interpretation may be based on as
secure a foundation as possible. In this exercise it is preferable to move not
only from the general to the particular but also from the complete to the
incomplete. The data from deep-sea cores satisfy both requirements so that, as
is argued by Kukla (1977), it would seem advisable to use these data as a base
against which to measure terrestrial data. These latter have a tendency to be
more localized in relevance but, at least in the more continuous sequences,
general trends can be recognized and correlated with those in the deep-sea
data. The geomorphological data tend to be both localized and incomplete and,
as such, must be interpreted with particular care. They do, however, have
undoubted importance for regional studies, especially when they can be fitted
into the general framework. For the region currently under investigation,
analysis of samples of terrestrial macrofauna from archaeological sites has also
provided insight into past vegetational changes.

The basic framework of climatic change during approximately the last
125 000 years is well established. Different lines of evidence show that after
being climatically similar to today, the remainder of the period was colder.
After some oscillations the temperature dropped considerably about 75 000
B.P. A subsequent partial recovery was followed about 20 000 B.P. by the last
maximum of the Last Glacial which is thought to have been the most severe.
During this time temperatures in South Africa might have been up to 10 °C
lower than at present on the interior plateau and high mountains, but were
probably in the region of 5 °C lower in the coastal regions. There is also a
possibility that there was less seasonal variability than at present. This climate
appears to have been favourable to grass, which seems to have comprised a
large part of the vegetation, at least on the coastal foreland and in intermon-
tane valleys, during glacial periods.

From about 12 000 B.P. conditions began to improve rapidly. By about
8 000 B.P. a state approaching equilibrium seems to have been reached.
Although it is clear that changes and fluctuations have continued to take place

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS ey

they are obviously on a much smaller scale than those previously experienced.
Both the oxygen-isotopic and the sea-level data indicate a reduction in the rate
of change.

MARINE EVIDENCE

The evidence for palaeoclimatic change during the Upper Pleistocene and
the Holocene from marine data has the advantage of being both essentially
complete and basically comparable in all parts of the world. Evidence for
changes in sea-level has long been studied, but the relatively recent advances in
the interpretation of deep-sea cores have had a profound effect upon palaeo-
climatic studies. These cores have the advantage of providing biotic and abiotic
material for study which, amongst other things, allows cross-checking of results
both from the cores themselves and from the sea-level data. In both cases
oxygen-isotope analysis, which measures the amount of '%O in calcareous
foraminifera and thus the surrounding sea-water, is of great importance.
Interpretation is based on the fact that during glacial periods not only is a larger
proportion of the earth’s water stored as ice at the poles, but also that ice is
relatively low in '8O. In compensation the oceans will be relatively high in 1°O
at the same time and sea-levels will be lower. In this way the oxygen-isotope
content of the water can be used to check whether contemporary low sea-levels
are due to changes in ice volume or to isostatic uprising of the land mass.

Emiliani (1955, 1966) originally conducted oxygen-isotope analysis in order
to establish a palaeotemperature sequence since he believed that changes in
'8O content were temperature dependent. He defined a climatic sequence in
which, basically, the even-numbered stages represent glacial periods and the
odd-numbered stages interglacial periods. Shackleton (1975) emphasizes, how-
ever, that these stages are related to the amount of ice present and not to the
climates that caused the ice. Moreover, whilst the marine record is divided into
roughly equal proportions of glacial and non-glacial time, there is evidence,
particularly at high latitudes, that full interglacial conditions were of much
shorter duration. Emiliani (1972) makes the point that periods when temper-
atures were as high as those of the present are exceptional and of short
duration. Shackleton (1975) concludes that the oxygen-isotope stratigraphy
provides a good framework but does not give sufficient information to allow
specific interpretation on a regional basis. In this general framework stage 5 is
correlated with the Last Interglacial, stage 3 with an interstadial in the Last
Glacial and stage 1 with the Present Interglacial. The intervening stages 4 and 2
represent the first and last maxima of the Last Glacial. Subsequent work by
Shackleton & Matthews (1977) has established the most reasonable time scale
for stages 1 to 7 and a correlation with three high-sea-level terraces in
Barbados. They conclude that Barbados II, the oldest and highest terrace, is
dated to about 125 000 B.P. and is to be correlated with standard isotopic
substage Se, representing the Last Interglacial sensu stricto. Barbados II
(105 000 B.P.) and I (82 000 B.P.), which also fall into stage 5, are thought to

328 ANNALS OF THE SOUTH AFRICAN MUSEUM

owe their present elevation to uplift of the land so that Barbados III represents
the last occasion when the continental ice-mass was as reduced as it is today.
On the basis of ages given by Shackleton & Opdyke (1976), the first glacial
maximum, stage 4, began about 75 000 B.P. and the last glacial maximum,
stage 2, at about 32 000 B.P.

As was mentioned above, not only was the ocean water high in ‘SO during
glacial periods, but also the quantity of water was reduced. If the effect of
tectonic and isostatic activity can be evaluated it should be possible to correlate
evidence for higher or lower sea-levels with past climatic conditions, although
differential rates of uplift can cause problems in interpretation. Problems also
arise from the fact that evidence for lower sea-levels is frequently submerged,
particularly for the Last Glacial when levels are generally thought to have been
more than 100 m below those of the present (Truswell 1977) during the last
maximum (Dingle & Rogers 1972). In southern Africa Butzer & Helgren
(1972: 160) note evidence of a shoreline at +5-12m which they believe
pertains to the Last Interglacial, while Davies (1971, 1972) records evidence of
such beaches all along the South African coast. The fact that the exact level
varies could be due to differing local conditions or to the beaches having been
formed during different stadia of the Last Interglacial. The evidence for low
sea-levels comprises submerged sediments, usually on continental shelves, and
drowned valleys and shorelines. Sediments laid down subaerially exist at
several tens of metres on many shelves, especially off the eastern United States
and western Europe, while drowned fluviatile patterns off the coast of France
have been discovered by the use of seismic plotting devices (Guilcher 1969: 83).
In South Africa similar work has been carried out on the Agulhas Bank, where
swathes of muddy sediment on the sea-floor are thought to indicate drowned
river-valleys (Dingle & Rogers 1972: 162). Whether these pertain to the Last
Glacial is perhaps more difficult to ascertain, although there would presumably
be some evidence if they represented more than one period of low sea-level.
Thereafter, during the Flandrian transgression, according to Shackleton &
Opdyke (1973: 46), sea-level rose rapidly from about 16 000 B.P. to reach its
present position some 6000 years ago, although Montaggioni (1976) cites
evidence from Réunion for a continuous rise in sea-level during the last 7 300
years to its present level. Butzer & Helgren (1972: 165) and Martin (1962: 25),
on the other hand, follow Fairbridge (1961, 1971) in suggesting that the
sea-level rose above its present level temporarily.

Faunal analyses of deep-sea cores have been based on changes in abun-
dance of individual key species and, more recently, on considerations of the
total fauna. As Shackleton (1975) has pointed out, the first method is too
simple and has proved unreliable, particularly in the older samples. Analysis of
all, or a large part, of the fauna represents a more sophisticated and reliable
approach. Of particular interest appears to be the work of Imbrie & Kipp
(1971) and Imbrie et al. (1973) which extracts quantitative estimates of surface
water conditions from data of planktonic microfossils. Analysis of core tops,

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 329

that is the modern data, forms the basis against which the fossil samples are
measured. Shackleton (1975) suggests that faunai analysis will prove useful for
within-stage changes but not for between-stage changes. Imbrie et al. (1973)
make the point, however, that oxygen-isotope analysis and their quantitative
faunal analysis are complementary because they measure different local respon-
ses to global climatic change.

Luz (1973, 1977) used a modified version of the Imbrie et al. (1973) system
to analyse the evidence from planktonic foraminifers in the Pacific Ocean. In
the process he noticed (Luz 1977) a distinction between high-latitude faunal
evidence and '*O evidence from lower latitudes. His evidence from the south
Pacific, the north Atlantic data of Sancetta et al. (1973), and the Camp Century
(Greenland) ice-core data of Dansgaard et al. (1971) all indicate an initial sharp
drop in temperature at about 75 000 B.P., followed by generally low temper-
atures during the remainder of the Last Glacial. In contrast, Luz (1977) notes a
general decline, with fluctuations, in temperature in the '8Q record as dis-
cussed, for example, by Shackleton & Opdyke (1973). He is presumably right
in intimating that latitude should not affect the evidence from oxygen-isotope
analysis, although there may be some time lag in stabilization of oxygen-isotope
ratios the further the position from the pole. Moreover, if the different analyses
are measuring different responses this could have an effect. However, it would
appear that, in fact, the differences may not be as great as is suggested. In the
isotopic record there is also a relatively rapid drop in temperature, or increase
in the polar ice mass, at the beginning of the Last Glacial. There is evidence for
this in the Caribbean and the Pacific (Shackleton & Opdyke 1973). Thereafter
Shackleton (1975) emphasizes that the establishment by Emiliani (1955) of
stage 3 corresponding to an interstadial is easily recognizable in the oxygen-
isotope record and that there is evidence for a climate significantly different
from that of the preceding and succeeding maxima. The data given by Sancetta
et al. (1973) also suggest a certain amount of recovery during the middle part of
the Last Glacial. Where these data differ from those given by Shackleton &
Opdyke (1973) is in suggesting that the first glacial maximum was as severe as
the last. However, it is interesting to note that in his generalized temperature
curve Emiliani (1972) shows precisely this situation. It would seem, therefore,
that although the details vary, the overall pattern is the same. In terms of actual
sea-surface temperature differences between glacial and interglacial maxima
there is remarkable consistency, with a suggested 3°C in the Caribbean
(Shackleton & Opdyke 1973), 5 °C in the south Pacific (Luz 1977) and Emi-
liani’s (1972) generalized curve showing an amplitude of about 6 °C. A similar
situation off the shores of southern Africa is suggested. Vincent (1972) has used
changes in the distribution pattern of planktonic foraminifers to postulate a
5 °C rise in temperature at about 10000 B.P. where she recognizes the
Pleistocene—Holocene boundary.

Hays et al. (1976), also using a version of the Imbrie et al. (1973) method,
have created a reconstruction of the Atlantic and western Indian Ocean sectors

330 ANNALS OF THE SOUTH AFRICAN MUSEUM

of the Antarctic Ocean at 18 000 B.P. One of the most notable features is that
there is far less variation in the volume of sea-ice from summer to winter at
18 000 B.P. than there is today. This would presumably indicate less seasonal
variability in climate. It is pointed out (Hays et al. 1976: 361) that there is
effectively no temperature anomaly south of Africa with the position of the
subtropical convergence and the Antarctic Polar Front being in substantially
the same position then as now in this region. However, advance of the latter,
particularly in the Atlantic, resulted in the constriction of the subantarctic zone
with a consequent steepening of the thermal gradient. This will have had a wide
effect, increasing winds and driving cold currents far north of their present
positions. Likewise, increase in sea-ice will have steepened the thermal gradient
to the equator, again causing stronger atmospheric circulation. There would
also have been a general contraction of climatic zones towards the equator
(Van Zinderen Bakker 1976) with the result that temperatures would have
been considerably lower, due in particular to frequent influxes of very cold
polar air in the winters. Moreover, Van Zinderen Bakker (1976) postulates
higher rainfall resulting from an increase in the influence of the prevailing
westerly winds. Bé & Duplessy (1976), on the other hand, provide evidence
from both ‘SO and faunal analysis to show that the subtropical convergence
did, in fact, move northwards to 31°S in the Indian Ocean during the late
Quaternary. This resulted in the weakening of the Agulhas Current so that the
consequent cooling off the east coast of South Africa was greater than that off
the west coast of Australia. Whether Hays et al. (1976) or Bé & Duplessy
(1976) are correct, it is clear that, in both cases, the evidence is in general
agreement with that from the oceans elsewhere in the world. It shows that
considerably colder conditions existed off southern Africa during ‘SO stages 2
and 4, that is, during the early and late pleniglacial of the Last Glacial.

TERRESTRIAL EVIDENCE

In a major attempt to correlate land and deep-sea data for past climatic
fluctuations, Kukla (1977) has pointed out that serious miscorrelations have
arisen from past failure to recognize that terrestrial records were incomplete.
Insisting that only complete, or essentially complete, sequences can provide
adequate information, Kukla (1977: 320) lists three types of land-based deposits
which are likely to provide evidence comparable to the oceanic record. These
are pollen-rich lake beds, continental ice-sheets and deposits of alternating
loess and soils. Van der Hammen et al. (1971) have analysed pollen from the
Netherlands and Macedonia to provide a detailed picture of vegetational and,
by implication, climatic change during the late Cenozoic in Europe. Work by
Wooillard (1978a, 1978b) on the Grande Pile sequence in France provides
evidence of glacial-interglacial cycles during the last 140 000 years which also
emphasizes the complexity and number of climatic fluctuations occurring during
that period. In both cases the evidence can be correlated with the deep-sea
evidence (Kukla 1977: 319; Woillard 1978a: 12). Recent work in Norway

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS aL

(Mangerud eft al. 1979) has correlated the Eemian (Last) Interglacial with
oxygen-isotope stage Se by means of analysis of pollen and marine fossils.

Cores from the continental ice-sheets of both the Arctic and Antarctic have
been examined (Dansgaard et al. 1971; Johnsen et al. 1972; Paterson et al. 1977).
A history of temperature variation has been based on 8'*O in the ice. This is
because the index is dependent on the air temperature at the time of deposition
of the snow of which the ice is composed. There are, however, other factors
influencing isotopic composition which cause Dansgaard et al. (1971: 37) to sound
a cautionary note on direct interpretation. In particular there are problems
connected with the interpretation of the Byrd Station, Antarctica, column
(Johnsen et al. 1972). There is, however, good general correlation between the
different ice-cores (Johnsen et al. 1972; Paterson et al. 1977), although they do
not agree in all particulars. Dansgaard et al. (1971: 52) and Johnsen et al. (1972:
433) have attempted to correlate the evidence from Camp Century, Greenland,
with accepted American and north European glacial terminology; Kukla (1977:
321) has correlated the Camp Century evidence with the Grande Pile evidence,
thus providing the link between the two lines of evidence.

The loess evidence has been discussed in some detail by Kukla (1975,
1977). This evidence is particularly important because the essentially complete
sequences can be correlated with the deep-sea evidence and with the classical
glacial stages (Kukla 1977: 322). Palaeoclimatic information is forthcoming
from analysis of the loess and interstratified soils and of their contained snail
faunas. In permanently unglaciated areas such as central Europe, between the
north European and Alpine glacial regions, a regular sequence of deposits was
built up, with unvegetated loess accumulating during glacial periods and for-
ested soils marking interglacial periods. The loess record has also shown that
the palaeoclimatic record is far more complex than was suggested by the
classical Alpine sequence which has normally been accepted. This record is,
however, in good agreement with the deep-sea record (Kukla 1977: 365).

The general indication is that interglacial conditions, specifically the Last
(Eemian) Interglacial, lasted from about 127 000 B.P. to about 105 000 B.P.
This is correlated with oxygen-isotope stage Se and the Barbados III high
terrace. Cold intervals at about 105 000 B.P. and 90 000 B.P. ('8O stages 5d
and 5b) interrupted a long, relatively warm period which lasted until about
73 000 B.P. ('8O stages Sc and 5a). This is correlated with Barbados II and I
high terraces and the early glacial period of the Last (Wirm or Weichselian)
Glacial. The Amersfoort and Brgrup interstadials of northern Europe can
apparently be correlated with '*O stages 5c and Sa respectively. The Camp
Century ice-core can be correlated with these events and a third interstadial,
equivalent to the Odderade of northern Europe, is also evident (Dansgaard et
al. 1971: 52). The lower pleniglacial, which is shown by Van der Hammen et al.
(1971: 394) to have begun about 60 000 B.P. but may have been earlier (see
above), may be correlated with oxygen-isotope stage 4 and is apparent in the
Camp Century core (at about 65 000 B.P.; Paterson et al. 1977: 511) as well as

332 ANNALS OF THE SOUTH AFRICAN MUSEUM

the Grande Pile column. The middle pleniglacial, including the Moershoofd
(50 000 B.P.), Hengelo (38 000 B.P.), and Denekamp (30 000 B.P.) inter-
stadials (dates from Van der Hammen et al. 1971: 395) is recorded in the
Netherlands pollen diagrams and can be correlated with '*O stage 3. In general,
this period is said to have been less cold and more humid than the preceding:
and succeding periods (Van der Hammen et al. 1971; Kukla 1977: 365). The
upper pleniglacial, '*O stage 2, is well recorded in the pollen and ice-core
sequences (Johnsen et al. 1972; Van der Hammen et al. 1971) as well as in the
loess record (Kukla 1977: 330). The late glacial probably began about 14 000 to
13 000 B.P. (Van der Hammen ef al. 1971: 396) with the Bolling interstadial at
about 12 400 B.P. and the Allergd interstadial from about 11 800 to 10 900
B.P. The colder Older Dryas and Younger Dryas occurred between and after
these interstadials, from 12 000 to 11800 B.P. and 10900 to 10000 B.P.
respectively. This represents the beginning of '*O stage 1. Similar oscillations
are visible in the loess and ice-core records. Thereafter, all lines of evidence
show rapid amelioration of conditions and a return of interglacial climate and
vegetation. There is some indication that the climate was warmer in the first
half of the Holocene than the second (Dansgaard et al. 1971: 48) although the
apparent evidence from the Netherlands is said to represent, in fact, the
increasing destruction of forests by people (Van der Hammen et al. 1971: 396).
The evidence from the Mediterranean (Tenaghi Philippon, eastern Macedonia)
is perhaps of greater relevance for comparison with the southern Cape and it 1s,
therefore, important to note that Van der Hammen et al. (1971: 394) have been
able to correlate this sequence with that from the Netherlands.

The only potentially continuous terrestrial sequences from southern Africa
come from pollen profiles. The first to be examined came from the interior of
South Africa. At Florisbad (Van Zinderen Bakker 1957; Van Zinderen Bakker
& Butzer 1973) the evidence suggests that very warm dry conditions with an
increased Asteraceae element existed from some time before 40 000 B.P. At
about 28 000 B.P. a series of oscillations from very arid to semi-arid warm
conditions took place. From about 25 000 to 19 400 B.P. an increase in grass
indicated a lower temperature, higher precipitation, or both. Livingstone (1975:
268) points out, however, that the 'C dates are not internally consistent, so
that the dating of the events is in some doubt. The profile from Aliwal North
(Coetzee 1967; Van Zinderen Bakker & Butzer 1973) covers the period from
about 12 600 B.P. to 10 000 B.P. and the observed oscillations have been
correlated with the late glacial oscillations observed in Europe (Van Zinderen
Bakker & Butzer 1973: 239). These fluctuations confirm the alternation of
warm dry periods with cooler wetter periods, which was indicated for Floris-
bad. Recent work at Wonderkrater in the Transvaal (Scott & Vogel 1978)
shows the same essential pattern during approximately the last 25 000 years.
Here a transitional period, dated to 11 000-9 000 B.P., is correlated with the
Aliwal North sequence and fluctuations in temperature and humidity are shown
to have continued into the Holocene.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 333

Of particular relevence to the southern Cape are two sequences which
have been studied in some detail. The earlier of these comes from Rietvlei on
the Cape Flats (Schalke 1973) and is said to cover the period from about 51 000
B.P. to the present, although the dating is insecure because of lying near the
maximum range of the '*C method. According to Schalke (1973) the middle
pleniglacial is represented by five intervals, two of which are thought to have
been wet and three dry. During the former, dated to approximately
45 000-40 500 B.P. and 36 500-33 000 B.P., a mixed Podocarpus forest resem-
bling that of the Knysna region today is suggested for the central part of the
Cape Flats. It is also suggested for the lower part of the upper pleniglacial in
the region of 28 500 B.P. Thereafter evidence suggests dry conditions in the
upper pleniglacial. During the dry phases and the Holocene, dune vegetation or
coastal fynbos replaced the forest. Schalke (1973) explains the changes in terms
of changes in humidity, whether due to increased rainfall or reduced evapora-
tion. Van Zinderen Bakker (1976: 183) is of the opinion that the evidence has
not been correctly interpreted. He points out, for instance, that except during
the Salt River interval (45 000-40 500 B.P.) the evidence for forest in the
region is not convincing (Van Zinderen Bakker 1976: 184) and even here the
impression is that the forest was probably restricted to river margins and did
not occur in the coastal area. There is, however, certainly evidence for change
in the fynbos and grass cover as well as in the dune and marsh vegetation.

The evidence from Groenvlei (Martin 1968, 1969) covers the period from
about 8 000 B.P. From then until about 7 000 B.P. the area appears to have
been covered with a strong Asteraceae element. Tree pollen indicates a
reduction in heath, perhaps by dunes, rather than an increase in forest. From
about 6 800 B.P. until about 2 000 B.P. the forest element is reduced, which
again could be due to sand or to a drier climate. Thereafter, first scrub and then
forest spread quite rapidly, either because of the removal of the sand barrier or
because of an increase in effective precipitation. It is thus not clear whether or
not there were fluctuations in the climate, although amelioration does seem to
have taken place at about 7 000 B.P. and again at about 2 000 B.P. At present
Groenvlei is just within the western limit of the Knysna forest and it is possible
that the evidence indicates, basically, the arrival of essentially modern condi-
tions in that area at that time.

Auxiliary evidence from macrobotanical remains from archaeological sites
is mostly confined to postglacial levels (Deacon 1972: 37) so that, as would be
expected, modern vegetation patterns are mainly reflected. There are, how-
ever, some apparent differences. At Boomplaas A, for example, Pappea
capensis fruits were found in levels dated to about 2 000 B.P.; this tree does not
grow within a distance of 10 km of the site today (Moffett & Deacon 1977).
Further evidence for vegetation changes at the end of the Pleistocene and
through the Holocene in the foothills of the Swartberg Mountains is provided
by the relative increase in the frequency of Acacia karroo charcoal found in
archaeological sites (H. J. Deacon 1979 pers. comm.) There have been signi-

334 ANNALS OF THE SOUTH AFRICAN MUSEUM

ficant changes in the dominant species in the woodland vegetation of this area
and, notably, A. karroo has been able to extend its geographical range in the
Holocene (H. J. Deacon 1979 pers. comm). This latter pattern is of wider
significance (Acocks 1975: 8) and Deacon considers it to be related to dynamic
adjustments in the distribution of individual taxa in response to warmer
Holocene climates.

Geomorphological evidence provides further climatic data specific to
southern Africa. Most particularly there is evidence for cold conditions during
the Upper Pleistocene, partly because, as Flint has pointed out (1976), this
evidence is more obvious and partly because most of the last 100 000 years
appear to have been colder than the present. At high altitude in the Drakens-
berg Mountains and Lesotho periglacial phenomena indicate temperature drops
of between 5,5°C and 9°C (Van Zinderen Bakker 1976) while oxygen-isotope
work on speleothems in the central Transvaal (Talma et al. 1974) suggests a
temperature 9°C lower than the present during the period 30 000-20 000 B.P.
On the south coast, or the coastal foreland of the time, frost-fractured debris
related to the Last Glacial has been described from Nelson Bay Cave (Butzer
1973; Butzer & Helgren 1972) and Die Kelders 1 (Tankard & Schweitzer 1974).
A postulated temperature drop of 10°C (Butzer & Helgren 1972) to account for
these phenomena is, however, perhaps excessive. Frequent very cold periods,
as suggested by Van Zinderen Bakker (1976), could possibly have been
responsible and would be more in keeping with the general evidence than an
average drop of such a magnitude. Evidence for higher temperatures during the
Holocene has been provided by isotopic analysis of molluscs from Nelson Bay
Cave (Shackleton 1973).

As far as changes in relative humidity are concerned it would appear that
the southern Cape was generally out of phase with the interior of South Africa
(Van Zinderen Bakker & Butzer 1973; Butzer et al. 1978). Inland, as was also
suggested by the pollen analyses, the evidence suggests wetter conditions
during colder periods. At Alexandersfonteinpan Butzer et al. (1973) have
postulated a rainfall approximately double that of the present at about 16 000
B.P., assuming a temperature depression of 6°C relative to that of the present.
Along the Gaap Escarpment a subhumid climate is also postulated for the
period 21 000 B.P. to 14 000 B.P. In the southern Cape, on the other hand, the
indication is that during the period approximately 70 000 B.P. to 10 000 B.P.
the climate was essentially dry (Butzer et al. 1978: 334). A wetter period is,
however, inferred for the middle pleniglacial at Nelson Bay Cave (Butzer &
Helgren 1972; Van Zinderen Bakker & Butzer 1973) and leaching of the
Boomplaas A deposits after the last glacial maximum might have been due
either to increased rainfall or to increased run-off (H. J. Deacon 1979 pers.
comm.). Evidence for wetter conditions at Die Kelders 1 (Tankard & Schweit-
zer 1974) has not been dated but may also refer to the middle pleniglacial.
Climatic change in southern Africa continued into the Holocene, as Butzer
(1974) has pointed out. Butzer et al. (1978: 333) have divided the Holocene of

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 335

the southern Cape into three units. The early Holocene, up to about 4 200
B.P., was relatively dry with a more open vegetation than that of the present,
and considerable aeolian activity. The period from 4 200 to 1 000 B.P. was
apparently considerably wetter, with an increase in bush and forest. Thereafter,
the effects of drier conditions were enhanced by increasing interference by
man. The earlier Holocene was wetter in the interior but after about 4 000 B.P.
the climate was similar to that of the coastal region. It is of some interest to
note the suggestion by Morner (1978) that changes in aridity may have been
due, at least in part, to palaeogeoidal changes which would have altered the
level of the ground-water table and not solely to climatic changes. Whatever
the method responsible, it is perhaps likely that the resultant conditions would
have been the same or similar. It need not, therefore, affect the type of
qualitative or relative assessments of conditions being attempted here.

The macromammalian evidence from the southern Cape has provided a
clear, although generalized, indication that during the Last Glacial the vegeta-
tion of the area must have been substantially different from that occurring in
the Holocene and the recent past. Remains of large grazing antelope and zebra
predominated in the glacial samples. Such species require relatively extensive
open grassland; they were not and could not have been present in the southern
Cape during the recent past. From this Klein (1972a etc.) has argued that
during glacial times grass spread over much of the area. Under interglacial
conditions the vegetation would have largely comprised scrub and bush, with
forest in suitably watered and protected areas. The predominance of small
browsing antelope in samples supposedly pre-dating the Last Glacial from
Klasies River Mouth (Klein 1976) and from Boomplaas A (Klein 1978) as well
as from the postglacial period from Byneskranskop 1 (Klein 1981) and Boom-
plaas A (Klein 1978) is taken as evidence for this change in vegetation. These
trends would certainly be in keeping with those recorded in Europe, although
fluctuations were apparently less extreme in southern Africa.

CORRELATION OF MICROMAMMALIAN AND EXISTING EVIDENCE

The general framework of climatic change during the last 125 000 years is
well established, as has been discussed above. This framework is based upon
quantitative data with wide application derived from more than one discipline
of marine science. Micromammalian evidence currently provides non-
quantitative data of restricted application. As such it must be placed within the
general framework before it can realize its full potential. Whether or not it may
be interpreted in terms of the framework also provides some test of its validity.
Beyond this it is expected that the micromammalian data will provide more
detailed regional information comparable with and complementary to that
already available from other sources. In order to assess the contribution of the
micromammalian evidence at this level it must be checked against that forth-
coming from the geomorphology and palynology of the area. Comparison at an

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ANNALS OF THE SOUTH AFRICAN MUSEUM

even greater level of detail is made possible in the present case by the existence
of macromammalian evidence from the same archaeological sites which have
produced the micromammalian material being examined in this study.

GENERAL FRAMEWORK

The general framework consists of the '*O stages of Emiliani (1955, 1966)
which have found validity on a world-wide basis, as was discussed above. To
this framework has been matched the major divisions of the last glacial cycle,
based on work done in Europe. Within these divisions fluctuations or oscilla-
tions of lesser intensity or shorter duration have been identified for the
northern hemisphere from various lines of evidence.

In the present section it is intended to show that the micromammalian
evidence from the southern Cape correlates well with this general framework.
Table 47 illustrates how the data from Boomplaas A, Byneskranskop 1 and Die
Kelders 1 can be assigned to the '*O stages and the major divisions of the
glacial cycle. It is not to be expected that there will be any detailed agreement
between southern African sequences and those from Europe. It is suggested,
however, that a generally comparable series of events appears to have occurred
in both areas.

The Boomplaas A evidence provides the clearest picture because the
sequence is the longest. The precise position of the lowest levels cannot be

TABLE 47

Generalized correlation of palaeoenvironmental sequences for Boomplaas A, Byneskranskop 1,
and Die Kelders 1 Middle Stone Age with 180 stages and major divisions of the last glacial

cycle.
BNKI DKIMSA

180) last glacial cycle

1 present interglacial

late glacial

ponctoeiiee Juanes He yy
ye, upper pleniglacial
—=— 32 000 BP.

——64 000 B.P. -

unit 3
4 lower pleniglacial wit So | a ne
—75 000 B.P.-
Sa early glacial 0 | ee ee

See Tables 44-46 for details of palaeoenvironmental units.
Dates after Shackleton 1975.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 337

ascertained at present, but it appears most likely that levels LOH and OCH
represent part of '*O stage Sa, the early glacial. This is because relatively mild
conditions apparently pertained at the time and it seems probable that this can
be correlated with the Odderade interstadial in Europe. Thereafter, the lower
pleniglacial, '*O stage 4, is represented by levels BOLS up to BOL1. Palaeo-
environmental unit 4 (Table 44) corresponds well with the middle pleniglacial,
'8O stage 3. Apparently warmer periods, such as at the time of levels BOL and
BOLI and again in levels BP2 and BP3, may possibly be comparable to the
interstadials Moershoofd, Hengelo and Denekamp. Although it is not possible
to be precise about the correlation, it is clear that fluctuations were occurring in
the south which were very similar to those in the north. Moreover, Van der
Hammen ef al. (1971: 395) note that the climate of the middle pleniglacial was
generally less cold and more humid than that of either the lower or the upper
pleniglacial; the Boomplaas A micromammalian evidence indicates just this for
the southern Cape. Palaeoenvironmental unit 3 can be correlated with the
upper pleniglacial, '*O stage 2. The moderate conditions recorded for level BP1
may correspond to an interstadial at this time in the European sequence but,
again, it is not possible to make any close correlation. Late glacial oscillations
are well documented in Europe with the Bglling and Aller@d interstadials
alternating with the Older and Younger Dryas stadials. At Boomplaas A a
period of amelioration in level CL, palaeoenvironmental unit 2, represents the
late glacial at the beginning of '*O stage 1. This was followed by a period of
fluctuation at the beginning of the Holocene, in level BRL, which may be part
of the same phenomenon that occurred in Europe.

The Byneskranskop 1 sequence may be correlated with '%O stage 1. It is
almost entirely Holocene, but radiometric dating shows that some of the lowest
levels were deposited during the Upper Pleistocene. The micromammalian
evidence (Table 45) suggests that levels 19 and 18 may be divided off in this
way and represent the late glacial. A certain amount of fluctuation, but of
lesser intensity, was apparent within the Holocene sequence here as it was at
Boomplaas A. This, however, will be discussed in the next section.

The evidence from much of the Die Kelders 1 M.S.A. sequence suggests
moderate conditions, as was discussed above. It would seem that palaeo-
environmental unit 3 (Table 46), comprising the period covered by levels 14 up
to 12, represents the last part of the lower pleniglacial, '"O stage 4. Palaeo-
environmental unit 2, the period covered by levels 11 up to 3, appears to be
correlated with the middle pleniglacial, '*O stage 3. The beginning of the upper
pleniglacial, '*O stage 2, is then probably represented by palaeoenvironmental
unit 1 but the site apparently ceased to be occupied before the last glacial
maximum. The suggestion that the micromammalian evidence indicates lower
sea-levels in the lowermost and uppermost levels than in the central levels
would be in accordance with this general division. The top two levels may
indicate the same interstadial conditions as those perhaps represented by
samples from Boomplaas A levels BP3 and BP2 but, as was pointed out above,

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ANNALS OF THE SOUTH AFRICAN MUSEUM

it is not possible to make good correlations with the European interstadials of
this period.

SOUTHERN AFRICAN EVIDENCE

It has been pointed out that the evidence suggests that climatic changes in
the southern Cape were different from those in the interior of South Africa
(Butzer et al. 1978). This is perhaps not unexpected in view of the modern
situation where the climate is different in the two areas. However, although
different in direction, the changes tend to have been contemporary, which,
again, may be expected. The micromammalian evidence shows some agreement
with the timing of change as indicated by the palynological evidence. It also
tends to support other evidence for southern Cape conditions.

It would appear that it is not really possible to compare the climatic
evidence from the Rietvlei palynological sequence with that from the micro-
mammalian data for Boomplaas A. The resolution is different in the two cases
and the dating, being near the limits of the radiocarbon range, is correspond-
ingly uncertain. It may be that the drier Milnerton interval could be correlated
with BPA level BOL, but there appears to be no correlation between the drier
Killarney interval and the evidence from the middle pleniglacial levels at
Boomplaas A.

At Byneskranskop 1 the evidence suggests that the period from about
6 400 B.P. to 3 900 B.P. was relatively dry. This agrees with the evidence for
Groenvlei for the second half of the Holocene; it is harder to reconcile the
evidence for the first half, although it is possible that the Boomplaas A
evidence is in agreement. Possibly the difference between the suggested climate
before and after 6 500 B.P. at Byneskranskop 1 is not great. Equally, it has to
be borne in mind that the estimations are relative and it is difficult, on present
evidence, to know what this would mean in absolute terms. It is also possible
that an adjustment may need to be made to the micromammalian interpreta-
tion, but this will have to await further data.

The types of vegetation which have been shown to have occurred on the
Cape Flats and at Groenvlei cannot be shown to have existed in the vicinity of
the archaeological sites. This is not unexpected, however, since the vegetation
is different today and the precise form of the vegetation tends, in any case, to
be very regional due to its reponse to differing conditions. It may ultimately
become possible to postulate what would be the equivalent vegetation for
different areas under different climates, but for the present there are insuf-
ficient data.

The geomorphological data provide evidence for changes in both tempera-
ture and humidity. As was mentioned, data concerning the former generally
indicate colder conditions. The micromammalian evidence from Boomplaas A
and Nelson Bay Cave gives convincing support for the other evidence for cold
conditions during the upper pleniglacial, especially the last glacial maximum.
There does not appear to be other evidence for a cold climate during the lower

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 339

pleniglacial, except that forthcoming from Die Kelders 1 which Tankard &
Schweitzer (1974) have recorded. The micromammalian evidence tends, how-
ever, to suggest that the period of maximum cold in the lower pleniglacial was
not represented at this site. It is difficult, on the basis of present evidence, to
reconcile this with the presence of éboulis secs in levels 12 up to 10. The wetter
conditions indicated for the middle pleniglacial at Nelson Bay Cave, and
probably Die Kelders 1, are also suggested for the Cango valley at that time by
the micromammalian evidence from Boomplaas A. There is, however, no
micromammalian evidence to compare with the geomorphological evidence
from Boomplaas A for wet conditions at the end of the upper pleniglacial.
Possibly these were caused by an increase in rainfall preceding the increase in
temperature during the late glacial and it may be that the situation existed for a
short period and, thus, did not affect the micromammals. Possibly, also,
increased run-off rather than increased precipitation could cause the pro-
nounced leaching (H. J. Deacon 1978 pers. comm.) but not influence the
micromammalian species. The geomorphological and palynological evidence
for the Holocene has been correlated by Butzer et al. (1978). The remarks
above in reference to the palynological evidence may, therefore, be taken as
applying equally to the geomorphological data.

MACROMAMMALIAN EVIDENCE

Comparison at the most detailed level may be made with the evidence
from macromammalian material examined by Klein (1972 et seq.) from the
sites which are discussed in this study. Klein’s pioneering work has been
hampered by small minimum numbers and by a lack of comparative data from
other lines of investigation; for this reason he has been obliged to restrict his
palaeoenvironmental interpretation of the data to general statements (Klein
1972b, 1976, 1978, 1981). The problem of numbers remains, of course, but now
there are comparative data from the small mammal evidence. It is, therefore,
possible to examine the macromammalian evidence from a different angle and
perhaps to provide complementary confirmation of the interpretation of the
two sets of data, thereby strengthening the validity of faunal investigations for
palaeoenvironmental reconstruction.

It is not appropriate here to consider in any detail the problems connected
with macromammalian samples and their interpretation. Many of the con-
siderations discussed above apply equally to macromammalian evidence and
may be considered in this connection. It is, however, necessary to mention a
few points which could have a direct bearing on the interpretation of the
present samples. In the first place, because the samples are small it is particu-
larly relevant to bear in mind that the absence of certain species may well be
due to the smallness of the sample rather than to changes in the environment.
The composition of the sample will almost certainly have been biased by
selection on the part of the people whose food debris the sample represents.
Moreover, there is the possibility that hunter and prey may not have occurred

340 ANNALS OF THE SOUTH AFRICAN MUSEUM

together in a particular area at a given time. In other words, if the game animal
is migratory and the hunter moves seasonally between the coast and inland, as
is suggested by Parkington (1972, 1976) for the west coast and has been
documented orally for the south coast (Deacon 1969: 163), it is possible that
man and animal could both visit the same place but at different times. While it
is perhaps unlikely that an efficient group of hunter—-gatherers would know so
little about game movements as to allow this to happen, there remains always
the possibility that these people could have chosen deliberately not to have
hunted certain animals.

Interpretation will undoubtedly become more refined when it is possible to
identify the high proportions of presently unidentified bovid material from
Boomplaas A (Klein 1978), Byneskranskop 1 (Klein 1981) and Die Kelders 1
M.S.A. levels (Klein unpublished). It is unfortunate that this material cannot
be used for interpretation at present because the groupings do not coincide
sufficiently closely with usable ecological groupings such as those of Jarman
(1974). Meanwhile, it is clearly necessary to group the samples of identified
material in order to obtain adequate numbers in the samples. Klein (19725,
1976, 1978, 1981) has been obliged to group the samples according to culture-
stratigraphic units because he had no other basis for division. This has the
unfortunate effect of forcing cultural change to coincide with environmental
change. The micromammalian evidence suggests, in fact, that this close correla-
tion is fallacious. For this reason, and in order to facilitate comparison, Klein’s
data were regrouped according to divisions indicated by the micromammalian
evidence. This exercise does not invalidate the general conclusions reached by
Klein but does allow a different approach to a more detailed interpretation. It
is clear, however, that grouping on this scale, whatever the basis, obscures
many realities of change and that there can be no substitute for adequate
samples as a prerequisite for reliable interpretation.

Boomplaas A

Table 48 gives the data for perissodactyls and artiodactyls from Boomplaas
A, grouped as nearly as possible in accordance with the micromammalian
divisions (Table 44). As Klein (1978: 68) has pointed out, the samples for the
lower half of the sequence are very small so that, until the full sample has been
analysed, the evidence is uncertain. Klein (1978: 68) has suggested that the Last
Interglacial is represented in the lowest levels, with a high incidence of smaller
bovids suggesting closed vegetation similar to that of the Holocene and the
present. He further suggested that the evidence indicates an expansion of grass
during the last glacial maximum and then a gradual closing of the vegetation
during the late glacial and into the Holocene.

In an area such as the Cango valley where there is complex topography, it
is necessary to determine where individual species are to be found. In the
macrofauna there are four species that are likely to occur on the hillsides rather
than on the valley floor. Tragelaphus strepsiceros (greater kudu), Redunca

341

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS

so1dods jOux yy,

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= 80°L os‘ZI 16 pr‘? ore ad O3IP]
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rr6l I8€Z 6T°9 17s evIZ 6 98S7Z gB0°TE j[ewis—“japur seprAog
Sah Cog nie ae ee. EES Zi P09 dooys dIsowop Sa14D S1IAQ
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Sp ATAVL

342 ANNALS OF THE SOUTH AFRICAN MUSEUM

fulvorufula (mountain reedbuck), Pelea capreolus (grey rhebuck or vaal rib-
bok), and Oreotragus oreotragus (klipspringer) all occur in rocky or hilly
country (Ferrar & Walker 1974: 142; Jarman 1974: app. 1), although the first is
not confined to it (Bigalke 1978: 1028). If the Equus species were E. zebra
(mountain zebra) it would also have occurred in rocky mountainous terrain
(Bigalke 1978: 1019). The remaining species were probably to be found on the
valley floor, or, perhaps, on gentler slopes such as are found near the head of
the valley. It seems, therefore, that the majority of the evidence refers to this
area which does not constitute a very high proportion of the ground in the
vicinity of Boomplaas A. Nearer the top of the valley, where the terrain
becomes less rugged, it may have provided a larger acceptable habitat for those
species normally to be found on plains. It has, of course, always to be borne in
mind that the numbers of individuals represented in the samples need not
suggest large herds, especially considering the long, or relatively long, periods
of time represented. The micromammalian evidence suggests that there was
always some grass on the valley floor, so that there might have been other
reasons for the changing composition of the macromammalian fauna.

There is undoubtedly a high proportion of Raphicerus spp. in the lowest
levels and their distribution within these levels follows closely the suggested
climatic fluctuations given in Table 44. It is possible, therefore, that the
evidence could indicate interstadial conditions rather than full interglacial
conditions. From this point of view it is notable that the total proportions of
small bovids (sensu Klein 1978) in the lowest levels is rather lower than that in
the Holocene levels, which would suggest that the situation was not entirely the
same in both periods. It would, of course, also assist interpretation if the two
species of Raphicerus could be distinguished. The presence of Alcelaphus
buselaphus caama (red hartebeest) or Connochaetes gnou (black wildebeest)
suggests fairly open grass or open mesic grassland or shrubs, with some trees in
the case of the former (Bigalke 1978: 1035; pers. comm.). Antidorcas marsupia-
lis (springbok) generally occurs in drier country with open, short grass or
shrubs (Bigalke 1978: 1025) but also makes some use of mesic grassland (R. C.
Bigalke 1979 pers. comm.). Pelea capreolus indicates short, open shrubby or
grassy vegetation on the hills (Bigalke 1978: 1030). This does not disagree with
the micromammalian evidence, and the apparent openness of the vegetation is
consistent with the suggestion that a glacial maximum occurred during the time
the BOL levels were accumulated.

The large mammal evidence tends to confirm the suggestion that the
material from OLP and BP levels indicates intermediate conditions. The
evidence is very slight but proportions of small and small medium bovids are
both intermediate. The high proportions of Alcelaphus buselaphus caama or
Connochaetes gnou are interesting and suggest that mesic conditions with open
grass or shrubs prevailed.

It may be possible that the preponderance of large medium bovids during
the glacial maximum is due to their having sought refuge either from the

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 343

extremes of temperature or from reduced grazing in the Little Karoo at that
time. If the Cango valley is considered as a refugium, it need not necessarily
mean that there was a great change in the vegetation as it affected these
species. In fact, the indication would be that less change occurred in the Cango
valley than in less sheltered places. Moreover, the low numbers of large bovids
involved (Klein 1978) would not have required extensive grass for their sus-
tenance. The reduction in the proportion of Raphicerus spp. would then result
from improved availability of the larger species. The presence of Syncerus
caffer (buffalo) indicates that there remained a certain amount of dense
vegetation during this period, as was also suggested by the micromammalian
evidence. Some endorsement of the hypothesis of apparent rather than real
change is provided by the hillside species which show a different pattern of
representation. Equus sp. (assuming EF. zebra), Pelea capreolus and Oreotragus
oreotragus all occur in approximately the same proportions during the glacial
maximum (upper pleniglacial) as they do during the later Holocene (DGL
unit). These species are not migratory (Jarman 1974) and may be taken as
indicative of prevailing conditions near the site during the periods under
consideration. According to this evidence there was little difference in vegeta-
tion during the two periods. If the hillside species are unable to reflect change it
could be argued that the changes in the valley floor species cannot be a simple
reflection of vegetational change at this site.

The beginning of a reduction in proportions of open-country species such
as Alcelaphus buselaphus caama or Connochaetes gnou and Antidorcas sp. in
the late glacial (CL unit) can presumably be taken as indicative of a reverse
migration of these species back to the Little Karoo as conditions ameliorated.
Although there is a proportional reduction in these species, there is an increase
in actual numbers of individuals represented and it may be that milder condi-
tions caused overcrowding which intiated the move out of the valley. At the
same time an increase in 7aurotragus oryx (eland) and of Syncerus caffer may
perhaps be taken as indicative of an increase in forbs and shrubs in the first
instance and of riverine bush in the second instance. This latter situation may
account for an increase in Hippotragus spp. which tend to need more cover
(Bigalke 1978: 1033). Both would be consistent with the micromammalian
evidence. It is interesting to note in this context that Tragelaphus strepsiceros
has so far been recovered only from deposits of this age and those next
youngest. The presence of this species indicates thickets and other dense
shrubby vegetation, probably on the hillsides. In general the macromammalian
evidence for this period gives a rather less clear indication of the intermediate
nature of conditions than do the micromammalian data, but it is still present
and thus confirms the pattern.

In the BRL unit Holocene conditions are shown by both sets of data to
have become established. There is continued reduction or disappearance of the
open-country forms (Alcelaphus buselaphus caama or Connochaetes gnou and
Antidorcas sp.). There appears to have been a higher than usual emphasis on

344 ANNALS OF THE SOUTH AFRICAN MUSEUM

hillside species from this time, at least until about 2 000 B.P. and possibly after
this. This phenomenon is probably connected with the mechanics of acquiring
food rather than with palaeoenvironmental change. Thus, if it is accepted that
large gregarious animals, which have non-concealing habits in an open habitat
(Estes 1971: 174), are potentially easier to find and kill than small solitary
animals, which have concealing habits in a closed habitat (Estes 1971: 174), and
also offer a better return for expended effort, then it is likely that as the larger
species disappear the hunters will turn to the next largest available species. In
this case this happens to be the hillside species. If this argument is correct, it
would suggest that in general changing proportions of the larger species may
reflect real changes in availability of species; the smaller the species the more
likely it is to be present throughout the period, and apparent changes in its
proportion to reflect only availability of the larger species. This would perhaps
not affect the overall interpretation, but it is important to bear in mind the
presence of such potential biases in the samples.

Byneskranskop I

Table 49 gives the percentage representation of perissodactyls and artio-
dactyls in the Byneskranskop 1 samples, grouped according to the results of the
micromammalian analysis. It is clear in overall terms that the larger ruminants
tend to predominate in the lower half of the sequence and the smaller
ruminants in the upper half, as Klein (1981) has pointed out. This change had
been taken as indicative of the replacement of extensive grass by a more bushy
vegetation (Klein 1981). Closer examination of the evidence suggests that both
the large and the small mammal data provide a very similar indication of past
environmental change. It is noticeable, for instance, that both Taurotragus
oryx (eland) (although not positively identified from the upper levels) and
Syncerus cf. caffer (buffalo), together with the undetermined ‘large bovid’
category, occur in approximately equal proportions in all three time periods
recognized. In the case of the first species this may be because it has very wide
habitat tolerance (Bigalke 1978: 1030), grazes and browses (Hofmann &
Stewart 1972: 236) and can do without water (Bigalke 1978: 1030). This would
suggest that it would be impervious to relatively minor environmental changes.
S. caffer, on the other hand, prefers dense vegetation, being essentially a
woodland species (Lamprey 1963: 76), and is a fresh-grass grazer dependent
upon water (Hofmann & Stewart 1972: 231). The riverine vegetation, which is
present throughout according to the micromammalian evidence also, could
presumably have proved a suitable habitat during the period under discussion.
The presence of this species in the area does, however, also indicate that there
was grass at all times, as is indicated by the micromammalian evidence.

Of the species which were more numerous in the lower levels, the majority
seems to have been dependent upon surface water and this may be taken as
confirmation that this period was relatively wet, as is suggested by the micro-
mammalian evidence (Table 45). Apart from Diceros bicornis (black rhino-

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 345

TABLE 49

Percentage representation of perissodactyls and artiodactyls at Byneskranskop 1 (based on
unpublished data, courtesy of R. G. Klein).

1-4 5-10 11-19
PERISSODACTYLA
ES ee -- 0,84 1,33
uummeeronicde midet. : . »§ « . «re » ww 0,68 1,68 i
rCHSIS™| SG ea — — 0,44
ne el) AY be PS Wa — 0,42 4,89
ARTIODACTYLA
em IS POYCUS . «2 we kl we 4,79 Dae 1,33
eememcwormsaciiopicus 2. . .« » «© «5 « ww 2 — — 0,44
Suidae indet. ar 5 Oil SE a ie eal ie 4,79 2,94 4,00
Hippopotamus Brephibias ee eee cer ity love 0,68 1,26 0,44
TS 6,85 7,56 4,89
ME el — 0,84 PSO)
SES — 0,84 2,67
MemremIOTIWIG. 5 wk ll 0,68 — 1,33
Hippotragus spp... ee oe ea 0,68 ZOE 4,89
Connochaetes sp. | Alcelaphus sp. oe FO eee, aoe ye 2,74 2,10 1p a
ES a — — 2
MEINIOCOIQGUS - 3. wll lw ls 2,05 1,68 1,33
MGOMIITCOIIDCSIFISS 2. ke ws 2,74 0,84 _
CVE TREIQHOUIS” 3) 2. 8 se wl ll 6,16 8,82 1:33
ec SS A a a Ue J 7 8,00
RS SI SS A Sr 0,68 0,84 0,44
Ovis aries : NS Ne Ed vig fed oS. Po Mee oe fies —- _-
Bovidae indet. —smailll Es ae ee ee 19,86 23,95 10,67
ediemeaum . << 2 € ww 4 os 10,27 2,94 4,44
Ieee median = . f « . 4,11 5,46 20,89
Peeeeee eee GP doy eM BU 753 9,66 7,56
N= 146 238 225

See Table 48 for English common names.
* Extinct species.

ceros), which is said to prefer fairly dense vegetation (Bigalke 1978: 1018) but
is certainly not restricted to it (personal observation), these species appear to
have a preference for open grassland (Equus cf. quagga (quagga), Damaliscus
dorcas dorcas (bontebok), Connochaetes spp. (wildebeest) or possibly open
savanna (Alcelaphus buselaphus caama (red hartebeest), Hippotragus leuco-
phaeus (blue antelope) (Klein 1974b: 113)). Redunca arundinum (reedbuck)
indicates taller, denser grass (Ferrar & Walker 1974: 143). This evidence agrees
with that from the microfauna which indicates that, apart from extensive grass
on the plains, there was probably a considerable amount of dense vegetation
along the river and between the river and Byneskranskop itself. Redunca
fulvorufula (mountain reedbuck) and Oreotragus oreotragus (klipspringer) sug-
gest the presence of both grass and shrubs on the hills which, again, is in
keeping with the micromammalian evidence. The occurrence of Phacochoerus

346 ANNALS OF THE SOUTH AFRICAN MUSEUM

aethiopicus (wart-hog), although represented by only one individual, is consis-
tent with the general picture, since it is an open woodland species (Lamprey
1963: 76) with a preference for abundant vegetation including short grass (Hirst
1975: 40). Superficially the later increase in Potamochoerus porcus (bush-pig),
which is indicative of closed bush (Bigalke 1978: 1019), would appear to
contradict the suggestion of wetter conditions during the earlier period. It may,
however, be as Klein suggested (1976: 83), that the earlier people were
disinclined to hunt pigs, which have a reputation for ferocity, although it is
perhaps doubtful whether they can have been harder to catch than Diceros
bicornis. Both could, in any case, have been scavenged—which would negate
the argument. It would seem that from about 6 500 B.P. conditions remained
essentially the same until the site was abandoned. This is not entirely in
agreement with the micromammalian evidence, but it is possible that minor
fluctuations have been reflected by the latter which would not have affected the
macromammals.

Die Kelders 1 Middle Stone Age samples

Table 50 gives the percentage representation of perissodactyl and artiodac-
tyl species in the samples from the M.S.A. levels at Die Kelders 1, grouped
according to the micromammalian evidence. Klein (1975: 265) has pointed out
that large bovids are not well represented in these samples. The high propor-
tion of small bovids would appear to agree with the suggestion that interpleni-
glacial conditions pertained, especially during the central period. There appear,
however, to be contradictions which cannot be readily explained. Connochaetes
sp. (wildebeest) occurs in rather higher proportions in the upper and lower
levels, but Hippotragus leucophaeus (blue antelope) and Damaliscus sp. (har-
tebeest) occur only in the middle levels. Possibly Hippotragus leucophaeus
indicates that there were some trees during the central period but not during
the earlier and later periods; the significance of the other two genera may
depend upon which species is represented. Syncerus caffer (buffalo) occurs only
in the middle levels but ?Diceros bicornis (black rhinoceros) occurs in higher
proportions in the upper and lower levels, despite the fact that both species
have a preference for fairly dense vegetation (Lamprey 1963). There are,
however, at least two differences; Syncerus caffer is a migratory grazer (Jarman
1974: 223; Hofmann & Stewart 1972: 231) whereas D. bicornis is a sedentary
browser (Lamprey 1963: 77). It is possible that suitable grass was available for
S. caffer only during the central period, or that its visits to the vicinity of Die
Kelders 1 coincided with those of the people only during that time, or that the
earlier and later samples are too small to include this species. The increase in
Tragelaphus strepsiceros (greater kudu) and in ?Pelea capreolus (grey rhebuck)
in the upper levels can probably be accepted as endorsing the micromammalian
evidence for increased scrub on the hillsides during this period on the basis of
the ecological data given by Ferrar & Walker (1974: 142) and Bigalke (1978:
1030). Redunca fulvorufula (mountain reedbuck) may, on the other hand,

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 347

TABLE 50

Percentage representation of perissodactyls and artiodactyls in Die Kelders 1 Middle Stone
Age levels (based on unpublished data, courtesy of R. G. Klein).

1-2 3-11 12-14
PERISSODACTYLA
a 2,78 1,16 2,78
I SS ee — 0,29 —
ARTIODACTYLA
Weroppoporomusamphibius..§ . «§ « « «© « 6» «© 1,39 1,74 2,78
MUUOGHST 2 lw ew 1,39 0,87 —
Syncerus caffer .. A ot ea te eee ae 1,16 —
Tragelaphus pireniiceros 5¢ MATS Sa We eee A oe les 1,39 0,58 —
TT a 8,33 4,93 13,89
Meaumcaorunamum. .-. .« « « «© « «© «© «2 so 1,39 1,45 —
Muemeerrmvorguia. lw ll — 0,29 —
Weeeperrarus lencopndeus® . . . 1. 6. 6 8 ee = 1,74 —
ENE Seller Chie ck ee a 2,78 1,45 2,78
Damaliscus sp. . » Wee se Wl ee ee — 0,29 —
Antidorcas cf. oustralis® a fi or Salen Sa Nolen} oul lle 2,78 1,45 —
Oeormeens orcoirdeus:. 6. kk lk 2,78 3,48 2,78
Wiaomicemmsmmcianolis.. . . 2 6 te we ll 16,67 33,04 16,67
?Pelea capreolus RE en Ciel Rm ms 6,94 4,06 —
os 2 | nc 22,22 20,87 21,518
= Meat we 1 6,38 5,56
faree MedIUM = ss, 1,39 4,93 5,56
ToS ee SS Sea ae ae 5,56 2,90 5,56
very large Et OO Mae eas kel ahs Pt 6,96 13,89
N= “72 345 36

See Table 48 for English common names.
* Extinct species.

suggest tall grass with shrubs on the hillsides (Bigalke 1978: 1031) during the
central period. Bigalke (1978: 1031) notes that this species is to be found in
more arid areas. This is interesting in view of the fact that the only specimen
from Die Kelders 1 M.S.A. levels was recovered from level 3, one of the levels
which it is tentatively suggested, on the basis of the micromammalian evidence,
might have been deposited at a time of a fairly dry climate.

In general, the micromammalian and macromammalian evidence is in
accord for this site, even to the extent that both sets of data suggest complica-
tions in the interpretation. The basic picture is reasonably clear but it would
seem that clarification of the details will have to await the acquisition of a
longer sequence with which the Die Kelders 1 sequence may be compared, or
more information on the species, or both.

MICROMAMMALIAN EVIDENCE FOR SITE CORRELATION
As a corollary to the main study the use of micromammalian evidence to
suggest generalized correlations of undated or insecurely dated sites was
investigated. Unlike biostratigraphies which require that stages in the evolution

348 ANNALS OF THE SOUTH AFRICAN MUSEUM

of various species be matched at different sites, the present correlations are
based upon changes in community composition and, therefore, upon environ-
mental change. It may also be possible to indicate approximate dates by
comparison of the micromammalian evidence with that from other lines of
research. Thus a base date for Boomplaas A may be suggested and, within
broad limits, dates for the M.S.A. occupation of Die Kelders 1, after this site
has been correlated with Boomplaas A. Such exercises are not generally useful
for Holocene sequences because it is normally possible to acquire good 'C
dates which preclude the need for indirect methods of dating.

Of the Upper Pleistocene sites in the southern Cape, only two have thus
far provided a sequence of micromammalian samples sufficiently long and
complete to afford the possibility of correlation on this basis. The Boomplaas A
sequence is absolutely dated to the limits of “C dating, but the length of the
period of occupation prior to this can only be estimated. At Die Kelders 1 no
reliable dates have been obtained for the M.S.A. levels. The fact that these are
separated from the dated L.S.A. levels by deposits representing an unknown
period of time when the site was not occupied, means that neither the date nor
the duration of the M.S.A. occupation is known. Tankard & Schweitzer (1974)
have, however, suggested on geological grounds that the period of occupation
was from about 80 000 B.P. to 35 000 B.P. Deacon (1979) considers that the
base of the Boomplaas A sequence may be dated to about 80 000 B.P., while
Klein (1978) has suggested, on the macromammalian evidence, that the date
may be nearer 100 000 B.P.

The micromammalian evidence tends to support the suggestion (Deacon
1979) that the age of the earliest deposits at Boomplaas A is approximately
80 000 B.P. As was mentioned above, it appears most likely that levels LOH
and OCH were deposited during the early glacial period which lasted until
perhaps 70 000 B.P. The micromammalian evidence suggests that in these two
levels intermediate conditions are represented, and it is perhaps possible to
equate these with the Odderade interstadial at the end of the early glacial in
northern Europe or '*O stage 5a which is characterized as relatively warm. Such
evidence is not, in fact, inconsistent with that from the macromammals (Klein
1978). It is possible that conditions during all the warmer intervals of '*O stage
5 were sufficiently similar to those of the Holocene not to be differently
represented in the macromammalian evidence. It would be possible for this
evidence to refer to a later rather than an earlier warm interval.

Variation in percentage of total variance accounted for by factors in the
analysis of Upper Pleistocene levels at various sites and changing representa-
tion of different species (Fig. 32) indicate a possible correlation between
Boomplaas A and the M.S.A. levels at Die Kelders 1. It is not suggested that
the actual proportions of species are the same at both sites or that factors
explain the same amount of variance at both sites. It is, however, suggested
that changes of proportion and amount of variance within each site display
similar patterns at the two sites. On this basis it would appear that the Die

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 349

o

O.saundersae

30

S.varilla

a 5 6
x. 10O:000 yrs BP.

Fig. 32. First alternative correlation of the Boom-
plaas A and Die Kelders 1 M.S.A. sequences. BPA
levels OLP to BOLI-3 and the DK1 M.S.A.
sequence correlated using individual species. (Solid
line = Boomplaas A; dashed line = Die Kelders 1.)

350 ANNALS OF THE SOUTH AFRICAN MUSEUM

FACTOR 3
%
90 F
"
La
80 om
; ,
\
;
70 ; ;
! \
' \
60 u :
FACTOR |
%
100

FACTOR 4

90

80

70

60
50
FACTOR 2
40 40

30

20

x 10000 yrs BP

Fig. 33. Second alternative correlation of the Boomplaas A and Die Kelders 1 M.S.A.

sequences. BPA levels BP3 to BOLI-3 and the DK1 M.S.A. sequence correlated using

variation in percentage of total variance accounted for by factors in the Oblique rotated factor
structure matrix. (Solid line = Boomplaas A; dashed line = Die Kelders 1.)

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 351

Kelders 1 M.S.A. levels were probably deposited during the same period as
Boomplaas A levels BOLI to BP3 inclusive. The approximate dates for this
period are 60 000 B.P. to 30 000 B.P. The latter is based on bracketing dates of
21 100 + 420 B.P. (UW 300) for level LPC and 32 400 + 700 B.P. (UW 304)
for level BP4. The former date is indicated by the fact that level BOL1 is
approximately midway between level BP4 and the base of the sequence which it
is suggested above is dated to about 80 000 B.P. C determinations UW 305
and UW 308 gave readings of greater than 40 000 B.P. for levels OLP1 and
BOL, while an informal date of 42 000 B.P. for level OLP1 was provided by an
unnumbered Pta determination. If these dates for the Die Kelders 1 sequence
prove correct, they might well not be incompatible with the bone apatite date
of 31 800 *39) B.P. (GX-1717) given by Schweitzer (1970) for level 5.

Establishment of the possible upper date proved more difficult than that of
the lower date. Correlation of M.S.A. level 14 at Die Kelders 1 with Boom-
plaas A level BOLI is good in both sets of diagrams (Figs. 32-33). On the basis
of the individual species (Fig. 32) the upper limit of the correlations was first
thought to lie in BPA level OLP which would have entailed a total M.S.A.
period occupation of about 20 000 years for Die Kelders 1. The levels analysis
(Fig. 33) indicated, however, that the best general fit was obtained by correlat-
ing BPA level BP3 with DK1 M.S.A. level 1. It was considered preferable to
accept this estimate since the factor analysis had involved the whole suite of
species. Moreover, such a correlation is generally consistent when applied to
the individual species. It is perhaps also more reasonable that the period
involved should be nearer 30 000 years in view of the fact that each level of
ccupation is separated from the next by possibly a considerable period of
non-occupation (see Fig. 5). It has been postulated above that much of the
M.S.A. occupation at Die Kelders 1 took place during the middle pleniglacial
of the Last Glacial. That the levels at Boomplaas A with which it is proposed
that they are correlated are similarly interpreted is seen as corroborating
evidence. Moreover, this is not inconsistent with the evidence from sea-level
changes reviewed by Tankard (1976b), which indicates an interstadial sea-level
at —20 m between about 47 000 B.P. and 20000 B.P. It is also in accord
generally with the specific geological interpretation that Die Kelders 1 was first
occupied during a very cold period but mostly during an interstadial (Tankard
& Schweitzer 1974, 1976).

PREHISTORY AND ENVIRONMENTAL CHANGE

In this section it is proposed first to discuss current thinking on the role of
the physical environment in the development of human culture. Then the
archaeological and environmental evidence, as interpreted from the micro-
mammalian data, is examined in an attempt to establish how the generalities
may apply in the specific case of the southern Cape. Particular attention is paid
to determining whether or not there is a pattern of correlation between

Los)
nN
i)

ANNALS OF THE SOUTH AFRICAN MUSEUM

environmental and cultural change in the sites examined. It must be empha-
sized that, besides being specific, this exercise is very tentative and will require
considerably greater quantities of more detailed data before anything
approaching a firm generalization can be reached. Mention should also be
made of the possibilities of using the palaeoenvironmental data to aid in the
detailed interpretation of archaeological sequences and of showing, by an
absence of environmental change, that a switch in resource utilization may be
due to choice rather than necessity, and that industrial development may take
place for reasons other than adaptation to environmental change. These are,
however, more properly archaeological problems and, as such, will not be
discussed further here.

GENERAL CONSIDERATIONS

There would appear to be general agreement that the natural (physical and
biological) environment has influenced the course of human cultural develop-
ment. What is not generally agreed upon is the extent and nature of that
influence. It would seem most likely, however, that, on the one hand, the
extent will have varied through time, being gradually reduced as men become
more technologically advanced; on the other hand, the nature of the influence
might well have varied according to the nature of the environment as well as to
the degree of technical development. From another point of view it may
perhaps be suggested that the nature of the environment could have affected
man’s ability to advance technologically. In any case, it could be argued that
environmental change will not only act as a catalyst for cultural change but also
make it easier for archaeologists to perceive the connection or relationship
between the two. More particularly, in the present study, the concern is to
assess the ways in which purely environmental change may be seen to have
affected cultural development. Because this is primarily a study of the natural
environment, only aspects of human culture possibly affected directly by
environment will be considered. The complexities of culture as a whole are not
relevant and, as such, are examined only generally in order to provide the
context for the main discussion.

A major general controversy has centred around the extent to which the
natural environment has had an overall effect on the total culture of a people
(Trigger 1971). One school believes, in effect, that natural environment will
determine the nature of the society’s economy which, in turn, will have a major
effect on the remaining aspects of that particular culture. Trigger (1971: 325)
has labelled this ‘determinant ecology’, and it would appear that one of the
main problems has been a tendency to take this idea to logical conclusions
which, at best, do not fit the evidence and, at worst, are absurd. Undoubtedly
the open-system ecology (Trigger 1971: 329) would seem to provide an intrinsi-
cally more realistic explanation because it emphasizes that a complex of factors
must be responsible for the final result which is, after all, complex itself. It
recognizes also that the human environment involves not only the natural but

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 353

also the cultural environment. This latter comprises interaction with other
groups and societies, with resultant acculturation. In the present study only the
natural, specifically biological, environment is to be considered, which would
suggest that only part of the problem can be examined. In fact, ‘open-system
ecology . . . assumes that developments affecting any one aspect of culture can
ultimately produce further adjustments throughout the system and affect the
system’s relationship with its natural environment’ (Trigger 1971: 330). This is
not the same as suggesting that the environment controls the culture but it does
indicate that a study of the relationship between the system and the natural
environment may prove more informative than may otherwise be expected.

The nature and extent of natural environmental control over the course of
cultural development appear not to have been static. For example, it should be
possible to assume that a group of people who subsist by hunting and gathering
are more directly affected by their environment than a group of people living in
a major city. Certainly climate will determine whether or not there is central
heating and/or air conditioning in the buildings, but this in itself implies a level
of control of the environment which allows almost total effective independence
from the climate. The gradual increase in efficient acquisition of energy up to
this level may be seen as basic to man’s changing relationship with his
environment (Klein 1979). In this process it is both the kind of energy as well as
the amount that are involved. At the beginning of the process the primitive
hunter-—gatherers may be supposed to exist at the stage of acquiring sufficient
energy from food to keep alive. Basically they will be obliged to seek food
where and when it is available and at this level it is to be expected that the
natural environment will exert considerable control over the activities of the
people. Slight modification of the environment in the form of fire, primitive
shelters and clothes may allow marginal expansion into previously uninhabit-
able areas, but it is likely that the availability of food will constitute the major
controlling factor for hunter-gatherer societies, as Harpending & Davis (1977)
have attempted to show with mathematical formulae.

Another point to be considered is that different types of environment may
perhaps have different effects on cultural development. By this is meant that
under harsh conditions various constraints will presumably operate that will be
absent under milder conditions. Alternatively, milder conditions will provide
more options than will rigorous conditions; the measure of choice of available
foodstuffs, mentioned by H. J. Deacon (1972: 33), will be greater. Yellen
(1977) suggests, for example, that the type of environment will affect the nature
of the society living in the area. Thus, under desert conditions where food
supplies are limited and unpredictable, the advantage will lie with groups that
are resilient, that is, adaptable rather than specialized and without too many
internal restrictions. Under milder conditions where resources are predictable
and more abundant, the advantage would presumably lie with specialists.
Through the archaeological record the possiblity exists of examining cultural
response to environmental conditions on a temporal as opposed to a spatial

os)
nN
=

ANNALS OF THE SOUTH AFRICAN MUSEUM

basis. Thus it is possible that under adverse glacial conditions groups of resilient
hunter—gatherers would have had the advantage. Their very adaptability could
also, thereafter, have best fitted them to deal with changing conditions at the
end of the glacial period. Under milder postglacial conditions, specialization
would presumably have been an advantage. Moreover, if, as has been sug-
gested above, the possibilities for specialization are increased under milder
conditions, the archaeological record might be expected to show increased
numbers of regional or local differences. If the people were unable to remain in
one area throughout the year it is to be expected that they would practise a
number of different specializations in different places, as was shown by Coe &
Flannery (1964). Yellen (1977: 265) notes that this phenomenon has been
observed in birds, which may be territorial in summer when food is abundant
but not in winter when food is scarce. As far as prehistoric people are
concerned, it may be that environmental change could well provide the incen-
tive towards technical development. It must, however, be noted that the actual
nature of that development would depend upon the people involved, as
J. Deacon (1978: 108) has pointed out. Whether or not social behaviour could
also be affected, is a more tenuous proposition.

One aspect of specialization may be the focusing of attention on smaller
areas which Coe & Flannery (1964) have called microenvironments. This is not
to suggest that one group of people necessarily concentrated on only one such
microenvironment; indeed, they manifestly did not. Butzer (1971: 7) also
makes the point that reconstruction of the immediate environment of a site is of
major importance. Such reconstruction makes it possible to establish in greater
detail the degree of specialization involved and, as in the case of Coe &
Flannery’s study (1964), the extent to which environmental differences might
have encouraged or delayed improvement in subsistence strategies. They have
suggested (Coe & Flannery 1964) that harsh conditions can delay progress until
some method is found of artificially ameliorating those conditions, such as
irrigation in an arid climate. Thus, people living under basically desert condi-
tions were unable to make the change to full dependence on agriculture until
they found a way of irrigating the crops; instead, cultivation became only one
part of their annual round. On the coast, however, the milder climate allowed
more and closer microenvironments which, together with agriculture, allowed
the population to remain sedentary and to develop village life much earlier than
the inland people. This is another aspect of the effect of environment on
cultural development, which is discussed above, and it would tend to illustrate
the interrelationship of all aspects.

An alternative possibility is that people will concentrate on certain plants
and animals, as has been suggested by Flannery (1968). Because these resour-
ces may be found in more than one microenvironment, different groups of
people may follow the same way of life in spite of living under slightly different
conditions, or one group may cross-cut several microenvironments in pursuit of
a particular resource. By the same token, change in the environment of one site

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS a)

over a period of time need not affect the economy of the people living there,
provided the relevant natural resources, plant or animal, are not affected. This
is an important point which suggests that only environmental change of consid-
erable magnitude will have forced people to change their economy. Changes of
lesser magnitude may, on the other hand, have provided the means or incentive
for voluntary change. It is possible that the micromammalian evidence may
provide some insight into this problem. It may give an indication as to whether
people were operating within or across microenvironmental boundaries and it
could perhaps also, by indicating the amplitude of environmental change,
suggest whether or not industrial changes were likely to have been initiated by
environmental change. Flannery (1967) used the micromammalian evidence to
provide background environmental data, but he did not tie this in with his
interpretation of human exploitation of resources (Flannery 1968). To do this
may provide some useful insight into the mechanics of subsistence.

As far as actual technological adaptation is concerned, it has been sug-
gested that some stone industries can be correlated with particular environ-
ments. For instance, on the basis of distribution, Clark (1963: 360) has
suggested that the Sangoan industry was connected with thicker vegetation and
higher rainfall. He notes later regional contemporary differences in culture
which he attributes to environmental differences, with a suggested correlation
between smaller tools and open country and vice versa (Clark 1963: 362). This
would seem to imply that the tools were used for working the vegetation in
some way. Mazel & Parkington (1978) have, in fact, suggested not only that
there is a correlation between high proportions of adzes and woody plants in
the western Cape, but also that the adzes were used for wood-working. There
is, however, still much to be done in the way of assigning uses to various stone
tools, so that this purely practical aspect has yet to yield much information
concerning environmental adaptation.

The open-system ecology of Trigger (1971) employs a very broad definition
of environment, as was mentioned above. It was also pointed out that in the
present context only a narrow definition was being considered. This implies that
studies of the natural environment can be of use only in selected aspects of
interpretation. For this reason it is as well to attempt to give some indication of
the aspects which data from micromammalian evidence may elucidate. In
general terms it may be suggested that these data will provide evidence to aid
interpretation of human ecology but not, at least for the moment, of human
biogeography; evidence for the relationship of people to their environment but
not of their distribution in the countryside may be provided. This is because the
data provide detailed information about small areas. This will enable some
assessment to be made concerning adaptation to, or utilization of, local
resources by the occupants of the site in question in terms of the contemporary
environment. It will also enable comparison of such adaptation at different sites
in different and similar environments. On the other hand, unless there is
evidence from a large number of sites, it is unlikely that micromammalian

356 ANNALS OF THE SOUTH AFRICAN MUSEUM

evidence will have anything to add on the subject of the distribution of human
beings or their movements from one area to another. The small scale of
micromammalian evidence is not appropriate to providing information concern-
ing activities conducted on such a relatively large scale.

THE SOUTHERN CAPE PROVINCE

It is to be expected that the generalities discussed above will find specific
application in the southern Cape Province. The purpose here will be to
investigate the extent to which the environmental data elicited from the
micromammalian evidence can be used to demonstrate a connection between
cultural and environmental change. In this exercise the main advantage lies in
the fact that the micromammalian evidence .is closely associated with, but
independent from, the archaeological evidence. The result is that the micro-
mammalian evidence is directly relevant but avoids the risk of false correlations
between human activity and environmental conditions and change. As has
already been indicated, it is at the detailed local level that the importance of
the micromammlian evidence will be felt. It is, therefore, at this level that the
data will be examined, although reference to the known archaeological frame-
work for the southern Cape will be made. In much of the following discussion
changes in the lithic assemblages and environmental change, as interpreted
from the micromammalian evidence, are compared in isolation. This has been
done purely as an aid to simplifying the argument; the lithic assemblage is
taken as representative of the culture to which it belongs only because it is
easier to discuss changes in a concrete entity than in an abstract concept. It is
not suggested either that the lithic assemblages represent the sum of archaeolo-
gical reality, as discussed by Hill (1972), or that there is a deterministic
relationship between technology and environment.

As a taxonomic convenience, the Upper Pleistocene and Holocene
archaeological sequence of the southern Cape, and the rest of southern Africa,
has been divided into two major parts, lasting respectively from about 125 000
to 40 000 B.P. and from about 40 000 B.P. onwards (Klein 1977: 120). The
earlier of these is the Middle Stone Age (M.S.A.), a term originally proposed
by Goodwin (Goodwin & Van Riet Lowe 1929), who described two industries
and a number of variants. Subsequently, Klein (1977: 120) has remarked on the
large number of different industries in the general M.S.A. category. Sampson
(1974) has attempted to instil some order by grouping these into complexes.
However, the mere fact that the material has been assigned to numerous
industries is indicative of the variation within the material. This, together with
the occurrence of sites in a wide variety of different habitats (Klein 1977: 120),
could be indicative of the adaptability of the people at that time. One example
is to be found at. Klasies River Mouth where the people made extensive use of
marine resources (Klein 1976: 83). At this and inland sites they apparently
preyed upon small or larger ungulates, depending on which was more readily
available, but showed a seeming reluctance to catch the fiercest animals (Klein

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS B57

1977: 120). Both Klein (1976) and Sampson (1974: 256) have noted that a
change in the most commonly represented antelope in the samples apparently
coincided with the introduction of Howieson’s Poort artefacts. It may be that
both this and the earlier Stillbay—although this has yet to be properly defined
(Sampson 1974: 257)—represent technological adaptations to environmental
change. In general, however, it appears that there was an appreciable trend in
development during approximately 100 000 years.

The apparent trend towards a cyclical effect in the M.S.A. industries may,
in fact, be explained by a need to adapt to oscillating conditions. Sampson
(1974: 248) remarked on the fact that the post-Howieson’s Poort industries
from Klasies River Mouth and Skildergat (Peer’s Cave) show a marked
similarity to the pre-Howieson’s Poort industries. The use of prepared cores,
which is a feature of the earlier cultures, is apparently absent from the later
levels at Klasies River Mouth, but was found at Boomplaas A (H. J. Deacon
1977 pers. comm.). It is possible that both the earlier and the later industries
may represent some form of adaptation to interglacial or interstadial condi-
tions. If the equation of the Last Interglacial with '8O stage Se is accepted, it
could be argued that M.S.A.I and perhaps M.S.A.II at Klasies River Mouth
might be dated to the Last Interglacial. Klein (1976) has suggested that
essentially modern conditions existed at that time, which is to be expected from
what is known elsewhere for that period. Again, in the later M.S.A. there is a
suggestion of conditions similar to those of the present. Klein (1978),;has
suggested that the evidence from the lowest levels at Boomplaas A indicates
essentially modern conditions and consequently suggests a date of about
100 000 B.P. for the base of the sequence. It has been proposed above,
however, that what is represented is probably an interstadial towards the end of
the early last glacial and that conditions, while definitely mild, were not the
same as those of the present. During the intervening period it is suggested that
a colder stadial in the early glacial occurred. Klein (1977: 120) proposed that
the Howieson’s Poort backed blades and segments may have been hafted
which, together with the increase in larger open-country bovids (Klein 1976),
suggests an adaptation to different conditions during that period. Whether or
not the Stillbay industry, if it exists, proves to be another such adaptation
remains to be seen.

It has been observed (Klein 1974) that archaeological deposits in the
southern Cape frequently exhibit a hiatus during the period when the final
M.S.A. may be expected to be replaced by the early Late Stone Age (L.S.A.).
One major exception has proved to be the Boomplaas A sequence (Deacon &
Brooker 1976), where the interface is represented. Although the late M.S.A.
material has not yet been studied in detail, its potential for providing evidence
of adaptive processes is considerable. The M.S.A. is thought to have lasted
until about 30 000 B.P. at Boomplaas A (H. J. Deacon 1978 pers. comm.).
This is in itself an important point because it suggests that the M.S.A. people
there were sufficiently resilient to cope with a considerable range of climatic

358 ANNALS OF THE SOUTH AFRICAN MUSEUM

conditions, including the first maximum of the Last Glacial. At present there
is no evidence to indicate that this latter event merited any major changes in
the stone industry. Detailed study may, however, yet reveal such a develop-
ment.

In the region of 40 000 to 30 000 B.P. the Middle Stone Age was replaced
by the Late Stone Age. The evidence from Boomplaas A suggests that the
transition to the first, as yet undescribed, L.S.A. industry took place before the
last glacial maximum. This is an apparent anomaly but it should be noted that
the micromammalian evidence indicates that conditions had been deteriorating
to a certain extent for some time before that change is recognizable in the
archaeological record. It may, therefore, be argued that the people had been
undergoing adaptation during the same period. If this were the case, it might be
merely accidental that the sum of the differences became sufficiently great to
warrant the recognition of a new industry before the contemporary environ-
mental trend had reached its climax. Perhaps an inherently more likely expla-
nation is that, in effect, cultural change is retroactive; that is to say, the sum of
the changes which are recognizable as a new industry represent the culmination
of adaptation to conditions which might already have begun to change again.
This might particularly be the case if the environmental changes constituted
repeated reversals of trends such as are envisaged for the Upper Pleistocene.
Thus in the case of Boomplaas A it may be that the early undescribed L.S.A.
industry represents the results of a period of adaptation to the interpleniglacial
conditions indicated by the micromammalian evidence. This may seem less
likely in view of the period of climatic deterioration prior to this. On the other
hand, conditions were apparently not very harsh and the period of time
involved may not have been very long (see Table 44). This hypothesis would
obviate the necessity of explaining why there were two different industries in
the same place both adapted to glacial maximum conditions, assuming a fairly
close association between industry and environment at this time.

The succeeding culture is the Robberg, which replaces the undescribed
industry at Boomplaas A at the height of the last glacial maximum. If this is
seen as a culmination of adaptation to environmental change, there is no
anomaly in the lack of coincidence between environmental and industrial
change. J. Deacon (1978: 109) has suggested that the change from one industry
to another will be marked by a period of accelerated change. There would
appear, however, to be no inherent reason why this need happen since, even
without a postulated increase in change, it is likely that the sum of differences
between industries will eventually be greater than the sum of similarities. In
fact, it would appear more logical to suggest that differences in rates of change
would be erratic responses to specific sets of conditions rather than part of a
fixed cycle. However, there is little merit in further speculation until more
information has been extracted from the material itself and it can be ascer-
tained that all stages in the development of the industry are represented in a
particular site. If such evidence is found, it is likely that detailed micromamma-

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 359

lian evidence for the contemporary environment would provide considerable
insight into the incentives for, and delays in, effecting industrial change.

According to the argument put forward above, under changing conditions
an industry could be expected to show its purest form and best adaptation to
prevailing conditions in its early stages. Subsequently, because the industry
might be supposed to become increasingly ill-adapted, changes would be
instituted that would ultimately require definition of another industry. It must,
incidentally, be pointed out at this stage that only high-level between-industry
change is being examined; low-level so-called stylistic change within one
industry is thought to represent response to quite different stimuli, perhaps
social or aesthetic (J. Deacon 1978). Although generalizations are inadvisable,
the evidence for the Robberg industry suggests that this scheme may have some
wider application beyond the Boomplaas A sequence. It has previously been
suggested that the Robberg industry represents an adaptation to glacial condi-
tions (Klein 1972a, 1978; J. Deacon 1978), which agrees with the suggestion
made above. It is to be noted, however, that at Boomplaas A the Robberg is
also temporally correlated with the late glacial, which was apparently a period
of considerable amelioration of conditions. It is logical to suggest, therefore,
that the Robberg must have become increasingly ill-adapted as conditions
changed. The evidence from Nelson Bay Cave is instructive in this respect.
Here the micromammalian evidence confirms that at least the earlier part of the
Robberg industry relates to glacial conditions. Unfortunately, there is no
micromammalian evidence for the later levels. However, the fact that J.
Deacon (1978) points out that the sample from level BSL, which she has placed
with the Robberg, is actually intermediate between the Robberg and the
succeeding Albany, is surely indicative of a situation similar to that postulated
for Boomplaas A.

The next major industrial change is recognized at about 12 000 B.P. in the
southern Cape at Nelson Bay Cave (J. Deacon 1978) and at Boomplaas A (H. J.
Deacon 1979). There is also a possibility of an industrial change at Bynes-
kranskop 1 at a similar time (Klein 1981) although Schweitzer & Wilson (1978)
point out that the paucity of remains in the lower levels makes comparison
difficult. At the first two sites, however, the industry has been identified as
Albany (J. Deacon 1978; Deacon et al. 1976). This industry is seen as an
adaptation to changing conditions and, as such, it is expected that it should
exhibit either a number of distinct sub-phases or a generality which would
ensure its suitability to a variety of conditions. This latter may be suggested by
the fact that at Nelson Bay Cave the Albany samples contain a smaller variety
of formal tools than do the Wilton or Robberg samples (J. Deacon 1978: 94). It
may, however, prove easier to establish the fine mechanics of adaptation at
Boomplaas A where the excavation of fine stratigraphic units should allow
analysis of discrete phases if such exist. It is interesting to note here that at
Boomplaas A the micromammalian evidence indicates fluctuations, albeit of
lesser amplitude, during the early Holocene at the time when the Albany

360 ANNALS OF THE SOUTH AFRICAN MUSEUM

industry was being employed. This may suggest a reason for the continued
usefulness of the industry after the beginning of the Holocene. It is also
possible that the unnamed pre-Wilton industry at Byneskranskop 1 will prove
to be a similar adaptation to somewhat fluctuating conditions. Since, however,
there is some possibility that conditions were changing more rapidly in the
Cango valley than at Byneskranskop 1 during this period, it may be that this
would have some effect on industrial development in the two areas. It may, of
course, prove impossible to separate the putative effects of this factor from
those of others, but it is still worth bearing in mind that it may exist.

The establishment of essentially modern, basically stable conditions in the
southern Cape may probably be correlated with the appearance of the Wilton
industry. It is suggested that the improved conditions will have provided a
greater range of possibilities from which the people might choose their modus
vivendi. At the same time it could be postulated that greater reliability of
resources would encourage specialization. H. J. Deacon (1972: 27) has criti-
cized Clark’s (1959: 189) characterization of the Wilton as a ‘period of regional
specialization par excellence’ as having ‘no basis in the understanding of the
effective environment in which the culture systems operated’. It may, however,
be that the fault les not in the concept but in the level of generalization. If
specialization is to be assessed, it should perhaps be examined at the level of
individual microenvironments rather than on a regional level. At the more
specific level there is much to recommend the hypothesis that the Wilton
culture involved considerable specialization. This process should probably be
seen in terms of multiple specialization by each group of people. The increasing
evidence for seasonal migration (H. J. Deacon 1969; Parkington 1972, 1976)
would suggest that this must have been the case. Klein (1974a: 275) has noted
that the coastal Wilton people exhibited an increasing tendency to exploit
marine resources and G. Avery (1974, 1976) has recorded concentration on
different species of shellfish by Pottery Wilton people. Inland there is evidence
for the exploitation of both selected plant foods and freshwater shellfish (H. J.
Deacon 1972). H. J. Deacon (1972: 34) also suggested that a low proportion of
segments and backed blades in the eastern Cape assemblages relative to those
from Zambia may indicate the use of traps to catch small bovids in the former
area, whereas projectile points were used to hunt larger bovids in the latter
area. Finally, H. J. Deacon (1972: 39) sums up the situation as an ‘apparent
trend towards an increasingly wider range of resources on an increasingly
intensive scale’.

The micromammalian evidence has nothing to add on any possible correla-
tion between culture and environment. There is insufficient evidence to allow
any spatial correlations because, as has already been mentioned, a large
number of observations would be necessary before this could be attempted.
However, such evidence as there is from Boomplaas A and Byneskranskop 1
provides some indication of the local environment of the site which could be
useful for checking possible specializations at each site. Changes within the

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 361

Holocene are hardly monitored by the micromammalian evidence and it is
suggested that changes must have been relatively minor. It will consequently be
necessary to acquire a more sensitive indicator for this period before it is
possible to determine whether or not there is any connection between environ-
mental change and industrial development. It is possible on logical grounds to
argue that the low-level changes witnessed in the Wilton (J. Deacon 1972) may,
in fact, represent adaptations to low-level environmental changes. It would be
interesting to find a means of assessing the more likely truth of the matter.

It is of considerable importance to point out, with H. J. Deacon (1972: 39),
that resource utilization cannot be seen simply as a response to environmental
conditions, or industrial development as direct cause and effect (J. Deacon
1978: 108). While it is most unlikely that this was the case at any time, there is
ample evidence from the Holocene for management of natural resources and
the environment. Evidence for the building of shelters and the use of fire, the
storage of food first in pits and then, presumably, in pots, and the later
introduction of sheep herding all point towards a greater independence from
environmental variables which, in turn, allows greater freedom of choice in any
given situation. It does, however, appear that the coincidence of a naturally
induced increase in options and an improved human technical skill could have
provided useful impetus for major cultural changes. That these changes
involved more than economic adaptation is suggested by the first appearance of
items of personal adornment and burials (Klein 1977) in Wilton contexts, which
presumably represent some sort of change in attitude towards the individual
and the group. Similarly, both Klein (1977) and H. J. Deacon (1972) remark
upon the evidence for changes in social structure and demography during the
later Holocene. All these aspects speak for the increasing cultural complexity
and, if one may so phrase it, civilization of the people responsible.

The available evidence suggests that, in so far as the lithic industry may be
taken as representative of a total culture, there may be some reason to suppose
that a correlation does exist between cultural and environmental change. This is
particularly the case with industries prior to the Wilton, which would be in
accordance with the suggestion that less technically advanced hunter-gatherer
societies will be closely dependent upon their environment. The evidence goes
on to indicate that the Wilton people displayed an increasing independence
from natural vicissitudes which may be expected to have as a corollary a
development increasingly independent of any direct environmental stimulus.
The converse is that later human activities almost certainly began to have a
significant effect upon the environment. The introduction of sheep herding to
the southern Cape at the end of the period under discussion in the study
(Schweitzer & Scott 1973; Deacon et al. 1978) must certainly have formed a
major step in this process, but it may well have begun before that time. For
instance, H. J. Deacon (1976: 174) has postulated that people may have
deliberately fired the vegetation to promote the growth of Watsonia sp., the
corms of which constituted an important source of food (H. J. Deacon 1976:

362 ANNALS OF THE SOUTH AFRICAN MUSEUM

162). This would undoubtedly have affected the vegetation, even if not to the
extent subsequently resulting from firing to improve grazing for domestic stock.
Klein (1972b) has suggested that improved hunting techniques may ultimately
have been responsible for the extinction of some bovid species at the end of the
Last Glacial. Although none of the evidence is adequate to prove any hypothe-
sis concerning the role of environment in cultural development, the data
discussed do suggest that the micromammalian evidence will prove useful to
any further work in this direction.

CONCLUSIONS

It was stated in the introduction that the principal aim of the present study
was to provide detailed information concerning the natural environment in
which past people existed. There is already considerable evidence to show that
micromammalian remains can be used to provide such information; it was not,
therefore, necessary to prove this point. There are, however, a number of ways
in which the basic material may be approached in order to extract environmen-
tal information and it was considered important to determine the potential of
each and the precise nature of the evidence provided. It was necessary, as a
preliminary stage in the interpretation of the data, to establish both the
approximate relationship of the available sample to the living community from
which it was derived and also the ecological significance of individual species.
Since micromammalian data provide one of several lines of evidence for
environmental change, this line must be fitted into, and checked against, the
general known scheme. This is particularly important because this evidence is
very local in application and does not necessarily come from uninterrupted
sequences.

There are considerable advantages to using micromammalian evidence for
the interpretation of the environment of prehistoric people. Not only is the
evidence contemporary but it also refers unquestionably to the neighbourhood
of the site from which the sample came. The evidence is, moreover, indepen-
dent of the archaeological evidence, which is extremely important, and it is also
capable of providing very detailed information concerning the environment at a
given time. Further, it has the potential, given the right controls, for indicating
both rates and amplitudes of environmental change through time. Finally, the
time lag in response to environmental change is almost certainly minimized by
the small size and consequent relative sensitivity and rapid breeding of micro-
mammals.

As far as the material itself is concerned, the initial good sample collected
by Tyto alba (barn owl) is thought not to suffer much differential destruction
because the specimens are of a similar size, and the fact that only the jaws were
counted provided each individual with an equal chance of being represented in
the sample. The relatively frequent occurrence of large samples greatly extends
the range of possible studies that may be conducted and allows reliable
statistical analyses to be performed.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 363

Disadvantages relate to the small size both of the area represented by the
evidence and of the animals themselves. In the first place the situation can be
rectified by the acquisition of greater numbers of samples. In the second case
the fact that the material may easily be broken or overlooked can affect the
completeness of the sample, but such biases can be reduced if great care is
taken in the collection of samples. Although large samples will allow studies of
population and community structure, they contain a degree of heterogeneity,
the elements of which it will not normally be possible to isolate. The level of
detail in interpretation will, therefore, be reduced, but this is an unavoidable
feature of fossil assemblages.

Of the three main lines of approach, one has been shown to relate
principally to vegetation changes and the others to climatic changes. The
former concerns changes in the composition of the community. Although it is
suggested that the immediate reason for changes in the proportions of some
species may be response to changes in other species, the ultimate cause of
changes is thought to have been shifts in the vegetational mosaic of the area in
question. Fluctuations in temperature and probably rainfall are reflected in
changes in the mean size of the individual in different populations. If the means
for different populations are compared, it is quite clear that some species
exhibit considerable variation in size both geographically and temporally. At
the community level there are also marked differences in structure which are
interpreted as reflecting changes in general climatic conditions at various times
in the past.

It is clear that there is considerable detail to be extracted from these
various lines of approach and the ultimate goal would seem to be a degree of
quantification of the data. At the present, however, there does not exist the
basic information relating to living representatives of the species involved.
When the data become available it should be possible to determine much more
precisely the vegetational shifts represented by the observed changes in compo-
sition of the small-mammal communities. As an aid to this, increased know-
ledge of interspecific behaviour should allow a better chance of isolating the
various mechanisms of change in community structure. Easier to collect will
probably be data concerning correlation of size variation and climatic factors.
Here again, though, it may be difficult to establish a direct link between, for
example, temperature and size, and even range and variability of temperature
could have different effects. There is, however, no doubt that considerable
advances must become possible once more detailed knowledge is gained of
existing small-mammal species.

Although the particular purpose of the present study was to furnish an aid
to the interpretation of human prehistory, it must be noted that the study of
micromammalian remains has considerable intrinsic interest. Apart from the
fact that the provision of evidence for palaeoenvironmental fluctuations and
conditions is important in its own right, the material also contains good
potential for the study of small mammals themselves. Just as information from

364 ANNALS OF THE SOUTH AFRICAN MUSEUM

modern representatives can aid interpretation of past data, so subfossil material
can provide insight into the biogeography, palaeoecology and general develop-
ment of extant species.

SUMMARY

Facts concerning the topography, climate and vegetation of the southern
Cape Province were given briefly as a background to the more detailed
descriptions of the individual sites which yielded micromammalian samples.
The material itself was discussed both in practical terms, from the point of view
of its collection and identification, and in more theoretical terms. These latter
concern not only how the material entered the cave, but also what biases may
have resulted from this method and subsequent taphonomic processes. Biases
inherent in micromammalian evidence were also discussed, the aim in general
being to establish what relationship any interpretation may bear to reality. The
basis which small mammals provide for interpretation was then examined. This
involved ascertaining as accurately as possible the habitat requirements of
individual species, after which each species was taken as representative of those
conditions. Control data from modern accumulations were then checked
against the known existing situation.

Existing evidence for palaeoenvironmental change is available from marine
and terrestrial geology and from the analysis of pollen and remains of animals
other than small mammals. These data were synthesized as a preliminary to
checking the interpretation from the small mammals and in order to provide a
framework into which the latter could be fitted. The micromammalian data
themselves were examined mainly from the point of view of community
composition. Changes in this provided good evidence of vegetational change
during the period under review. Changes in community structure, measured
with various indices of diversity, gave some indication of general relative
climatic conditions at different times. Changes in the mean size of the individ-
ual in various populations of selected species also suggested that there was
climatic change during the last 80 000 years. An overall reconstruction was
based on all these aspects and provided a picture which could be readily
correlated with other lines of evidence and appeared to be generally accurate as
well as more specific and detailed than most evidence.

The possibility exists of using the pattern of environmental change thus
acquired to correlate archaeological sequences which are undated or insecurely
dated. If the patterns of change at two sites can be matched, the likelihood
exists that they represent the same period of time, subject, of course, to gross
temporal controls.

The application of the evidence acquired from micromammals to archae-
ology could be fairly wide. Knowledge is gained of conditions in the immediate
vicinity of the occupation site. It was shown that, given sufficient material, it is
possible to provide detailed evidence of change which is tied securely to the

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 365

archaeological sequence but from which it is independent. It was postulated
that this could make it feasible to establish whether there exists any correlation
between environmental change and alterations in prehistoric technology and
economy.

ACKNOWLEDGEMENTS

The present study, which is an amended version of a thesis accepted for
the degree of Doctor of Philosophy at the University of Stellenbosch, resulted
directly from the advice and encouragement of Prof. R. G. Klein, University of
Chicago. The constructive criticism of Prof. H. J. Deacon, University of
Stellenbosch, and of Prof. Klein during their supervision of the research is
appreciated. Prof. Klein also generously made available various unpublished
data. Mr G. Avery is thanked sincerely for his help in all aspects of the work.
Thanks also go to Prof. Deacon for making available the Boomplaas A sample,
and to Mr F. R. Schweitzer and Dr Q. B. Hendey for allowing access to the
remaining samples which are lodged in the South African Museum. Modern
samples were collected by Miss K. Scott and Mr Avery; Mr T. N. Pocock
kindly provided a checklist of the Windheuvel material examined by him. Mr
T. P. Volman, University of Chicago, and Dr N. J. le Roux, University of
Stellenbosch, were of great help with the factor analysis but cannot be held
responsible for any errors. Thanks are also due to Dr G. de Graaff, National
Parks Board, for generously offering access to his unpublished manuscript, and
to those who kindly answered requests for information; individual items are
acknowledged as personal communications in the text. The typing of much of
the manuscript by Mrs M. Scheiner and Mrs P. Wallendorf is greatly appre-
ciated.

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6539 035

6. SYSTEMATIC papers must conform to the International code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific namé must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
comma separates author’s name and year
“semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.

In describing new species, One specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:

Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid- tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. ‘Smith, 15 January 1973.

Note standard form of writing South African Museum registration numbers and date.

7. SPECIAL HOUSE RULES

Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to in the text

~)

e.g. ©... the Figure depicting C. namacolus ...’; *. . . in C. namacolus (Fig. 10)...’

' (b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. DuToit but A.L.du Toit; Von Huene but F. von Huene

(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a
book or article, such as
‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.

Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

D. M. AVERY

MICROMAMMALS AS
PALAEOENVIRONMENTAL INDICATORS AND
AN INTERPRETATION OF THE LATE
QUATERNARY IN THE SOUTHERN CAPE
PROVINCE, SOUTH AFRICA

-

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