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ANNALS OF THE ANNALE VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

VOLUME 85 BAND 85

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

VOLUME 85 BAND

Hae SERUSTEES OF THE DIETIRUSTEES VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

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

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EIST OF TCONTENTS

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Avery, D. M.

Micromammals as palaeoenvironmental indicators and an interpretation of the Late

Quaternary in the southern Cape Province, South Africa. (Published January

BEI Me ey Re ere ete Ie ee ee ects Nese Mee SiMe Nee Seou pees SLs 183

DINGLE, R. V.

The Campanian and Maastrichtian Ostracoda of south-east Africa. (Published

“CUS NOLS ras EES) Na SRS Cae a a a he re Ye Se 1

NEW GENERIC NAMES PROPOSED IN THIS VOLUME

Dutoitella Dingle, 1981 ......

Klingerella Dingle, 1981 .....

Ponticulocythere Dingle, 1981

@) ejylei es! (si ce} je 4) [ele eine: ie) es tw) cee U6) is) .0)) a) (| elim) es) via) @) fellie) 'w! elie! ie}, (e! \eliculimiapine: e| (ee =! je) ieice) peice) ie

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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 moliusques. 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

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(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 85 Band

October 1981 Oktober

Part 1 Deel

THE CAMPANIAN AND MAASTRICHTIAN

OSTRACODA OF SOUTH-EAST AFRICA

R.V. DINGLE

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na gelang van die

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OUT OF PRINT/UIT DRUK

2 (135-8) S245 toe) 53, ss

6 tpi) 1s 8) OD: 7), 1043)

1102) 5; 7. tpi), 15(4-5), 24(02)..27, 31-3), 326), satan)

EDITOR/REDAKTRISE

Ione Rudner

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 019 2

Printed in South Africa by In Suid-Afrika gedruk deur

The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk.,

Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

THE CAMPANIAN AND MAASTRICHTIAN OSTRACODA OF SOUTH-

EAST AFRICA

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

LDL LOM. 5 oh eae Re Fee ee an ae ai, ee nee ane 2

Regional stratigraphy and sampling localities..................... 3

DE IETAE EG CES CTC TLC eee ose a 12

2 SPECIE GY bs SS eR Ta cee ee ea ec ce ee ee 131

EMAC OOPS ery hae at 8 A ee tek cue ee OS Rex 132

INGRUMEMI CaN ace ce ti aed ce a garg ee oe oe 132

BH-9 borehole and Zululand outcrops .............. 132

ROR a at ae esc het BS A = 146

Campanian—Maastrichtian palaeogeography of

southern Natal—Zululand Basin................. 149

JEVTISLEU CU) 211 E21 ie, 2 ek ee Gee ek ay OE er 152

NAO Re Pk Rin, Ae i tt 153

ING CUSHOAINP one fo Geter Ce Me en Wome ae Ee 154

WiteamId eee oe ee eee ge oes he Shee pee 156

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

mp ANNALS OF THE SOUTH AFRICAN MUSEUM

Southern area; Agulhas Bank. 4o002 . sees osteees Fe eee Fe 156

SUMMIT yee ere. ee Ss at ah oie hoe 159

POStratiaia playa: peri in ere Te SRN IR de ehh Ee 160

SOUR CASH AURIC A. 2092 ec ak oR Ses SO. aed gaa = Rk nama 160

Phylogeny—species distribution (appearances) ....... 160

Octracodwonal schome (50 2424 ac os 6 2a 163

Phyloveny—highertaxal 2.7: 2065.0"). 52. Seo eee 166

Phylogeny—species distribution (extinctions)......... 170

Comparison with other Gondwanide localities............ 170

Western Australias. 2.5 oiss4. 40s oo 6 OS a 172

Bast Aiea fet Fe ihe ae 8 cites 6 174

Wiest Alina <3 4 taro: Bane oan on oo 174

South AMECal ey seie ark Bote ee, oaks do £5

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

main

1 sediment

basins

- 2kmisobath

Fig. 1. Distribution of Campanian—Maastrichtian sediments in south-east Africa (shaded).

Numbers refer to sampling localities: 1—sample 818, Agulhas Bank; 2—lIgoda 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-1 zamba Igoda Camp Bank

Cytherella sp. . : : : : x x x x <

Cytherelloidea contorta . : , x x

C. umzambaensis : : : : x »<

C. griesbachi ‘ : : x<

C. mfoloziensis . : : : é x

Platella africana 2 : . x x

Bairdoppilata andersoni . ; : x x x x x x

B. andersoni aequalis : x

B. africana . 4 : : : ‘ x

B. cf. africana. ; : : : x

B. sp. A : : ; x

Bythocypris Polardsbavenss : : x x x Xx

IR SO, : . 4 . : x

Paracypris snmnzennlvocrats : x x x

P. zululandensis . ; x x Xx

P.sp. A : : : , : ; Xx

PrSps ’. 5 : : : x

Pontocyprella atbolnensis 6 x x

POSPil : f : : : Xx

Pariceratina ene jx

Eucytherura? pyramidatus <

Cytheropteron brenneri x

C. cf. westaustraliense x

Pedicythere fragilis x

Apateloschizocythere vaneleretilaen ; *

A. laminata : <

Amphicytherura qmneen : ‘ : x x

A. zululandensis : 4 : : x

A. armatus . : : ; : : x

A. sp. A : : : ; x

Klingerella aranearius , x

Hutsonia? sp. . : . : : x

Krithe nibelaensis : : : x x

K. sp. A ; ; : : : : <

Pondoina igodaensis . : : x

Xestoleberis luciaensis . : : x x x

Buntonia? sp... : : ; x

Brachycythere sicarius x *

B. longicaudata x x x x

Pterygocythere lencoolann” x

Ponticulocythere biremis x

Agulhasina quadrata . x

Unicapella sacsi ; ! x x x

U. reticulata : , ; ; x

Dutoitella dutoiti : : : : <

D. mimica . . : ie Xx x

Haughtonileberis hone inont x

H. vanhoepeni_. : 5 é x x

H. fissilis é : d x x x

H. nibelaensis . : ‘ ; ; x x

Rayneria nealei x

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 5

85(1) Zulu- Umz- Needs Agulhas

land BH-9 JC-1 zamba Igoda Camp Bank

Cythereis klingeri : ; : : x x

C. transkeiensis . : : ? x x

Oertliella sp. A. ‘ : : : x

O. pennata . : ' : ; : x

O. africana ‘ : : : : x x

O. maastrichtia . 4 : : x

Gibberleberis elongata : : : x

G.sp.A : ; : ; A : »

Trachyleberis minima : , x

T. zululandensis . : : : : x x

T. schizospinosa : ; : : x

Hermanites kennedyi : : : x x <

TIGKCUS =. : : ; : : x

WY ci.arcus A : : : . x x

Parvacythereis monziensis : : x

RPospinosa . : : : : : x<

Curfsina monziensis . : 5 : x

Cativella? dubia é x

Phacorhabdotus? anomala : : x x

eesp. cA... : : ; x

Australileberis stangerensis : : x

Paraplatycosta reticulata . ; Se

Indet. sp. | x

Iindet: sp. 2 x

Indet. sp. 3 x

Indet. sp. 4 x

Indet. sp. 5 x

Indet. sp. 6 x

Indet. sp. 7 ; : : : ; x

Indetesp. ~. : : : : 2 x

Totals . ‘ : i 44 28 11 5 15 6 15

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 Bronnimann, Globotruncana

arca (Cushman), G. stuarti (de Lapparent), and G. aegyptiaca Nakkady, which

indicate a Middle—Upper Maastrichtian age (Postuma 1971). This is corrobo-

ANNALS OF THE SOUTH AFRICAN MUSEUM

sisuapjaqiu si4aga]1uolysnv fy

SISUADIIN] S149Ga]O]SAX

SISUIPUD][NINZ S1dagajAYIVA J,

DUDIIA{D D]J21]149E

DJAOJUOD vaplojjadayIAD

SISUAPUB]NINZ vANAaYyAoIYdu py

DUNUNY S1Adga]AYIDA F

sisuadogspavy aid Ssiddkooylag

1uo0sAapuv vo] 1ddopslwg

‘ds pyjasaysad a

DIDPNDIBUO] AADYIAIAYIDAT

dds siuddavivg

1498U1] SIAAIYIAD

SHIADIIS AAAYIAIAY IDA

SIJISSY S1Aagapluojysnvoyy

SISUBDQUIDZLUN DapPlOpjasIYyAD

1uadaoyund Si4agajluojysnv fT

IYIDGSAIAS DAPIO[JadIYIAT

DIDBUOJA S14AGAAIGGID

Vv ‘dso

DJDUUad DI]21]149—C

1UOJYBhdY Sl4aqgajuo]ysno YY]

Dpiun} vDAnsaYy aolydup

JajDOU DIAQUADY

Jaquinu paq

Aj] eD0]

SUOISIAIP 988}S

818

MIII

IR OX KK

MIS Kk OK OX

oes XX SS xX

WS eK OK OX

MII

MN KK KM K XK

NS KO XX

MW KK XK XK

Ie IS OR KK

1/3

1/2

KK NM KX X

Dil.

iol

i TK AS OX

mS KK XK X

NN SM KK XK

CIV

XS XK XK X

CIil

MSM K KX X X

110

WOO AK XxX OX

K XX XX Ve aaa

OX DK Ok OK

BH-9 89.0

WTI OK OK OK OK

ISS PS GIES KOK COX

XO OX KIS PG EX eee wae

92.3

Ded

100.0

KOO NK KX K OX

KM NM KK KX KX X

x eM K KX KR Kear ae ee ae OX

106.0

OO) Bk XK

KX KX MK KX KKK KKK XK 2 aaa

SANTONIAN

CAMPANIAN AND MAASTRICHTIAN OSTRACODA

DIDUIUID] addy JAIOZIYISO]aJDdp

DIDINIIJAA DISOIA JD) ddA

pipaponb ouisvyjnsp

DIDUIOUD ¢ SNJOpqoYysoIDY

psoulds SladayJAIDAAD

psouldsoziyos sidaga]AYyov4 | *

DINU BjJA]110jNG

¢ ‘ds jopuy

I ‘ds Jopuy

149UUIAG UOAIIAdOAIYIAD

ISUAI|DAJSNDISAM "JD UOAIJAOABYIAD

p ‘ds Jopul

SNIAD ¢ SaJUDUAA HY]

DINSAIY DUIJDAIIIADY

SNUIMG AABYJAIO[NINJUOCT

SNIADIUDAD D]JAAIBUIL YY

SIIBDAL AABYIAIIPAT

DONG REAP ENLD)

7 ‘ds jopuy

DIDJOIIUD] AABYIAIOBA AA

SISUAIZUOU S1dAIYIAIDAAD

V ‘ds vansaydoiydup

snjopiuoddd ¢ vaAnsdaysaon”z

SISUAIZUOW DUIS{AND

OTe OU

DUDIIA{D D]jAID]

SISUIDJAGIU IYJIAY

sisuanjaqiu vjjasddojuog

sisualzojofiu vaployjas4ay aD

DIJYIIAJSDDUL D]JA1]J4IC

IUDJYIDJIU adaYyJAIOZIYIS OJ aJDdp

Vv ‘ds s14aqa]42qq19

2 satiate Consens

ae LEE NN SSS ISIS SS SY PPO TW Se

KK KR K K KK KK

Pr a Xx

x Fea

as &

x Erg

(oc Co

One

Xx x oMs

Ore

x Bey

x x ECs

oss

x x Bee

BOS

eX A ao°

x x fo ESa

4 aes

NS Khe ESS Beas

< mx aM

x x eens

3 Ze

SESS SOC as SS a

BS a

SoS oy OX < er

CNX

x << x popisuce

x ec

css

x sete

3:28

ros

NCSC NOX MS SX ex Ea:

x x aS) ee x ZESsa

: ee

Sa oS fae Ce a XC ROS SG

SO Se rh Ce be oe

SOG BREE X ES BK EK

» aay em taal OP a x SESE

CAMPANIAN AND MAASTRICHTIAN OSTRACODA

ANNALS OF THE SOUTH AFRICAN MUSEUM

DipUIuD] adaysAIOZIYISO]aIDd py

DIDININJAd DISOIAID]ADADT

vjvaponb ouispyjnsp

DIDUIOUD ¢ SNIOPGoYsOIDY

DSOUIS SIasIYIAIDAAD

DsoUuldsoziyas Sl4agajAyIod I”

DINU DIJAOING

€ ‘ds Japuy

I ‘ds Japuy

IMaUUasg UOsACOAIYIND

ISUAI]DAJSNDISAM “JO UOLIIAOLIYIAD

+ ‘ds japuy

SNIAD SAJIUDUAAFT

DINSALY DULIDADINADY

SIMA AADYIAIO/NINUOT

SMIADAUDAD D]JAAISUIL

SILSDAL ALBYIAIIPAT

DIGUP ¢ DIJAANDD

7 ‘ds Japuy

DIDJOIIUD] AMY IMIOBL AD

SISUAIZUOW SIAAIYIAIDAAD

V ‘ds pansaydorydiup

snjopnuvadd ¢ DANAAYIAONT

SISUAIZUOU DUIS{AND

Mloinp vpyjalioing

DUDIIA{D B]]AIP]

SISUADJAGIU AYNAY

SISUIDIIGIU D]JAAUAIOUOg

sisualzojofiu vaployjasaysi<D

DILYIIAISDDU DB] 21] 14IEQ

IUD] YOoJIU atayiAIOZIYISO]ajvdp

W “ds 4092/4999!

Smipulan nansauiciuidupw

SUSU APUDLINTZ SIMIGH/ AY POLL

DUBIA{D PYJAPIIO

DIAOJUOD vaPlO[JasayIAD

sisuapunjnynz bansayscony dup

DUNUILU SIAIGAAYIDAL

sIsuadDgSpABy IIA SIAUAIOYING x

luossapuv byojiddopalwg x

‘ds pyjasayia yd x

DIDPA_IBUO] AddYIAIAYIDAT

dds s1udtopavg

Masulyy Sla4ayl Dy

SNIADIIS AADYIAIAYIOAT

SIJISSY SlAagajiuojysandoyyT

sIsuapquinzuin vaplojjasaysid D

IMadaoyuva S14aqgajuojy sno pT

1yoDqsalas vaplojjasaylA Dy

DIDZUO]A $14aga]4agqIN

Vv ‘dso

piouuad vj] I1]1490

MO]YsnNDY Sl4aqgaplUojysno pT

ppiuin} vandoylcorydiup

la]pau blaaudoy

Jaquinu paq

AVRO}

SUOISIAIP a8e1S

< X X X

Se SPYERY

xX XX XK

x XX

x x

SCA

ea)

= =

X >

x xX

Xx x

x XX

x x

x xX

eos 52

x xX Xx

xxx

x Xx

x xX xX

x x X

ral

Cll

x x

110

GIS

= 2S ES

88.0

87.0

BH-9 89.0

82.0

cil

TABLE 2

92.3

97.5,

100.0

Distribution of ostracod species in samples from the Campanian and

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

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

102.2

106.0

110.0

SANTONIAN

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

EAST LONDON

my = NIBELA

OQ ST LUCIA

NEEDS lsLower Quarry

CAMP |

= \Upper Quarry

Igoda

outcrop

ened fe eiatatelstetatotabatetatercte aes main road

‘Nn

SIAN OCEAN

Fms

St Lucia

Mzinene

IEEE EEG

RIES SRS

QS 6 SSS 556

Makatini

+ Lebombo

UN RICHARDS volcanics

: J: t} BAY

HARBOUR Oo abies 20

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.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA

(9261) HOL nq “uonewso, preydty 104 uonses ody 7y BD sJoysi0q ‘yuRg seylnsy ‘(8/61) YOO] 2» JosUILy

‘duie} spoon pur eposy *(//61) Kpouuay 2 1osurpy ‘equiezwig ‘(p/61) WOT 7 OL nq ‘T-Of ‘(O861) FI3UIC ‘6-H ‘sone SuTMOTTOy oy) JOIZe ore

SUOI}DAS 19Y}O BY} JO sfle19q “(S/6]) J93UI[y 2 Apouusy Jo Aydeis}e13s pue UoT}e}OU dy) VSN puLN{NZ UI s1oqunuU poq puke AjITedO] ‘T sINBI 0} JoyoI

sloquinu Ay yeoo] UleY] ‘BOLPY 1sevo-YyINOsS UT soNIeso] UeNYoINseepy—ueiuedueD poduies wo sassouyoIy] sJoyoIog pue suOTdeS poInseay “¢ ‘S14

ajdwes >

NVINOLNVS

jyUusWasSeEg + 4 eI9N11S

E ueiuojuRs = equezwy

epob|

6!d>f_.,. + paeydiy

toeedig

Ebidmye ’*

es ST ea Ssuoljeuso4

a8 i +444 | ‘hits

NVINVdWVO

NVINVdWV9D

ie) 05 en mm CST

]

4 +

= a)

+ +

—

0061

HHH

I ,

tit

HHH

ec:

iss

Halal

Mitt

(aN

0081 00L

tll

is}

as)

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deo

OOLL "4

equiezwp

)

(eo)

NVILHOIYLSVVW

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 J 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 III. 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 Ill. 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, «

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

CAMPANIAN AND MAASTRICHTIAN OSTRACODA

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: ; (BaaRIOYyIAD % UBL) SISUADIIN] S14agajOJsaX

SNIDUAD “py

V dsp

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ppiwny bansay aoydiupy

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(BosRIOYyIAD % UPd) MasuI/¥ Sla4sdy aD

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Vv ‘ds ‘oO

pbyoUuuad Dj]a1]J49E

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1uUadaoyUva * LY

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ple ¢ [se Qi Ox Cx Ox pi C# (Bposev.}sO [e10} %, UBILU) SIsuadDgspaDYyaiA SIAdKIOYIAG

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q1qeIs 9[qeisun usdo —s UOT}RT- ~—- MOT] BYS

alo} is)

PdqdIISou

“purlninZ “6-H Pure RAGIN “IZOJOJJI ‘IZUOJ WO.y soIdods pure eidUdsd pe}99]9s JO UONNQIYsIp [e}UDWUOIIAUA

(so1j9lU) SyJdap 19}¥M po} eLUI}Sa

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.

ostr.

CYTHERELLA

CAMPANIAN

SANTONIAN

40 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, arid 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 (be)

Genus Cytherella Jones, 1849

Cytherella sp.

Fig. 5A—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.

(i111) 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 ‘7

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

locality

ostr.

assembl.

S CYTHERELLOIDEA PLATELLA

MAASTRICHTIAN

)

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

Rigs: 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 5a: 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

Fics 7€

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 II (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 ps) |

Fig. 8. Cytherelloidea mfoloziensis sp. nov. A. Holotype, SAM-—K5664, locality 20—7/1,

Mfolozi River, LV, Maastrichtian I]. 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 QF25

K5665 0,47 0,28

K5666 0,50 0,28

_K5667 0,50 0,28

K5668 0,45 O25

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 300.

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 O29

K5670 0,49 Ore

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 PoDOCOPINA 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).

CAMPANIAN AND MAASTRICHTIAN OSTRACODA

BAIRDIACEA

stage

locality

ostr.

assembl.

& BAIRDOPPILATA BYTHOCYPRIS

>20°7/o

Bythocypris

>80 Vo

Bdirdoppilata

7p)

c rn ————

5 © ——_

S eo =

o > 4

ce! © rH

{= 2

S mo}

fe)

Zz i

<q ,

IIISANTONIAN

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 III (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 = 300y.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 27

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 = 300y, others = 100u.

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

andersoni

e Ds:

andersoni aequalis

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-K5675, 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 Munster sp. var. aequalis var. nov. Chapman, 1916: 115, pl. XV, fig.

17a—b. (SAM-2736/20. )

Bairdia subdeltoidea Minster, Chapman, 1916: 114-115 (no ilustration). (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, 5a, 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 100u.

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 Inoceramus 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

PARACYPRIDIDAE

stage

locality

ostr.

assembl.

PARACYPRIS] PONTOCYPRELLA PARICERATINA

MAASTRICHTIAN

TOT

hirsuta

nibelaensis

—————

————

—T

—

————]

===

—————

———

—a

————J

——

——I

——J

—

—,

———s

——_

CAMPANIAN

SANTONIAN

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 IIJ (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 835m (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 100y.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA : SH,

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

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 100.

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-—KS5687, locality 113-3, Nibela

Peninsula, RV, Campanian IV.

Scale bar 90u.

Dimensions (mm)

length height

K5685 0,91 0,40

K5686 0,94 0,44

K5687 0,87 0,38

Age, distribution, palaeoecology

Campdnian IV to Maastrichtian [II (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 4]

Family Bythocytheridae Sars, 1926

Genus Pariceratina Griindel & Kozur, 1971

Pariceratina hirsuta sp. nov.

Fig. 19A—D

Derivation of name

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

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 O35

K5689 O55 0,30

K5690 0.55 Doi

Other material 0,60 2

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—K5689, 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 = 104, D = 30y, others = 100p.

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 Miller, 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.

Pig 19E

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 aS)

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) nein

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 = 30y.

Age, distribution, palaeoecology

Late Maastrichtian I to Maastrichtian II (Mfolozi River, Zululand). C.

brenneri occurs in ostracod assemblages Sa, 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 47

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 = 100w.

48 ANNALS OF THE SOUTH AFRICAN MUSEUM

Dimensions (mm)

length height width

(excl. spines) (over ala) (over ala)

K5694 0,46 0,27

K5695 0,45 E22

K5696 0,45 ?

K5697 0,45 0533

K5698 0,41 Cet

Other material 0,40 0,25

Age, distribution, palaeoecology

Maastrichtian I to Maastrichtian II (Mfolozi River, Zululand). P. fragilis is

found in ostracod assemblages 5a, 5b, and 7 (Fig. 22D), and although it

constitutes between 1 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. 3

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).

Jes

CAMPANIAN AND MAASTRICHTIAN OSTRACODA

CYTHERURIDAE SCHIZOCYTHERIDAE

3 CYTHEROPTERO APATELOSCHIZOCYTHERE|S

?2EUCYTHERURA PEDICYTHERE | AMPHICYTHERURA

locality

assembl.

MAASTRICHTIAN

brenneri

cf westaustraliense ie

CS]

pyramidatus

no)

armatus

CAMPANIAN

tumida

II|]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. 1. 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.

50 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

(2100-200 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. 238

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, 1.e. it inhabited outer-shelf and upper-

slope environments (Table 4).

CAMPANIAN AND MAASTRICHTIAN OSTRACODA a

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 110,0m, 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 = 100y.

a2 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, the

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

§ 3

Fig. 24. Amphicytherura armatus sp. nov. A. Holotype, SAM-K5701, locality 20-3, Mfolozi

River, LV, Maastrichtian I. B. SAM-—KS5702, 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 = 10yu, others = 100y.

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

GC, Poa

—

°" @

@ @

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 30y.

~—

Dimensions (mm)

length height

K5701 0,38 0522

K5702 0,40 05

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 55)

Description

External features. In lateral view, small quadrate species with rounded AM

and symmetrical, acuminate PM with a caudal process. DM and VM straight.

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

(2200 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

56 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 26. Apateloschizocythere. A-B. A. mclachlani sp. nov., holotype, SAM—KS705, 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 III. C. SAM-K5706, LV.

D. SAM-K5707, internal LV, E. SAM—K5706, sieve plates over normal pore canal apertures

LV. F. SAM-K5707, MS LV.

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

CAMPANIAN AND MAASTRICHTIAN OSTRACODA a7

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.

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

2 a

Sts -

cial 8) ie ae

n = Q& KLINGERELLA KRITHE XESTOLEBERIS ons

aranearius

: so

CAMPANIAN

III|ISANTONIAN

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-K5709, RV, locality 20-5, Mfolozi River, Maastrichtian I

SAM-KS5710, 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-K5710,

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 = 10u, 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.

-—

Fig. 29. Muscle scars, Klingerella aranearius gen. et sp. nov., SAM-K5710, locality 20-5,

Mfolozi River, LV, Maastrichtian I.

Scale bar 30u.

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 Q;52 O27

K5711 0,53 0,27

K5712 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 Kr6émmelbein

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 100y.

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-K5719, 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 = 100u.

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 055 0327

K5720 0,60 0,31

K5721 0,70 0,30

K5722 0,61 Q:34

K5723 0,70 O27

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 kostiacae 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-K5726, 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. luciaensis 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-K5727, locality 20-1/3, Mfolozi River, dorsal view RV(F) detail

PTE(G), Maastrichtian I.

Scale bars: B; G = 10u, C = 30p, others = 100u.

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 O25

K5725 0,40 O27

K5726 0,42 5. 0530

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 5a (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-

—

CAMPANIAN AND MAASTRICHTIAN OSTRACODA

BRACHYCYTHERIDAE

= PONTICULOCYTHERE

BRACHYCY THERE PTERYGOCYTHER

locality

assembl.

t=)]

ostr.

MAASTRICHTIAN

6 w

- ~~ -_

bY =

s &

3 gs 2

co -_

9 2

& c

p r=) S

= &

)

= ¢

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:

LIS),

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.

V2 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

Krommelbein, 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 (>S00 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 (&:

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 100u.

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-K5731, carapace, locality 20-1/2, Mfolozi River, Maastrichtian I

SAM-K5732, 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. Jongicaudata. 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 U5

K5732 0,75 0,45

K5733 0,70 0,40

K5734 0,68 0,38

CAMPANIAN AND MAASTRICHTIAN OSTRACODA WS)

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-K5730, 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 = 30, others = 100u.

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 Pferygocythereis 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 77

Holotype

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

Paratypes

SAM-K5736, LV, locality 20-8, Mfolozi River, Maastrichtian II

SAM-K5737, 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-—KS5735, 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 = 100p.

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 50y long) on its posterior

end.

80 ANNALS OF THE SOUTH AFRICAN MUSEUM

ai Phe

Se <6, SO re ee SS

= « 7

&& ee e

3 sae ee se

2 Ne ee eens needy Denti SEED SEN semen MONEE ORNs

Fig. 37. A-B. Agulhasina quadrata Dingle, 1971, TBD, 818 Alphard Formation, Agulhas

Bank, Maastrichtian III. A. SAM-K5738, LV. B. SAM—-K5739, RV. C-E. Unicapella sacsi

Dingle, 1980. C. SAM-—K5740, locality 20-7/1, Mfolozi River, LV, Maastrichtian II.

D. SAM-K5741, 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 = 10p, others = 100y.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 81

Age, distribution, palaeoecology

A. quadrata is known only from the Maastrichtian III (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 Grindel, 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 Idiocythere triebeli from the

Upper Campanian—Lower Maastrichtian of California may belong to Herrigocy-

there, as suggested by Griindel (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

UNICAPELLINAE

S& UNICAPELLA DUTOITELLA

stage

locality

ostr.

assembl.,

MAASTRICHTIAN

7 ;

-_ |

reticulata >

CAMPANIAN

INISANTONIAN

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 : é . Narrow large, prominent small, foveolate 0,55

dome- small rounded, to coarsely

shaped discon- reticulate

tinuous

Paleoabyssocythere 7 Massive) aa lance: 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 Griin-

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-—KS5611) is now considered to be an

artefact caused by slight distortion of the dorsal valve edge during sedimentary

compaction.

Age, distribution, palaeoecology

Campanian JI—Maastrichtian II (Mfolozi, Monzi, and Nibela outcrops,

Zululand and Richards Bay BH-9 borehole), Maastrichtian HI (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 II. 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 Rugen,

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 200.

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 Aflanticythere; 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

H.definita

H lO

(ar oety

)

Loe coe

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 posterior 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-K5743,

RV. E. SAM-K5744, internal LV. F. SAM—K5745, internal RV.

Scale bars all 100y.

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 dorsal 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 O57 0,33

K5743 0,55 032

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 II (Monzi, Mfolozi and Nibela areas

Zululand) (Fig. 383D—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 Aftlanticythere

maestrichtia Benson, 1977.

Holotype

SAM-K5747, LV, sample 818, Agulhas Bank, Maastrichtian III

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 = 100y.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA O91

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 = 30y.

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

KS5751 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.

92 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus C A.maestrichtia

' FP. OSG

5, ee

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—KS5742, locality

20-1/2, Mfolozi River, Maastrichtian I. B. Dutoitella mimica gen. et sp. nov., SAM—KS5747,

TBD 818 Alphard Formation Agulhas Bank, Maastrichtian III. 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

= B

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-K5749,

TBD 818 Alphard Formation, Agulhas Bank, LV, Maastrichtian III.

Scale bars 60.

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 I 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

Ma)

€

bs]

Be}

—

haughtoni

fissilis

vanhoepeni

CAMPANIAN

III|ISANTONIAN

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 II (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 III 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 nibelaensis 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-K5757, 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-KS5755, RV.

E. SAM-K5756, internal LV. F. SAM-—K5757, internal RV.

Scale bars all 100.

98 ANNALS OF THE SOUTH AFRICAN MUSEUM

Dimensions (mm)

length height width

K5754 0,65 0,31

K5755 0,58 0,30

K5756 0,62 0,34

K5757 0,67 0,32

K5758 0,62 0,25

Other material 0,65 2 OLS i

Other material 0,72 0,30

Other material 0,61 0,32

Other material 0,57 0,25

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 5a (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.

Ne)

CAMPANIAN AND MAASTRICHTIAN OSTRACODA

assembl.

stage

ole a essa locality

ostr.

OERTLIELLA

MAASTRICHTIAN

WI is S 15. 25° 35

c Ss

= — = =

® Q bod Oo

a n cs fe

il 3 sb

; sa

| : 4a

= A B Cc

= 5

WIISANTONIAN 3 3 5 615

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).

100 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 48. A-D. Oertliella. A. O. pennata Dingle, 1980, SAM-K5759, BH9 102,6 m, 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-K5762, locality 20—-1/3, Mfolozi River, LV, Maastrichtian I.

Scale bars all 100u.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA oi AOe

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 0233

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 5a, 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

Fig. 49. Oe6ertliella 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 = 10u, others = 100p.

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

O. 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 0,33

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 II (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

(?>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.

stage

locality

ostr.

assembl.

HERMANITES CYTHEREIS GIBBERLEBERIS

kennedyi

elongata

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. E. 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 100.

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 0529

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-—K5769) 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 0,68 035

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. Sir

Rayneria nealei Dingle, 1980: 55-57, figs 23E-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

Igoda (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 IJ 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-KS774, locality 110-14, Nibela Peninsula, RV, Campanian II.

Scale bars all 100u.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 111

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.sp.A

Fig. 53. Sketches of Gibberleberis species showing main rib patterns. A. G. elongata Dingle,

1980, SAM-—K5660, BH9 124,0 m, RV, Santonian III. B. G. sp. A, SAM-K5774, locality

110-14, Nibela Peninsula, RV, Campanian II.

Scale bars 200u.

G.elongata

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.

i2 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 100.

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 113

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 1 676 m), 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 (T. schizospinosa) constituted a remarkable 40 per

cent of the cytheracean population (33% of the total ostracod population).

114 ANNALS OF THE SOUTH AFRICAN MUSEUM

stage

locality

TRACHYLEBERIS PARVACYTHEREIS

schizospinosa

W—)

CAMPANIAN

minima

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 US

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). 7. 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 100u.

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. 1a.

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 _ a

e \

0,5 fo oY

/ 7

° a

= aes we

2 / on

2 | zululandensis CS."

—@ @@ bd Saas

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. T. zululandensis Dingle, 1980, SAM-K5779, locality 20-5,

Mfolozi River, RV, Maastrichtian I. C. Length v. height scattergram of adult 7. zululandensis

(dots) and 7. schizospinosa (circles). D. T. zululandensis Dingle, 1980, SAM-—K5780, locality

20-5, Mfolozi River, MS RV, Maastrichtian I.

Scale bars: A-B = 200u, D = 30p.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA +4 aly

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 Griindel, 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-KS5782, 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-—K5783, 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 = 100p.

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 0:22

K5782 0,45 O21

K5783 0,47 0,25

K5784 0,46 0,23

Other material 0,44 0,24

Other material 0,42 — 0:22

Other material 0,44 0,22

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 121

= a =

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

Be) Bee oe B

J ie J G4

monziensis spinosa

“2

(

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 30p.

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 1253

Paratypes

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

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

Fig. 61. Curfsina monziensis sp. nov., locality 21-1, Monzi, Campanian V. A. Holotype,

SAM-K5790, LV. B. SAM-K5791, RV. C. SAM-K5792, MS RV. D. SAM-K5792, internal

RV.

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

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 Sy 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 UY) 0,31

K5792 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

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 100yu.

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

= ¥f Costa

=} ®

2 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 0,53 0,30

K5794 0,50 0,26

Age, distribution, palaeoecology

Maastrichtian I to Maastrichtian II (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. 65A

Remarks

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

Oertliella.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 27

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-—K5797,

dorsal view RV, anterior to right. D. SAM—K5797, dorsal view PTE RV. E. SAM-K5796,

MS RV. i

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 100.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 129

Age, distribution, palaeoecology

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

Indet. 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. sp. 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 Igoda 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 1676 m). 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 Sd

(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 (4-5)

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 133

Maastrichtian Campanian Santonian

IV il

assemblage

population

= . . °

similarity Zo

7 trends

N° species als

diversity “

Bairdiacea

sé higher

taxa

B.richardsbayensis

405 B-andersoni ee Brachycythere spp. dominant

Lo Tar + EEEE == Haughtonileberis species

—_——"™, we bah oe" feet \ fu=—. spp.

oa can ae Mie ey a pas)

aN a al SE ill mee

borehole &

20 21 113 110 ha BH-9 locality

8 7-37-2,7-1,5 3 1-31-21-11 1311 3 1 0 211914 2 bed number

R.nealei

A.tumida

H. haughtoni

O.p

|S a aaahospeni +. — |

Seer [Fae] C. umzambaensis

T. minima __

al A.zululandensis

C.contorta

ae a] O. africana

een |e Sea: se alitandensis

____ Sn Be 2 eee) ee X.luciaensis

= H.nibelaensis

ile

U.sacsi arene

ee |) a armatus

O.maastrichtia

ee | |. C.mfoloziensis

ee |e | pnibelaensis

me ee L K-nibelaensis

mae sp Patricana

| «iD. dutoiti_|

C.monziensis

E? pyramidatus

P.monziensis

P.lanceolata

P. fragilis

K.aranearius

P.biremis

P. hirsuta

H? arcus

lLsp4

_C. cf westaustraliense

C.brenneri

— hs

l.sp3

fe eres a ea eee eae Sse

(a) fe

a!)

a no)

Fal

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 xX 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

ulomp “q uoimp “qd ujomnp ~“q

1SIDS “(Q 1SIDS “f) 1SIDS “(|

SNJDULAD * SNIDULAD “fy

DIDpnvI1BUo] “F DIDpNdIIBUO] “{ DIDpNVIIBUO]. JF

SNIADIIS “GF

SISSY “HT

1498U1]4 “D Masuyy Qo 1498U1] 4 “D

sisuavion] *“X SISUaDIIN] *X

DIJYIIAJSODU °C DIJYI1AJSDDU *C DIJYIIAJSODU °C DIJYIIAJSDDU °C)

4 iUadaoyuna *

sisuaDquivzuin * Dy sisuapquiozuin *+)

14IDqGSalss “D

DuvI1A{D °C DUuDIIA{D “CO

sisuapjaqiu °F] (ajdures [<)

4 SISUBPUDIN]NZ *p (45> Al[eroues)

ppiuni “pp ~ : 2 * IOUT

ne eee eee nee ee eee eee ene eee aan

sisuapjaqiu "yy ;

sisuanjaqiu “dq

DIJYIIAJSDDUL °C

sisuav1on] *X sisuav1on] “X SISUBDIIN] “X

1SIDS “()

1Apauuay “Hs 1dpauuay “Hs 1Apauuay “HT

SIISSY *H

14asUul]y *d 1498U1] “D

4 UadaoyUuna “FT (ajdures [<< ur)

SnuADoIS *g (%0I-s)

sisuakogspavyid “g sisuadogspanyrl4 “gq sisuakvgspavyold “gq * ites AleyI9 TL,

INSU

1u0sAapuD “g

sisuakDgspavyaid ‘gq sisuadDgspavyrid “g sIsuadvgspavYyd}4 *g

Ayjeoid A ASODS <i)

sisuavjaqiu *H

‘ds pyjasayid ‘ds pjjasayidD

uou juojysnoy *H (% 07-01)

DIDpNDIIBUO] “{ ~ S * Arepuosag

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1uossapuo ‘g ‘ds vjja4aysid "ds pjyjasayiady mMossapuv *g 1UOSAapuDd “g{ - (Y%OT<) Jueurmo0gd

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9 ATaV L

135

CAMPANIAN AND MAASTRICHTIAN OSTRACODA

] ‘ds ‘jopuy

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1dpauuay *H

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sIssy *“H

SIIBDAL “d

sisualzuoul “q

+DIQNP ¢ OD

SN1ADIUDAD “YY

+DINSATY “d

1dpauuay *H

sisuaizuoul ‘

‘Vv ‘ds 'y

SISUBDZWUIDGUIN *D

4DIDJOIIUD] “df

SISUAIZUOUL *

p ‘ds ‘yopuy

¢ ‘ds ‘yopu]

‘V ‘ds Vy

snuadlg “d

SNIADAUDAD *

1yaDqGsalds *D

1AOUUIAG *)

SIN SDAL “dd

SISUADQUIDZUIN *)

DJ DpnvI1B8Uuo] *F

pudd1d4f{0 °C

SNJDUIAD “py

sisualzojoful *)

sisuapuvjnjnz * I.

QSV[QUIDSSE UO O} POJOIA}SOI 4

smuaag ‘d

sisuavjaqiu *d

SNIADAUDAD “Y

1yo0qGSal4s “dD

HOINP

14auuadg “D

4DID]NINAA *f)

SIDA, “I

14a8ul]y “D

sisuapquivzuin *+)

sisuapjaqiu *¥

sisuaizojoful *D

sisuapudjnjnz *T

v ‘ds ‘oO

SOSE|QUIOSSE OM} O} POJOII}SOI y

SNIADIIS °F

DIDpnvdIIBUO] *F

pudI1A{D “d

‘Vv ‘ds 79)

pJDsBUuo]a *D

sisuanjaqiu *“Y¥

IUD] YODIJOU *Y

puvaidfo °C

sissy *H

SNJDULAD “Wf

sisuaizojoful *D

4 DIAOJU0D “2D yO4OJUOI *D

SISuapUuB]njNZ * Sisuapub]n]nz * (o]dues [)

4+ DUIUIM * WAse)

y a1e1

‘Vv ‘ds ‘O

a ee ee eee

TABLE 6

Composition of faunal assemblages 4-7 from Campanian-Maastrichtian of Zululand.

4 5

assemblage 4a 4b Sa 5b 6 7 2

Dominant (>20%) . B. andersoni B. andersoni Cytherella sp. Cytherella sp. B. andersoni

Cytherella sp.

Secondary . 3 . B. longicaudata

(10-20% H. haughtoni none

Cytherella sp. Cytherella sp.

H. nibelaensis

U. sacsi typically

B. richardsbayensis B. richardsbayensis B. richardsbayensis iS

B. andersoni eS

Tertiary S : . B. richardsbayensis B. richardsbayensis B. richardsbayensis 2

(5-10%) B. sicarius 9

(in >1 sample) H. vanhoepeni* 2

C. klingeri C. klingeri ical

H. fissilis iB

H. kennedyi H. kennedyi H. kennedyi iS

U. sacsi =|

X. luciaensis X. luciaensis X. luciaensis >

O. maastrichtia z

P. nibelaensis io

K. nibelaensis e

Minor... .. A. tumida =

(generally <5%) A, zululandensist a

(>1 sample) H. nibelaensis Ss

O. africana O. africana

C. griesbachi

C. umzambaensis — C. umzambaensis

H. vanhoepeni*

O. maastrichtia O. maastrichtia O. maastrichtia O. maastrichtia

X. luciaensis X. luciaensis

C. klingeri C. klingeri C. klingeri

H. fissilis

B. sicarius

B. longicaudata B. longicaudata B. longicaudata

A. armatus A. armatus

U. sacsi U. sacsé U. sacsé

D. duroitt D. duroit D. dusoitt

rg Seine: SITE AIOE A ai chanc ree

ps — 4 = TP ONOE > TS nyotozte: ‘ = =

~ zululandensis T. zululanden

P. lanceolata

P. hirsuta*

K. aranearius

C? dubia*

P. monziensis

P. fragilis

rare g 5 . . Ov. sp. A. O.sp.A

(<5% T. minimat

(1 sample) T. zululandensis T. zululandensis T. zululandensis T. zululandensis

C. contorta* C. contorta*

C. mfoloziensis C. mfoloziensis C. mfoloziensis fol

A, armatus A. armatus fe

H. fissilis H. fissilis 5

O. africana O. africana z

A. melachlanit+ >

K. nibelaensis K. nibelaensis 2

G. elongata z

G. sp. A. is)

P. africana =

B. longicaudata B. longicaudata z

B. sicarius B. sicarius 4

C. umzambaensis — C. umzambaensis C. umbamzaensis — C. umzambaensis 2

C. klingeri Q

P. fragilis P. fragilis S|

U. reticulata+ z

C. brenneri C. brenneri C. brenneri °

D. dutoiti 4

C. griesbachi C. griesbachi 2

K. aranearius K. aranearius 3

P. nibelaensis S

P. biremis P. biremis P. biremis >

A.sp. A. A. sp. A. A. sp. A.

Indet. sp. 3

Indet. sp. 4 6

P. monziensis P. monziensis

P. lanceolata*

H. kennedyi H. kennedyi

Hi? arcus*+ =

C. cf. west- ia)

p a

australiense

Indet. sp. 1

* restricted to two assemblages * restricted to one assemblage

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 5b 6 7 to

Trachyleberis minima ; Es one

Amphicytherura zululandensis

Apateloschizocythere mclachlani . a

Unicapella reticulata

Curfsina monziensis : : : *

Cativella ? dubia ‘ : ,

Pariceratina hirsuta -

Hermanites ? arcus

Cytherelloidea contorta . : : * % two

Haughtonileberis vanhoepeni . *

Pterygocythere lanceolata ? : * *

Cytherelloidea griesbachi ; * Es * three

Haughtonileberis nibelaensis . : * % %

Amphicytherura armatus : : # *

Klingerella aranearius

Cytheropteron brenneri :

Ponticulocythere biremis ; : %

Pedicythere fragilis .

Parvacythereis monziensis , ‘ * *

Ye KH

Se Se se Se Se

Dutoitella dutoiti . : : A % a x ke four

Platella africana 2 : : * % % *

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 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

ee ig /// ~«-Cytheracea

Cage 4 ag line

Sse 6 : 5a

Bythocypris” ~ ey,

line ke / 6

Gee : fe

( 1j@e« ° 2)

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 mportant 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

sacsit 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 merely 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 (5a), 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’ (Sa 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 5a),

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 (5S-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 poehele: 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

Pnibelaensis X.luciaensis B.richardsbayensis

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

B&C 50 C

e - H? arcus

H.kennedyi

turnover

O.maastrichtia D. dutoiti

B&C 50 26C 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 Sb 6 a

Oertliella africana : : 1 1 3

Trachyleberis zululandensis A i ; : 1 ] i 2 2 6

Apateloschizocythere mclachlani _ . 5 : 1

Unicapella sacsi : : ; , ; 4 2 3 4 6

Dutoitella dutoiti : : : i : : 1 B: 3 6

Pedicythere fragilis 1 1 2

Cytheropteron brenneri ] 1 1

C. cf. westaustraliense 1

Pariceratina hirsuta 3

Indet. sp. 2 1

No. of recordings per assemblage . 3 a d 12 9 26

No. of species per assemblage 2 4 6 6 3 8

No. samples per assemblage . : | 5 3 4 4 i)

No. records per sample per assemblage 0,4 1,4 2.3 3,0 233 ae0

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 5b 6 V

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:

204 : : ; . SB. andersoni B. andersoni B. andersoni B. andersoni Cytherella sp. B. andersoni

Cytherella sp. Cytherella sp. Cytherella sp.

specialized types . : : H. nibelaensis

T. minima A. laminata U. reticulata none C. monziensis C? dubia

A. zululandensis P. hirsuta

HZ. arcus

other characteristics:

Cytheracea é 3 . 50-60% 35-45% 30-50% 20-25% ~20% 20-30 %

Bairdiacea . : : . 15-407 30-40 % <40% >40% 30-40 % 40-50%

Cytherellidae ; : . 10-20% 10-25% 25-30% 25-35% 40-50% 20-30%

Bythocypris richardsbayensis <IKUYA <10% <10% usually >10% >13% SBA

Faunal diversity . 5 P <20% (low) >20% (high) ~20%(med.) <20% (low) <NOWA <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 Sa. 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 5b (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. Sa 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.

rata

Sill)

Allie oN : WG,

a ef Cee

BLIND SPECIES BH-9 AGES ZULULAND OUTCROP AGES

Bac ae C Bec a C Bac " Cc

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 Sa, 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 Sb)

during the period Santonian III to Campanian II (approximately 3 m.y.) can be

traced in BH-9 (Fig. 70B).

ostracod

assemblages

MAASTRICHTIAN

= “Bythocypris line’

)

CAMPANIAN

a inianen whats ies ieee NSS es nie aie =e =) me =e. @) = ae

Nee ee

lil SANTONIAN

eecoeoegGeeeeveeeeseeneee eee

eeeeRDPeeeeteoeeoeeeeseeseseee

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

Bythocypris ? sp.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 147

Krithe sp.

Cytherella sp.

. Bairdoppilata cf. africana

Australileberis stangerensis

Dutoitella mimica

Phacorhabdotus ? anomala

Pas Dw A

TABLE 10

Distribution of Ostracods and suggested palaeoenvironments in Campanian-

Maastrichtian section of JC—1 borehole.

Paracypris ? sp.

Indet sp. 7

Indet sp.

charophytes

x KK

Inoceramus prisms

No. ostracod valves

NN WwW

NWNOA LOO

—

(=|

cD)

He)

uo)

Za) {Su

?shallow-water,

?restricted environment,

essentially barren

rapid return to moderately

deep water

(?200 m)

shallow, major, fresh-

water influxes

moderately deep water

(?200 m) evidence of rapid

shallowing at top

depth gradually increasing

upwards to moderate-

water depths (?100-—200 m).

Establishment of sparse

ostracod fauna

?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 Bairdoppilatal

Cytherella fauna associated with abundant Inoceramus prisms

1 895-1 934 m (early Campanian)—barren of ostracods, abundant Inocer-

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 1 811-1 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

~ /

: |

5 NIBELA

ist

b) “vl

rugged hinterland

Tugela Delta

coast

eeeee Shelf edge

-- present coast

100 km

(ee |

™bo mts

SSS SS SS SSS SS SSS

SSS SSS SSS

SSSSSS

Pee BE

pee MONZI/MFOLOZI

> 6 @

SPS

Tugey,

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 Jnocer-

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 depth) versus time. This allows an assessment of

CAMPANIAN AND MAASTRICHTIAN OSTRACODA Si

rapid minor rapid rapid

regr. regr./transgr. transgr. transgr.

| | slow transgr.

————

water

depth ostr.

(00m) assembl.

6&7

—_——-| —_—_ —_ —

44 5b

ee ee aos ats, tae 4b—

?- -—Igoda 4a

?_ —~ Needs Camp

Hl I V lV Il Il

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 I/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 richardsbayensis 2 2 397,

Cypridacea : . Paracypris umzambaensis . 9 12

P.sp.A 5 2

Cytherellidae . . Cytherella sp. : 4 4 4%

Cytheracea ; . Cythereis transkeiensis a t

Hermanites kennedyi . 4 4

H ? cf: arcus 1 1

Pondoina igodaensis 14 [5

Hutsonia ? sp. 2 Z Siva

Buntonia? sp. . ee WR 2 2

Brachycythere longicaudata 14 15

Xestoleberis luciaensis 2 2

Indet sp. 5 Ps 2

Indet spp 6 6 J

Dominant (>20%) . : . Bairdoppilata spp.

Secondary (10-20%) : : . Paracypris spp.

Pondoina igodaensis

Brachycythere longicaudata

Tertiary (5-10%) : ; : . Cythereis transkeiensis

Cytheracea: total of 54 specimens = 57% fauna

Trachyleberididae : = 20%,

Brachycytheridae . i « Zon

Cytherideidae ; : S226 %

Progonocytheridae ; spend lA

Xestoleberididae . , ee we

Buntoniidae . : : RR i ay

Indet. é : : : a nOy,

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 £35

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 )

Cypridacea : . Pontocyprella sp. , ; : 2 4 f

Cytheracea 2 HH Pek, arcus : : : ; l 2 6

Indetsp.6 . A 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 Miinster . . 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? cf. 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 . . : : 5

Bairdoppilata cf. andersoni . : : E 2

Amphicytherura tumida A ; : : 3 2

Cythereis transkeiensis . : : . fragment

Sample from bed Pi 9

Brachycythere longicaudata . é : : ps

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-water

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 . 2 |

Cypridacea : . Pontocyprella nibelaensis . : 8 4

Paracypris sp. ; : ; : 2 1

Myiherciiidae . . -Platellaafricana. . . . 1 Fy 10

Ciiherculasep: “iidler som, Of 19 Oh

Cytheracea : ‘ Unicapella sacsi . 1] ]

Dutoitella mimica : 21 10

Phacorhabdotus ? anomala 1 l

Agulhasina quadrata 65 30

Paraplatycosta reticulata 9 4 83

Trachyleberis schizospinosa 73 33

Parvacythereis spinosa 7 3

Krithe nibelaensis ; 2 ]

Apateloschizocythere laminata . >) 2

Dominant (> 20%) : . Trachyleberis schizospinosa

Agulhasina quadrata

Secondary (10-20% ; : Dutoitella mimica

Tertiary 66-10%). ; ; ; 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.

%eo 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 IJ 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 1s

further borne out by the curve which plots percentage of species in a stage

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 161

re 4 .N

ee i \

80 0 ie atin d .

‘lee 208 .

! \ . “7. »

F / Fa |

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60 Ce © similarity

! .

40 A

‘ ‘ ge 8 _: population

sigs os 7) appearin

es SNS a cs Pon heads g

Mili Mil MI CV CIV Clll Cll 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. II Camp.I Camp. II Camp. III Camp.1¥V Camp. V Maas.I Maas. II Maas? III

no. species extant . : 21 24 24 17 DD, 24 SZ 29 15

mibtenitedi( 4) 9) . . ~ 14 (67) LIGA) 19 (79) 16 (94) 16 (73) 20 (83) 21 (66) 23 (93) 8 (53)

festuicted (7%) . . . 1 (5) 2 (8) 2 (8) 1 (6) ) 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 i) 8 (33) 1 (6) 2 (9) 3 (13) 5 (16) 21KG72) ?

Similarity

no. species ‘ : P 28 29 25 22 26 35 34 36

common species ‘ ‘ 17 19 16 16 20 21 27 8

% Similarity : y é 61 66 64 73 TA 60 79 22

no. species ; é > 45 60

common species : P 17 21

% similarity 3 , é Sant./Camp. similarity 37% Camp./ Maas. similarity 35%

no. Species extant . P 22: 41

inherited (%) . f : ? 21 (51)

restricted (%) . : 3 ? 20 (49)

appearances (% total) 20 (49)

disappearances (°% total) 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

nealet, 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 O6ertliella 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 II. 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

CAMPANIAN AND MAASTRICHTIAN OSTRACODA

167

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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

MaastrichtianI . .. : 23) Be 72

Campanian V ‘ ; : 1G) ee Wl

Campanian IV ; : : 14 DD 64

Campanian III : ; : 11 17 65

Campanian II : é : 16 24 67

Campanian I . : 16 24 67

Igoda : ‘ : ‘ : (2 17 vl

Mili Mii MI CV CIV Clll Cll Cl

Fig. 79. Number of extant species of Trachyleberididae as a percentage of total number of

cytheracean species in Campanian I—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 interestimg 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 . 3 eA ORO Pee OZ I 834

Cytheruridae . : : : i a at 2577 Slay

Schizocytheridae_ . : = ae oe, fo * 6% Se l4ee

Trachyleberididae : 5 MID KOS 7 44% Wn SAY

Cytherideidae gee Oey, OF 0 li haesie

Xestoleberididae oY OF 50 LS

Cytherettidae . ee egibcuust 7 0 ee O LP Gy/A Ne SA

Bythocytheridae . ‘ a @ i 260/ Se

Indet. . : ; OF 750 Oy, [ie Ses

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. WI Camp.I1V Camp. V Maas.I Maas. II Maas. III

Brachycytheridae ain oF NEG DNB DSi DSiVad 2) ile 8) BIZ PA MVIYA 0 O

Cytheruridae : = 050 0 O 0 O 0 O I YZ 3) BS D IY 0 O

Schizocytheridae SEA I IA 2 18% i PGA DMD, 2 OU A NZ i GZ

Trachyleberididae . 12 74% 13 82% 7 64% N64 O59 6 1043.7 8 42% Oni

Cytherideidae 0 O 0 O 0 O WIA i OY, V4 lh “S92 1 tiey,

Xestoleberididae 0 0 0 O 0 O I WZ I OZ il aiyZ hesie4 I

Cytherettidae . , 0 0 0 O 0 0 0 0 a) © ie ie i S% 0 0

Bythocytheridae 0 0 0 0 Onu OPO OPO l 4H, il 39% 0 0

Indet. . 0750 I) 0 0 0 0 0 0 lh 4% WP SA. TAO ao

16 16 11 14 17 23 19 5 ©)

c. Total cytheraceans: Zululand and Agulhas Bank

Campanian Maastrichtian

Total number of species , =» 30 29

New species appearing ; FO? NOs 7 14°. 48%

Total number of genera ; fia Wh 23

New genera appearing . : ; 1 MEGA A ATS,

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; Krommelbein 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

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ge chart for Campanian—Maastrichtian ostracods in Zululand (BH-9, and Monzi, Mfolozi and, Nibela outcrops) and

lon ran

. 80. Borehole explorat

‘igure 77 for explanation of left and right-hand columns.

Agulhas Bank. See fF

172 ANNALS OF THE SOUTH AFRICAN MUSEUM

W Africa Tanzania

C-9°/o, M-7°/o C-14°/o, M-17°/o

Td. EC

SE Africa

— Perth

-356 Carnarvon

9 s % 7

\ Xs

| ls Z A

Argentina W Australia

IM- 5°/.,mM-14°/o C-27°/o |

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,*Cytherelloideaand *Platella . 2?

Bairdiacea . tBairdoppilata and +Bythocypris : 9%

Cypridacea ; : . tParacypris and +Pontocyprella_ . ’ : Ay

Cytherideidae . Ln ethane Co 11a are : : : ak tee 50/

Eurotundracythere : : ; : Dee ce

Cytheruridae . : . *+Cytheropteron : 107%,

Oculocytheropteron : : <A

CORRCHAG Te ee i ee ed t2 7,

Paracytheridea : : : ; : 2 ce yA

*+Pedicythere . A : ea yA

Bythocytheridae ... B ythoceratina eae oe ne ee WA

Cytheralison . ; : 5 bay fe 3%

Monoceratina : : , : ; ‘ 1 ai

Progonocytheridae .. inclatargeyiiere jah ee ee 4% 4%

Schizocytheridae . . +Apateloschizocythere . : : 12% Wea

Pectocytheridae : Premunseyella ; : : : ay A Pa A

Trachyleberididae .. GON el Bah eee oer yee ee Te [hes

+Hermanites . k ; : 4 j : Ss

Karsteneis : : : : ; : Teg

Toolongella . ; 3 , : : : 6%

Anebocythereis teen! ; ag A yay A 2m

*+Curfsina : : ,; ; : t pega

Limburgina . , : ibe A

*+Oertliella : : 4%

+Trachyleberis : ; TA

Orthonotocythere . : : : : Ary,

Hystricocythere . ' Eas tr

Genus B : 2 ; . Le

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), O6ertliella 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 1)

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 i ‘ : %

*Trachyleberis :

*Genus C (Dutoitella)

*+Cythereis

+Phacorhabdotus

* Krithe

Genus B ; : : :

+ Bairdoppilata : : : ‘ *

*+Cytherella

Bs *Cytherelloidea

yw KH oe

% } X% y

Similarity with south-east Africa (*):

Campanian: 5 genera E. Afr., 24 genera SE Afr., 29 total, 4 common

Maastrichtian: 7 genera E. Afr., 29 genera SE Afr., 36 total, 6 common

14%

lia

lll

b. genera recorded in west Africa (after Apostolescu 1961, 1968)

(number of species in parenthesis)

Campanian Maastrichtian

#(1) *Brachycythere . ra : (3)

*(3) Ovocytheridea E J

*(6) Buntonia : : : : : *(4)

#(10) **Cythereis’ . ; : ‘ : (2)

(3) Cophinia : : ‘ : : #(1)

*(1) Nanocythere : , (1)

Dactylia : : : f : *1(1)

Protobasslerites . : : *(1)

Soudanella . : ; : #(1)

Isobuntonia . ; ; 5 #(1)

Anticythereis : 3 *(1)

=CT) ‘Clithrocytheridea’ : z

#(3) Veenia (Nigeria) : : : #(5)

(2) +Cytherella . : : : : *(2)

(1) Sphaeroleberis . , : (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 Jagiiel 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 Jagiiel Formation):

+Cytherella . ; , ; Gy

Jonesia ? ; : : ‘ 5 : : bey

Cytherura ; : : : : : : SA

Paracytheridea : : ; : iA

Trachyleberididae . . Alatacyihekeno- 3.75. See 8a}

*+Trachyleberis i ed LWA F

Piajacyihereis >" 5 ie Se ee 2 a 17

Wichmannella é : : : 4 : 297,

Similarity with south-east Africa (*):

8 genera Argentina, 29 genera SE Africa, total 37, 2 common = 5%

b. Middle Maastrichtian (Upper Jaguel Formation):

+Cytherella . : : , : : 4%

+Bythocypris ? ; : Ky

+Paracypris. 4 : : : ‘ AA

Trachyleberididae_ . ‘ Togoina : : ; : : : LA

+Trachyleberis ; : ; 5%,

Actinocythereis . : , : Is

Acanthocythereis . : : : : IT,

+Cythereis? : : ; GEA 48%

Henryhowella ; : : 2

Protocosta . ; p ; , , Bw

Veenia (Nigeria) . : : : : ; IS

Wichmannella : : ‘ é : 2 Leh

Hemicytheridae : ; Bradleya? ; . : ' : : ‘ A, 1°

Anticythereis eC Wig Ars 70

Pectocytheridae ; Munseyella . : : ys,

Progonocytheridae .. Mosaeleberis?: .°! . 3 = ee RAN

Tumidoleberis Jae sg Ree nn yA i ay,

Sphaeroleberis? . ; : : Wis

Cytheruridae . . . +Cytheropteron : ’ LA

Hemicytherura : ; ; : ; : bs } Dy,

Cytheromorpha? . : : ; A NOA

Schizocytheridae . +Amphicytherura? . ; : iy

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 TT

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.

REFERENCES

ALEXANDER, C. I. 1929. Ostracoda of the Cretaceous of North Texas. Bull. Univ. Texas Bur.

Econ. Geol. 2907: 1-137.

ALEXANDER, C. I. 1933. Shell structure of the ostracode genus Cytheropteron and fossil species

from the Cretaceous of Texas. J. Paleont. 7: 181-214.

APosTOLescu, V. 1961. Contribution a |’étude paléontologique (Ostracodes) et stratigraphique

des bassins crétacés et tertiares de l'Afrique occidentale. Revue Inst. fr. Petrole 16:

779-867.

178 ANNALS OF THE SOUTH AFRICAN MUSEUM

AposToLescu, V. 1963. Essai de zonation par les ostracodes dans le crétacé du bassin du

Sénégal. Revue Inst. fr. Petrole 18: 1675-1694.

Bairp, W. 1845. Arrangement of British Entomostraca, with a list of species, particularly

noticing those which have as yet been discovered within the bounds of the Club. Hist.

Berwicksh. Nat. Club 2: 145-148.

Bairp, W. 1850. The natural history of the British Entomostraca: 1-364, London: Ray Society.

Bate, R. H. 1972. Upper Cretaceous Ostracoda from the Carnarvon Basin, Western Australia.

Spec. Pap. Palaeont. 10: 1-85.

BaTE, R. H. & Bayuiss, D. D. 1969. An outline account of the Cretaceous and Tertiary

foraminifera and of the Cretaceous ostracods of Tanzania. Proceedings 3rd African

Micropaleontological Collogium 1968: 113-164, Cairo: National Information and Docu-

mentation Centre.

Bate, R. H. & Rosinson, E. eds. 1978. A stratigraphical index of British Ostracoda. Liverpool:

Seel House Press.

Benson, R. H. 1971. A new Cenozoic deep-sea genus, Abyssocythere (Crustacea: Ostracoda:

Trachyleberididae), with descriptions of five new species. Smithson. Contrib. Paleobiol. 7:

1-25.

Benson, R. H. 1977. The Cenozoic ostracode faunas of the Sao Paulo Plateau and the Rio

Grande Rise (DSDP Leg 39, Sites 356 and 357). Initial Reports of the Deep Sea Drilling

Project 39: 869-883. Washington: United States Government Printing Office.

BERTELS, A. 1973. Ostracodes of the type locality of the Lower Tertiary (lower Danian)

Rocanian Stage and Roca Formation of Argentina. Micropaleontology 19: 308-340.

BERTELS, A. 1974. Upper Cretaceous (lower Maastrichtian?) ostracodes from Argentina.

Micropaleontology 20: 385-397.

BERTELS, A. 1975. Upper Cretaceous (middle Maastrichtian) ostracodes of Argentina. Micro-

paleontology 21: 97-130.

BERTELS, A. 1977. Cretaceous ostracoda—South Atlantic. In: Swain, F. M. ed. Stratigraphic

micropaleontology of Atlantic Basin and Borderlands: 271-304. Amsterdam: Elsevier.

BosquEt, J. 1854. Monographie des crustacés fossiles du terrain Crétacé du Duche Limbourg.

Mémoires de la Commission pour la description de la Carte geologique de la Neerlande 2:

53-126.

Brapy, G. S. 1880. Report on ostracoda dredged by H.M.S. ‘Challenger’ during the years

1873-76. Rep. scient. Results Voy. Challenger (Zool.) 1(3): 1-184.

Brapy, G. S. 1898. On new or imperfectly-known species of ostracoda, chiefly from New

Zealand. Trans. zool. Soc. Lond. 14: 429-452.

Brapy, G. S., CrosskEy, H. W. & Rosertson, D. 1874. A monograph of the Post-Tertiary

Entomostraca of Scotland including species from England and Ireland. London: Palaeon-

tographical Society.

BRENNER, P. & OeErTLI, H. J. 1976. Lower Cretaceous ostracodes (Valanginian to Hauterivian)

from the Sundays River Formation, Algoa Basin, South Africa. Bull. Cent. Rech. Pau 10:

471-533.

But er, E. A. & Jones, D. E. 1957. Cretaceous ostracoda of Prothro and Rayburn salt domes

Bienville Parish, Louisiana. Bull. geol. Surv. La. 32: 1-65.

CHAPMAN, F. 1898. On ostracoda from the ‘Cambridge Greensand’. Ann. Mag. nat. Hist. 3:

331-346.

CHAPMAN, F. 1904. Foraminifera and ostracoda from the Cretaceous of East Pondaland, South

Africa. Ann. S. Afr. Mus. 4: 221-237.

CHAPMAN, F. 1916. Foraminifera and Ostracoda from the Upper Cretaceous of Need’s Camp,

Buffalo River, Cape Province. Ann. S. Afr. Mus. 12: 107-118.

CHAPMAN, F. 1923. On some foraminifera and ostracoda from the Cretaceous of the Umzamba

River, Pondoland. Trans. geol. Soc. S. Afr. 26: 107-118.

CoryELL, H. N. & FieLtps, S. 1937. A Gatun ostracode fauna from Cativa, Panama. Am. Mus.

Novit. 956: 1-18.

CorvELL, H. N., Sample, C. H. & JENNINGS, P. H. 1935. Bairdoppilata, a new genus of

ostracoda, with two new species. Am. Mus. Novit. 777: 1-5.

Damortte, R. 1974. Un noveau genre d’ostracode de |’Albien du Bassin de Paris: Matronella

n.g. Importance des empreintes musculaires dans les attributions génériques chez les

Trachyleberidinae. Paleontographica 146: 181-189.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 179

Deroo, G. 1966. Cytheracea (Ostracodes) du Maastrichtian de Maastricht (Pays-Bas) et des

regions voisines; résultats stratigraphiques et paléontologiques de leur étude. Meded. geol.

Sticht. S(C) 2: 1-197.

DINGLE, R. V. 1969. Upper Senonian ostracods from the coast of Pondoland, South Africa.

Trans. R. Soc. S. Afr. 38: 347-385.

DINGLE, R. V. 1971. Some Cretaceous ostracodal assemblages from the Agulhas Bank (South

African continental margin). Trans. R. Soc. S. Afr. 39: 393-418.

DINGLE, R. V. 1973. Post-Palaeozoic stratigraphy of the eastern Agulhas Bank, South African

continental margin. Mar. Geol. 15: 1-23.

DINGLE, R. V. 1976. Palaeogene ostracods from the continental shelf off Natal, South Africa.

Trans. R. Soc. S. Afr. 42: 35-79.

DINGLE, R. V. 1978. South Africa. In: MOULLADE, M. & Nairn, A. E. M. eds. The

Phanerozoic Geology of the World II, The Mesozoic A: 401-434. Amsterdam: Elsevier.

DINGLE, R. V. 1980. Marine Santonian and Campanian ostracods from a borehole at Richards

Bay, Zululand. Ann. S. Afr. Mus. 82: 1-70.

Ducasse, O. & GrekorF, N. 1976.-Quelques ostracodes de l’éocene inférieur du sud-ouest de

Vocéan Indien: Site 246, Croisiere 25, ‘Glomar Challenger’, Deep Sea Drilling Project.

Revue Micropaléont. 19: 134-152.

Du Torr, S. R. 1976. Mesozoic geology of the Agulhas Bank, South Africa. Unpublished Ph.D.

thesis, University of Cape Town.

Du Torr, S. R. & Leitn, M. J. 1974. The J(c)-1 borehole on the continental shelf near Stanger,

Natal. Trans. geol. Soc. S. Afr. 77: 247-252.

Eacar, S. H. 1965. Ostracoda of the London Clay (Ypresian) in the London Basin; I. Reading

District. Revue Micropaléont. 8: 15-32.

FirsTBROooK, P. L., FUNNELL, B. M., Hurtey, A. M. & Smitu, A. G. (n.d.). Paleoceanic

reconstructions 160-OMA. Washington: National Science Foundation.

GrosDIDIER, E. 1979. Principaux ostracodes marins de l’intervalle Aptien-Turonien du Gabon

(Afrique occidentale). Bull. Cent. Rech. Expl. Prod. Elf-Aquitaine 3: 1-35.

GRUNDEL, J. 1973. Zur Entwicklung der Trachyleberididae (ostracoda) in der Unterkreide und

in der tieferen Oberkreide. Teil I: Taxonomie. Z. Geol. Wiss. Berlin 1, 11: 1463-1474.

GrRUNDEL, J. 1974. Zur Entwicklung der Trachyleberididae (ostracoda) in der Unterkreide und

in der tieferen Oberkreide. Teil II: Phylogenie. Z. Geol. Wiss. Berlin 2, 1: 61-71.

GRUNDEL, J. & Kozur, H. 1971. Zur Taxonomie der Bythocytheridae und Tricorninidae

(Podocopida, Ostracoda). Monatsber. dt. Akad. Wiss. Berlin 13: 907-937.

Herric, E. 1965. Zwei neue I[diocythere-Arten aus dem Campan, Insel Riigen (Ostsee).

Geologie 14: 1224-1235.

Hix, B. L. 1954. Reclassification of winged Cythereis and winged Brachycythere. J. Paleont.

28: 804-826.

HInTE, J. E. vAN.1976. A Cretaceous time scale. Bull. Am. Ass. Petrol. Geol. 60: 269-287.

. Hoven, J. C. 1964. Upper Cretaceous ostracods from California. Palaeontology 7: 393-429.

Howe, H. V. & CHamsers, J. 1935. Louisiana Jackson Eocene ostracoda. Bull. geol. Surv. La

5: 1-65.

Howe, H. V. & Laugrencicn, L. 1958. Introduction to the study of Cretaceous Ostracoda. Baton

Rouge: Louisiana State University Press.

Jones, T. R. 1849. A monograph of the Entomostraca of the Cretaceous Formations of England.

London: Palaeontological Society.

Jones, T. R. 1884. Notes on the Foraminifera and Ostracoda from the deep boring at

Richmond. Q. JI geoi. Soc. Lond. 40: 766-777.

Jones, T. R. & HINDE, G. J. 1890. A supplementary monograph of the Cretaceous Entomostra-

ca of England and Ireland. London: Palaeontographical Society.

KENNEDY, W. J. & KLINGER, H. C. 1975. Cretaceous faunas from Zululand and Natal, South

Africa. Introduction, stratigraphy. Bull. Br. Mus. nat. Hist. (Geol.) 25: 263-315.

KLINGER, H. C., KAUFFMAN, E. G. & KENNEDY, W. J. 1980. Upper Cretaceous ammonites and

inoceramids from the offshore Alphard Group of South Africa. Ann. S. Afr. Mus. 82:

293-320.

KLINGER, H. C. & KENNEDY, W. J. 1977. Upper Cretaceous ammonites from a borehole near

Richards Bay, South Africa. Ann. S. Afr. Mus. 72: 69-107.

180 ANNALS OF THE SOUTH AFRICAN MUSEUM

KLINGER, H. C. & KENNEDY, W. J. 1980. The Umzamba Formation at its type section

Umzamba Estuary (Pondoland, Transkei), the ammonite content and palaeogeographical

distribution. Ann. S. Afr. Mus. 81: 207-222.

Kuincer, H. C. & Lock, B. E. 1978: Upper Cretaceous sediments from the Igoda River

Mouth, East London, South Africa. Ann. S. Afr. Mus. 77: 71-83.

KROMMELBEIN, K. 1964. Ostracoden aus der marinen ‘Kiisten-Kreide’ Brasiliens. 1: Brachycy-

there (Brachycythere) sapucariensis n. sp. aus dem Turonium. Senckenberg. Leth. 45:

489-495.

KROMMELBEIN, K. 1972. Remarks on marine Cretaceous ostracodes-of Gondwanic distribution.

Proc. 5th African Colloquium on Micropalaeontology 1972: 539-551, Madrid: Revista

Espanola de Micropaleontologia.

LATREILLE, P. A. 1806. Genera Crustaceorum et Insectorum 1: 1-303. Paris.

Lusimova, P. S. 1955. Ostracoda of the Middle Mesozoic formations of the central Volga area

and the Obshehego Sirta. In: LuBimova, P. S. & CHABAROVA, T. H. eds. Ostracoda of the

Mesozoic sediments of the Volga—Urals region. Trans. All-Union Petrol. Sci. Res. Geol.

Explor. Inst. (V.N.I.G.R.I.) (N.S.) 84: 1-189. (In Russian.)

Makripes, M. 1979. Foraminifera of the Upper Cretaceous Mzamba Formation, Transkei,

Southern Africa. Unpublished M.Sc. Thesis, University of Witwatersrand.

MaNDELSsTAM, M. I. 1960. Arthropods. In: Ortov, Y. A. ed. Elements of Palaeontology. 8:

1-515. Moscow: Geological and subsurface prospecting: Scientific and Technical Edition.

(In Russian. )

Maup, R. & Orr, W. 1975. Aspects of post-Karroo geology in the Richards Bay area. Trans.

geol. Soc. S. Afr. 78: 101-109.

McGowran, B. & Moore, A. C. 1971. A reptilian tooth and Upper Cretaceous microfossils

from the Lower Quarry at Needs Camp, South Africa. Trans. geol. Soc. S. Afr. 74:

103-105.

McKENZIE, K. G. 1967. Recent ostracoda from Port Phillip Bay, Victoria. Proc. R. Soc. Vict.

80: 61-106.

McLacuian, I. R. & McMILLAN, I. K. 1979. Microfaunal biostratigraphy, chronostratigraphy

and history of Mesozoic and Cenozoic deposits on the coastal margin of South Africa.

Spec. Publs geol. Soc. S. Afr. 6: 161-181.

Moore, R. C. ed. 1961. Treatise on Invertebrate Paleontology. Part Q, Arthropoda 3.

Lawrence: University of Kansas Press.

MorkHoven, F. P. C. M. van. 1962. Post-Palaeozoic Ostracoda. Their Morphology, Tax-

onomy, and Economic Use. 1. General. Amsterdam: Elsevier.

MorkHOoVEN, F. P. C. M. van. 1963. Post-Palaeozoic Ostracoda. Their morphology, taxonomy,

and economic use. 2. Generic descriptions. Amsterdam: Elsevier.

MUELLER, O. F. 1785. Entomostraca seu insecta testacea, quae in aquis daniae et norvegiae

reperit, descripsit et iconubus illustravit. Lipsiae et Havniae.

MULLER, G. W. 1894. Die Ostracoden des Golfes von Neapel und der angrenzenden Meere-

sabschnitte. Fauna Flora Golf. Neapel 31: 1-404.

Musaccuio, E. A. 1979. Datos paleobiogeograficos de algunas asociaciones de foraminiferos,

ostracodos y carofitas del Jurasico medio y el Cretacico inferior de Argentina. Ameghi-

niana 16: 247-271.

NEALE, J. W. 1975. The ostracod fauna from the Santonian Chalk (Upper Cretaceous) of

Gingin, Western Australia. Spec. Pap. Palaeont. 16: 1-81.

NEUFVILLE, E. M. H. 1979. Upper Cretaceous-Paleogene marine ostracods from the Sergipe-

Alagoas Basin, north-eastern Brazil. Bull. geol. Instn Univ. Uppsala 8: 135-162.

OERTLI, H. J. 1963. Mesozoic ostracod faunas of France, Leiden: Brill.

Pokorny, V. 1964. Oertliella and Spinicythereis, new ostracod genera from the Upper Cre-

taceous. Vest. ustred. Ust. geol. 39: 283-284.

Pokorny, V. 1967. The genus Curfsina (Ostracoda, Crustacea) from the Upper Cretaceous of

Bohemia, Czeckoslovakia. Acta Univ. Carol. Geologica 4: 345-364.

Postuma, J. A. 1971. Manual of Planktonic Foraminifera. Amsterdam: Elsevier.

Puri, H. S. 1954. Contribution to the study of the Miocene of the Florida Panhandle. Part III

(Ostracoda). Bull. geol. Surv. Fla 3: 217-309.

Puri, H. S. 1955. Hermanites, a new name for Hermania Puri 1954. J.Paleont. 29: 558.

Reuss, A. E. 1846. Die Versteinerungen der b6hmischen Kreide Formation 2: 59-148. Stuttgart;

Schweizerbart.

CAMPANIAN AND MAASTRICHTIAN OSTRACODA 181

Ramsay, W. V. 1968. A new morphological aspect of the ostracode genus Cytherelloidea

Alexander. Micropaleontology 14: 348-356.

REYMENT, R. A. 1960. Studies on Nigerian Upper Cretaceous and Lower Tertiary Ostracoda.

Part 1: Senonian and Maastrichtian Ostracoda. Stockh. Contr. Geol. 8 1-238.

ROSENFELD, A. & BEIN, A. 1978. A preliminary note on recent ostracodes from shelf to rise

sediments off north-west Africa. Meteor ForschErgebn. 29: 14-20.

Sars, G. O. 1866. Oversigt af Norges marine ostracoden. Forh. VidenskSelsk. Krist. 7: 1-130.

Sars, G. O. 1888. Nye Bidrag til Kundskaken om Midlehavets Invertebrat fauna 4. Ostracoda

Mediterranea. Arch. Math. Naturv. 12: 173-324.

Sars, G. O. 1922-1928. An account of the Crustacea of Norway 9. Ostracoda, Parts 1-16:

1-277. Bergen: Bergen Museum.

StESSER, W. G. & Mites, G. A. 1979. Calcareous nannofossils and planktic foraminifers in

Tertiary limestones, Natal and eastern Cape, South Africa. Ann. S. Afr. Mus. 79: 139-158.

Simon, W. & BARTENSTEIN, H. 1962. Leitfossilien der mikropaldontologie. Berlin-Nikolassee:

Gebriider Borntrager.

Soekor, 1976. Structure of the Mesozoic succession of the Agulhas Bank. Johannesburg:

Technical Department, Southern Oil Exploration Corporation.

SYLVESTER-BRADLEY, P. C. 1948. The ostracode genus Cythereis. J.Paleont. 22: 792-797.

SYLVESTER-BRADLEY, P. C. & BENSON, R. H. 1971. Terminology for surface features in ornate

ostracods. Lethaia 4: 249-286.

Swaln, F. M. 1946. Upper Jurassic Ostracoda from the Cotton Valley Group in northern

Louisiana; the genus Hutsonia. J.Paleont. 20: 119-129.

TRIEBEL, E. 1941. Zur Morphologie und Oekologie des Fossilen Ostracoden. Mit Beschreibung

einiger neuer Gatungen und Arten. Senckenberg. Leth. 23: 294-400.

TRIEBEL, E. 1952. Ostracoden der Gattung Cytheretta aus dem Tertiar des Mainzer Beckens.

Notizbl. hess. Landesamt. Bodenforsch. Wiesbaden 6: 15-30.

WHATLEY, R. C. & STEPHENS, J. M. 1976. The Mesozoic explosion of the Cytheracea. ADI.

Verh. naturw. Ver. Hamburg 18/19: 63-76.

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Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15SA

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Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

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R. V. DINGLE

THE CAMPANIAN AND MAASTRICHTIAN

OSTRACODA OF SOUTH-EAST AFRICA

VOLUME 85 PART 2

ay GS

OF THE SOUTH AFRICAN |

~” MUSEUM

CAPE TOWN.

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

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

FiscHER, P.-H., DuvAL, M. & RAFFy, 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

rant 2, Deel

MICROMAMMALS AS

PALAEOENVIRONMENTAL INDICATORS AND

AN INTERPRETATION OF THE LATE

QUATERNARY IN THE SOUTHERN CAPE

PROVINCE, SOUTH AFRICA

DMA 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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word uitgegee in dele op ongereelde tye na gelang van die

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Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 8000

OUT OF PRINT/UIT DRUK

(2623, 528), (1224-5, 8 Caps) 0 3 roe

GC, t=pib)y 14). 8. 922-7) 103)

11(1-2, 5, 7, t=prit); 154=5), 24(2),27, 31(1-3), 32), 232450)

EDITOR/REDAKTRISE

Ione Rudner

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 028 1

Printed in South Africa by In Suid-Afrika gedruk deur

The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk.,

Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS

AND AN INTERPRETATION OF THE LATE QUATERNARY

IN THE SOUTHERN CAPE PROVINCE, SOUTH AFRICA

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

WETPOGU EEO Ay Re ae Se ae ee ee See ae 184

TEE STUDS Coy ore ee 37 ile See ae oe ee en ee a Se 186

“WEE S0IEY . . ogg ates Sie ae endl ee een? Shh Beate 189

Pee AO PaCAISHIGS Siar. weet ho U, Slus Meee iS eon tes oe 189

Pe LDDETE SSS Se ee tte ee cee ee ee re 200

LE EDIE og et RS a ee seaport Se om nee Per 207

Pe HEVAOn ACCUMMMAUOM 204 oe tae Seca wee es Snes FFs ee eet 207

IMUSIC OUCCHIOUE 16 Fs chen dyed as wisleiamebeeuels che else's DOs Ob ae pee 209

LETTE BETS agi se ge 2 OREO ee ee 210

Mie aan OF CxcaVvale@ Samples 2. 8). Ge ee eee eee ee ee 213

Whe nature Of micromammalian-evidence....5..5.2.56.020. 0. eee tee 217

PE LOOGIS ovo GUIS ole Sy a ane BP ge ee A 228

Mam uiatORtOl MUBUTUI NUMBERS . 5.2. sn 4 oes ness ie ce eee ee ee 228

Multivariate statistical analysis of primary data....................... 229

EMOECSSES MOM MeEpPHetNe ANALYSES 62 5.57.0... bases vee ee oe ee 234

hensuration. and -caleulation-of MGICES\): 2.5.25 ..666 . feeb es lee Bob ees 235

DEE THOSE CSET so Soe cae CRUG es ie Meet eae eh ea on rene ee 238

CMO ONG CISERMON fee, Ge Le ek eae ee ete eee eels Oe 238

Taasis On MilephetaniOn : 1k fee ess oo Soe eee oes does oon 243

Farcepictalion of conmarative Samples) 22 2c jn. 2 as 2 Pe. eek 246

Micromammalian evidence for palaeoenvironmental change................ 253

PMideNce roll COMMUNILY COMPOSHION . 1. oa4.25--- 522-52 eee le eee 253

Evidence tien iiiedan Size Variation = 6.2 2... ei. ee ee See i Day.

PVidence rom: SpECIeS GIVETSILY 20.659). Peeks. 2s Soe A ees Bw ecklees » 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

Boamplaas Aree cee ca) os eterna en Ae eM aye tee eee ae 314

Byneskranskopel sos ac8:..055 0 Wert eaten cee erat ais > ey Maas ne noes 318

Die Kelders 1;Middle\Stone Age samples) s..¢5.. see: «2 ee 320

Coastal:samples:.. bch ake arays oe ete tadet caters ace: eke ee 322

Holocene andimodeni‘samiples... 2.235522. -be 0.65) + 05 eee 828

Upper Pleistocene andmoderm samples). 05. 2a. n4 54)... 4 eee 324

Existine evidence tor Late Quatemany environments <..5>..5.-.55 ere 326

Marine evidence 20... na av anctces eater ae Whee eee eee oo

Terrestnial 6Vide@nce spice raha Aoeitn cine qin ios ss 4s oan er 330

Correlation of micromammalian and existing evidence...................... 335

General’ framework 22) Sires an ace 25 oo Sanlous aus gone Seale eae er 336

Southern Aifvicam €vidence . . oo). ssh abs = eae oo eee eee 338

Macromammealian*evidence. . . .2.cnc. a sais, ees ee 5 6 ey)

Micromammiahan evidence tor siteicormelation. 22. >...2 27..-+.- oe 347

Prehistory and environmental change... 455.) -..44. 44-2 on eee 351

Géneral. considerations 22. Ss seta ae a ee aa Se eee 352

The southern-Cape Province: 0. oe. 6. woe lous 5 See ak ee 356

CONClOSIONS yao.) sade 55, nce al ae She el oe 362

SUMMIMABY, soy) Sis aa ec resis: san lope tel gah waaay a hs edi ae Pe aes Ae ae 364

ACKNOWlEGGEMENIS:: «0. 6's... ong aie sae anal SeSeeoe Boles, GPaae oe ene ee 365

IREfGheMCGS: taka oss ene o> kes hoeas oe aad ee ee 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 EF. 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

[]100-200mm KY 400-600

= 200-400 Z 600 +

z +

a SSS,

VSENG SSS TEAS =X

DS Yi = NUIT NONG

s CT,

‘ fynbos _ renosterveld Strandveld

Sw mountain /arid = coastal =

NE HT Ny temperate forest

Gao coastal mountain ma sreaiitn gaa 2

Ly. S

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 eft 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

HE 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.)

below datum

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 191

years BP

CBM

Sree 1630150

DGL

ISIO=75

pie I700*55

BLD 1955-65

BLA_ 6400:75

BRL 9100-135

104257125

re 124802130

12060

(ei

GWA/

HCA

LPC 22000

YOE=

a ee Sa @/>40000

OLP 42000

\ nn, ti, LMP Sa ihe vv Vv v

vvvvvy ‘vy Viv

»>40000

BOL

Be pee ey oe Bone oe

BSE EE Ge Bu

OCH Po Bsa ee 8 Oo a a oO

Se se eo o) ea

Fee eee ES fo a oe

14200240 UW30!

21100t 420 UW300

32400*700 UW304

historic occupation

burnt sheep dung

UW337

prehistoric herder

UW 307

UW338

UW336

occupation

Wilton

UW306

UW4I0

UW 4ll

UW4l2

Pta

Robberg

early Late Stone Age /

Pta late Middle Stone Age

UW305

Pta

Middle

Stone Age

UW308

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

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MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 193

leaf mould

partly disturbed

marine shell 3

concentration

)

compoct ashy layer 6

I2 black ash

tortoise shell

concentration

15 ash, brown soil

i 4 ash

19 ash, brown soil

—_—_ —

a — —

ae aie ia a ea ee

bedrock

0) Oro. Mm

Fig. 4 Section 028/29 at Byneskranskop 1 (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 Boomplaa:

DGLI DGL2 BLD2 BLD2A BLD3 BLD3A_ BL BLA BRL B

INSECTIVORA

C. duthiae Fe eee a. er. Pei eae 0,42 1,08 — 0,78 0,71 1,06 0,16 0,48

E. edwardi Stat et eh ee, Seer UES 1,26 1,08 0,99 0,78 0,71 1,42 0,99 2,40

M. varius ST iiata: cop ete <c) ee eee ey a LO 8,82 4,30 3,47 6,07 4,48 3,90 6,73 5,29

ONAVOSOFEX oe ec) es ee OS = — 0,50 0,47 0,47 0,35 0,16 —

Se Vanilla) a. Gaehep tls ey ee ao AOD 0,42 — — — 1,18 0,71 1,64 1,92

GCicyaneae 2 Oe a is ea i SE 4,20 B23 1,98 2,18 2,3 4,61 1,64 1,92

G) flavescens: ‘5% 5. Ss. ee TE 13,87 9,68 14,85 12,15 11,56 13,12 11,00 10,58

sub-total’ ?.°". 2.7 6 cee & 2A8S 28,99 19,35 21,78 22,43 21,46 25,18 22,33 22,60 2

CHIROPTERA

R. clivosus — — 1,08 — 0,16 0,24 — _— 0,48

R. capensis — —_ — —_ 0,31 — —_ 0,16 —_

?M. lesueuri . _ _— — _ _ _ —_ _— —_

INESITICOLIOnS Te VO iy eS CSN _ _ _ —_— — — 0,16 —

E. hottentotus _— — — — 0,16 — — — —

E. capensis _ _— _ _— — _— _— _ —

M. schreibersi —_— —_— _— _ — _ —_ _— —

sub-total — — 1,08 — 0,62 0,24 — 0,33 0,48

RODENTIA

Cahettentotus® Bay. = 24 See = CEA 11,76 17,20 24,26 18,69 16,27 12,41 12,97 6,73

As: SUDSPINOSUS#) «a -os ae ce ee ONS9 0,84 — — 0,16 0,47 2:13 0,66 0,48

As namaquensis’<s- <%= 3 > 3. = (124 10,50 9,68 7,43 7,01 8,02 7,09 9,03 14,42 1

IDicincomtuss. =e) 5" Ss Ges cs — — 0,50 0,31 0,24 — 0,16 —

Mminutoides- 5 a i. 7s Ss = AGIS 0,42 — 0,50 0,47 0,94 0,35 0,99 0,48

IBRVCLICQUXE. ee Aas ee te re) nS —_— 1,08 2,48 0,62 0,47 1,06 0,82 3,37

IR PUMIIO® te et Ce, Sees ante esol 2,94 — 2,48 2,34 4,01 2,48 0,99 0,96

Mealbicaudatus 3 5 7. = *. = = ets 5,04 6,45 4,95 5,76 6,60 8,51 6,40 35107

T. afra Se) ee Psp notes ys) oe es ee — — — 0,16 0,47 0,71 0,49 —

G. paeba — — — — 0,16 0,24 — 0,16 0,48

D. melanotis . — 0,42 — — 0,16 0,24 0,35 0,33 0,48

D. mesomelas — — — — — — — — —

?Dendromus . 0,59 0,42 1,08 0,50 0,16 _ 0,35 0,33 —

S. krebsi . — 0,42 3,23 0,50 1,25 2,36 3,19 1,64 2,40

S. campestris . 1,78 1,68 1,08 0,99 1,40 1,42 1,77 0,49 0,96

O. laminatus . 1,18 0,84 1,08 — 0,31 0,47 0,71 0,49 0,48

O. saundersae 7,10 5,88 10,75 6,93 8,10 (BOS) 6,74 8,21 6,25 t

O. irroratus 34,32 28,99 26,88 25,74 27,26 26,89 25,89 31,36 32,69 2:

O. unisulcatus 5” ee re 2 ee MES 0,84 1,08 0,99 2,49 1,65 1,06 1,81 0,96

GROculanis. (SS esl ow can ee ee SS _ — — 0,16 — — — —

sub-total) --). = <2 <0 a5 < 7ost5 71,01 79,57 78,22 76,95 78,30 74,82 77,34 76,92 7

IN Ht cae PS eto. Pe eer, eehOe 238 93 202 642 424 282 609 208

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

CL3 GWA LP LP LPC

, BRL4 BRLS BRL6 BRL7 CLi CL2 BG TBF GWA GGU GGL LPM _ 2B

| edie? O72 194 3,60 523° 2,89- 3,13 3,27 3,59 2,74 3,25

oo 203 -1,04 1,44 — 0:72 0:28: 0,14 — — — 0,11 —

eee es.10 «89648 )86=—«6,22—S «5,81 0s «13,67. «15,00. 26,17 25,28 © 30,39-- 22,99 22,11 39,61

= 0,25 — — O22) 26, ‘023 3,99 "7,95" 4,25. . 3:02 - 642 4,14

B meer 207. 21,67 «381,08 3= 1,442,144 s,52 1,69 «2,61 2S s«O86)=Ss 1,16 ~~ -1,30

iS eer 27s (23300 3,11 «2,58 3=— 2,16 ~—Ss«1,36 = = = = = =

5 Seemercose 7.51 = 5,74 7,53 10,79 «4,32, 1,79 4,28 2,29 2,30 2,53 - 1,95

Nene eee ee

i Gereme 1924 «19,95 18,90 19,14 34,53 27,50 36,50 39,33 42,81 32,76 35,05 53,25

hime 025 026 £«-- ee. a See Os 016) 058 O14 O01. ee

| <a = a a ae a eee Main? ee oe

= Ee a nes G77. 653 - = ie = ps. JON

iw om ve ze re xi ae: = a ee 504) as

[eee 026 — saat os oe a a on eats i ee

| ee a cai Es a cm =a af af is AQUI Se

Meo or 624 = 072 of 014 032 033 057. 053 —

| 96 11,39 1451 16,51 20,43 10,79 22,73 7,30 425 3,92 5,60 4,00-- 1,95

ma 06. 10-1002, “072. 045.028 024- == 029 021 0,65

Puen 1s44. 1088 789 10.75 1007 614 179 161 261 4172 1.79 » 1,95

Seo 025 428026 «2048 ss O14, 9. 008, © Otay © 0:11 eae

mens! 026 O48 022 — OP aie ie ee 029 0/53 0,65

| oss 259 «191 «| «043 Ci72.s—Cit Si 14 16S 33 29s«éS3 O65

topes! 120 O86 144 182 0355 -032 065 043 084 — 1,95

[esos 337 «(31119444 408 029 0,42 0,65

gs 026 024 - A: a 2 ti a pe on aN

| — Ose 006. 04. ae = am a = = rae a

iS Merl) 026 024 043 0,72 «068 1,10 080 1,31 . 1,58 1,16- - 4,30

Poe O78 024- BW MIA ce ae go Ragga. te nae

Ges = 024 es 2a" (953 O80 033° 029. 0:53 ~.. ~1.95

i. oe ae = a a ae es as a a ae

Mopeeiameiss 048 108 072 136 014 O16 — O14 — _

697 1494 11:40 9533 13,76 1727 1409 39,12 39°57 30,72 41:38 39,89 24,68

28,48 25.32 30,83 3660 29:03 2014 21959 702 618 12:42 100 9,89 6,49

if Peni 026 0.24 108 —-—- 023 427 49538. 458 359 4.32 3,90

| ees = 4) 22si«i72si”si Ci ti titi

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

! 460 395 386 418 465 139 440 726 1246 306 696 950 154

! Byneskranskop I (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.

194 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 2

Percentage representati

‘on of species in samples from upper levels in main excavation at Boomp

Daas

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 195

= CL3 GWA LP LP LPC

DGL! DGL2 BLD2 BLD2A BLD3 BLD3A BL BLA grid BRLS BRL6 BRL? CLI CL2 BG TBF GWA GGU GGL LPM 25

INSECTIVORA

e = 0,42 1,08 — 0,78 0,71 1,06 0,52 0,72 1,94 3,60 5,23 2,89 3,13 3,27

E cdwordl iis 126 OR 0.99 OTE O71 142 yp Ca a a eee iitae

Mi wadlus sofort ft gio 882 430-347 607 448319073. fo fio 68 «6,22, S81 13.67 1500 26,17 25,28 30,39 22,09 anil 39,61

Sey oh eb aie oe ee a A Ce er Ob se 2226 023399795425 “al02 wean “F'14

Geet, 8h 6 we oecme Oo Sind Ue — — — 118 0,71 164 sa7 «01S 2,07 1,67 1,08 1,44 1,14 1,52 169 2,61 0,86 116 1:30

C. cyanea. sy = 2 catia) yai20) °3123/aigs: iis: 2/36) alcrumtica S373 233 ML 5B 21636 2s gi oe Bh La

Gganea of lt EE Langa 1387 9681485 12151156 13,12) t}00 Oe ids 751574 753—('10,79 4327912822930 2,839

sub-total 24,85 28,99 19,35 21,78 22,43 21,46 25,18 22,33 me 1924 19,95 18,90 19,14 34,53 27,50 36,30 39,33 42,81 32,76 35,05 53,25

CHRP IER 1,08 0,16 0,24 6 0,

iv — - : _ ; , - = 0,25 0,2 _ = — 14 0,16 0,33 =

Be capensh SS = = of 5 = oie ms = oe Ea ee

2M. lesueurl = = = = — - - a = = = = = = te =

M. tricolor = = = = ir = = 016 = - 0,26 0,72 0,23 - 0,16 —_ 014 O11 =

E. hottentotus = = = = 0,16 = =- — = = - — = 0,14 =

E. capensis. = _ = = = = = - = 020 0,26 = = OL

M. schreibersi = = = = = = = == = = = = = = Ol

sub-total = — 1,08 = 0,62 0,24 = 0,33 048 43 -0,25—(0,78 0,24 - 0,72 0,68 0,14 0,32 0,33 0,57 0,53 =

RODENTIA 3 Oe 86

. hotte! Fe eee eee eS ThE) .26 18,69 16,27 12, 12,97 1196 11,39 14,51 16,51 20,43 10,79 22,737,304,

Saale OE cue op 2 oes eyni059) 0'SAU =e Eee 1016 10.47/92 oe (Oe yas ge Mt eo 7022072 Os aR Ogee iS gist te

Ainanaquensis’ © fe 2 ae = (Uy2a) 10,50" 19,68) 47/43 S701 8,020 7) 903 1442 isl 1544 10,88 = 7189 «10,75 1007S «6,14 s,79s6l 261 s72,—Ss«1'79S 1195

D, incomtus 3 Old Deore — = 0,50 0,31 0,24 — 016 — 022 0,25 «0,26 «= (0,48 — = 014 0108 = O14 9 O11 2

Pe omoes kk kk OB O42 SOT 094 = 35 0990s 02 051 0,26 0,48 = — 028 032 — 0,29 0153 0,65

Piverredixi Mees eee, ere) BLS 1,08 248 0,62 «0.471106. = 082337 435-253-259 91 0,72 0,91 0,14 0,16 = 0,33. 0,29 0,530,685

eC ee re eee nh ar a cc rn

Baa lcausains 58 ol eae heirs = = a O16 047 O71 0,49 a — 025 0126 ee LAO 280.24 a 0,29 0,42 (0,65

Gapaebalaas = = = 016 (024 —) loemmod — 025 0,26 = = FS = = = =

Bele = = Se a Le Ge aE GD oe Ee] ti i

?Dendromus 0,42 «1,08 += 0,500, 16 = 035 033 = 22.0250 cr we <= — H — —

S. krebsi. 042 323 «0/50 «1,25 2,36 3,19 642,40 87025 = ee AEE EU EB ED) ES NES

S. campestris» 168 ©6108 = 09940142177 049096 = =— = = = = a = = = =

(ONlamtnatus ewes «<. sipks hs Ps O84 «108 = 031047719 87 101,55 072 1,36 0,14 0,16 vs =

Otsatndersae) = <= SG 270) 1588) 10,75" 16,93 |8;10, 77155° 16,74) gala lamtogs 697 14,94 11,40 17.27 14,09 3911239157 30,72 41:38 39,89 24,68

Ouirroraus . 1 1 1 1 ttt 34,32 © 28,99 © 26,88 = 25,74 27,26 © 26,89 25,89 31,36 32,8 848 25,32 30,83 20,14 21/59 7/02 «G18 12:42 10,20 6,49

Ou unisulcaus 3. 1 1.) 1 TMB O84 108099249 1,65 1,06 1,81 086 $52 177 0,26 wy 7033 ata Sag ase 19128 ae

Giaculariste we Cases se eee = = — O16 — = i pein 0,72 0,23 014 024 — 0,29 —

q 19,8

sub-total . . . . . . « « 75,15 71,01 79,57 78,22 76,95 78,30 74,82 77,34 76,92 80,51 79,27 64,75 71,82 63,36 60,35 56,86 66,67 46,75

Sa ae 208 460

N Enns 169238: 93 202—S— 642i 282 09 395386 1399 440 «+726 +~«41246°«#306~S*«SG 154

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

1700 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 lattes

sample comprised a considerable number of subsamples based on lithological

units observed in the excavation.

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 3 m of

ane spanning the last 12 500 years without major breaks. The stratigraphy

8 4) comprises mainly dark-grey soil interspersed with ash bands and a

a a band each of shell and tortoise bones (Schweitzer & Wilson 1978).

leek Be as is divided broadly into two industries, a ‘pre-Wilton’ in

Fimiee sid , and a Wilton in levels 9 upwards (Schweitzer & Wilson 1978).

nese es of these major divisions into a total of five subgroups are

highest se ave not yet been refined. Pottery was recovered from the

The microfaunal samples from the pre-pottery levels is adequate (Table 4);

that fro

m 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 . ‘ 1,15 0,75 1,42 0,99 2,59 0,55 0,05 0,0 =

M. varius bela Spill 5,44 3,90 6,73 5,63 9,44 25,61 23,72 39,61

? Myosorex ‘ 0,26 0,47 0,35 0,16 0,28 0,34 6,49 4,87 7,14

S. varilla . 3 0,26 0,47 0,71 1,64 1,24 1,30 1,62 1,28 1,30

C. cyanea aes ne 3,83 2,25 4,61 1,64 2,48 2,33 — — —

Ga flavescens) ss.) 3554: 11,91 13,12 11,00 7,09 5,68 1,47 2,41 1,95

sub-total . . . 24,52 22,05 DSS 22,33 19,92 23,05 38,29 35,45 53,25

CHIROPTERA

R. clivosus 0,13 0,19 — — 0,28 — 0,15 0,20 —

R. capensis — 0,19 — ,16 — 21 — 0,05 —

M. lesueuri — — — — — — — —

M. tricolor — — — ,16 0,11 ,14 0,10 0,10 —

E. hottentotus — 0,09 — — — — — 0,05 =

E. capensis — — — — 11 — — 0,05 —

M. schreibersi — — — — — — — 0,05 —

Subsiomal 5 ¢ 5 WLils 0,47 — 0,33 0,51 0,34 0,25 0,51 —

RODENTIA

C. hottentotus . . 16,35 17,73 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

D.incomtus . . . 0,26 0,28 — ,16 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

P.verreauxi. . . 1,15 0,56 1,06 0,82 3,32 1,03 0,15 0,41 0,65

Rey DUNLILO ene ONG 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

T. afra Ae CRS — 0,28 0,71 0,49 0,17 0,07 — — —

G. paeba amie — 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 1,95

S. krebsi . = 8 0,77 1,69 3,19 1,64 0,79 — — — —

S. campestris meas; 1,28 1,41 aA 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 6,74 8,21 9,85 12,93 39,40 38,99 24,68

O.irroratus . . . 29,76 27,11 25,89 31,36 28,36 28,11 6,49 10,40 6,49

O. unisuleatus . . 1,15 2,16 1,06 1,81 1,58 0,48 4,97 4,10 3,90

G.ocularis . . . 0,13 0,09 — — 0,17 0,27 0,20 0,20 —

sub-total . . . 75,35 77,49 74,82 77,34 79,57 76,61+ 61,46 64,04 46,75

ING 783 1066 282 609 1777 1462 1972 1952 154

+ Includes Muridae indet. 0,07 %.

DGL = DGL — BLD2A

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 197

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 1 456 + 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 (KRMIA)

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. KRMI1A 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 II 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 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 4

Percentage representation of species in samples from Byneskranskop 1.

1 2 3 4 5 6 7

INSECTIVORA

Ce ASIAN CAE ea — — 3,33 1,41 1,92 3,05

E. edwardi . Re pe ie er ane — — 1,67 — — —

IVE SUORUUS: Fae a Se te Lt 8,70 — 10,00 17,46 20,05 19,08

Savaniila™- . Se. oe Es 24S — — 1,67 5,65 7,42 9,92

C. cyanea ATs. oy ee ee teh 4,35 — 1,67 — — —

Cy flavescens 2. «© « 3% £°* » 4486. 13,04. 20,00: 16,67 (2eivSs 14,01 8,40

sub-total we ew Uw. 35,14 | 26,09 - 20,00 - 35,00 37,10) SAS aie 46

CHIROPTERA

IRE CLIVOSUS < Pas as oad aes, © ns 1S — — 1,67 0,26 0,55 —

IRECGPCNSISY *, 2° 3 ePWES 4 FONSS 4,35 — 3,38 0,26 — 0,76

MiSTHICOlOP ce oS Gee 8 ye ee SS — — — 0,13 — —

EE NOUenIOLUS © eo ee wt — —_ — 0,13 — —

TEEMCOPCHSIS 2. «ef koe we ee bn!) — — — — —- ——

Muschreiberst. is 2 Oe eS — — — 0,26 0,27 —

sub-total” 3 ea ee IO 4,35 — 5,00 1,03 0,82 0,76

RODENTIA

Cahotieniows a) se — — — 0,64 — 0,76

Gecapensise.” =< 2 ee 38 ee. Sel 21,74 40,00 10,00 4,62 SL 7,63

VAS SUDSPINOSUS 5 “= 8 ek ek —- — — 0,39 0,27 #£3—

Ai MAMGQUCNSIS: .bees 5 os ey 8S — — — — — —

DDVURCOTNEUS: bc se oe | — — — — O27 —

M. minutoides eh pe ie ||P — — an 1,41 0,55 0,76

IPRVERLEQUXL” "so Ee ee oe SRALOS — — 3535 3,59 2,47 3,05

REDUMNUIO gon 8 eee AY sy 4) Sa 8,70 20,00 Gy Desi 1,65 0,76

Maalbicaudatus = 2" 2) 4 '. 15,4 4,35 — 5,00 3,98 3,30 53

MORO Dac hie a Ped a ean wee ee OAM 8,70 — 5,00 Bei. 1nGS 3,82

‘Demelanots .) ae .2- ON A ESS — = — 1,80 0,82 1,53

D. mesomelas . . 2. 6 9 ee — ~- ~- 0,26 0,82 0,76

(Dendromus “5 3. 5. =. » 1,35 — — — “= — —

SKreEDSE- eee a tS 8,70 — 11,67" 204a 19:51 19,08

OO laminatus . -.2. 6) 3. SS — — — 0,90 0,82 0,76

Ox saundersaé 2-30 +. -+i 3 « 96 4,35 — 11,67 1207 7 eer Dt

Ov irroratus; .2.-%s «= . 6,76 13504, 20:00... TikGy 5p 5,49 6,11

sub-total ooute hs «© 4+ « 62,16 69,57 60,00 60,00 ‘GIES > S5>-ees 5

ING Sate Son ee a Be 74 23 5 60 We 364 131

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

i \IICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 199

MSGS «(962 «10,71 S7,79 «13,64 20,47 «S«9.77S«s«12,50«s«10,58 S541 13,85

Mma) 20,13) «32,14 24,68 26,14 37,43 33,33 27,78 31,73 18.92 3241

1,48 a O64 1,19 = -- a = _ = = 0,28

9,24 — — — — — —- — — — — 0,28

= = = = 1,30 == 0,58 = = “ = 0,28

- a = = = — — = 1,39 == = 0,28

9,95 — 28 1,19 1,30 = O58 |. 10:57) — 2.78 “= _ 1,94

a4ae 1.30 © ©: 11,28 os a a TAT = _ a a= 0,28

Seeeeesoe.0s «68,33 «10,39 Ss 5,68 =Ss(i5,85 0 s2,87 «5,56 «8,65 sé, (6,65

‘s was — = a= = OSS. RIS) 139-> 1.92 = 1,39

—_ — — = = = a = = 0,96 oe —

= = = = — — = — _ — = 0,28

Rigs. 1,30 1,92 == 130 - hi4 0,58 = = 0,96 ~- 0,83

P45 «1330 =~ 1,28 = PCO ela, OR OST ©) 139% + 11,922 2.9023. 21,94

SSO) 9 2.56 ~2,38 , 2,60 _ ede AIO 278" » 3.85 - 9, 2.708.305

Bieesteeass «65,95 93,90 3,41 3,51 2,30 1,39 = 240" = 229

S@eeeiesgs 49 «5.95 11,69 1250 1228 6,32 833 .6,73- 81° 6,65

1,38 = Wome 238 1,30 1,14. 058 0,57 == 0,96 = 0,28

bk Ve aa = — ia.” O58 OST) 1.39). - 0,96 = 0,28

0,71 = poe) 2:38 1,30 a 0,58 0,57 a= = — —

Semmes 1190 7,79 455 3,51 13,22 4,17 3,85 2,70° 5,26

1.48 2,60 = POS eeOr 7. WRIT, 230). 27S. 0,96./ “270.1539

ei eee ote 16.67 15,58 25,00 17,54 19,54- 18,06 17,31 24,32- 19,94

£51 5519 8,97 Seon e098 P59 ADS: 37D (22,22 «319,23 ~-- 27,03" 15324

D,57 58,44 68,59 66,67 74,03 73,87 61,99 66,09 69,44 68,27 81,08 65,65

421 77 156 84 ai| 88 171 174 72 104 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.

198 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 4

Percentage representation of species in samples from Byneskranskop 1.

1 2 3 4 5 6 7

\!WCROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 199

INSECTIVORA

C. asiatica . 2,70 = = a 14 1,92 305 2,56 pl ey 18 BS 402 1,39 3,85 2,70 222

ect ear 13,51 8,70 — 10,00 17,46 20,05 190g nae 1218 1310 7,79 909 12,28 9,779.72 1442811 15,24

8. varilla i a KD Op Gioy ST RIG 9512 et 222 1 a 2 eee Oe

: ae = 4,35 — 1,67 = a 2 es = = =

Gicyanedine# tyiten i) sve . , = ay

Cflescens . . . - + + 1486 13,04 20,00 16,67 12,58 14.01 8,40 Cie et LOT TS eee 13,85

subtotal . . . - + + 3514 26,09 20,00 35,00 37,10 43,41 40,46 4,56 30,13 32,14 24,68 26,14 37,43 33,33 32,41

CHIROPTERA

R. clivosus . 5 6 comes Js = _ 1,67 0,26 0,55 = = 0,64 0,57 1,39 — — 0,83

R.capensis. . - » - - - 41,35 4,35 = 3,33 0,26 = 0,76 - 0,64 1,19 0.28

M.tricolor. . - +» + + = 0,13 — eS sy “a i = — Ea

E. hottentotus. . - + + 2 = = = = 0,13 — = = 0,28

IENCGPENSIS!:) es — = a = — 1,30 — 0,58 — — — — 0,28

M.schreibersi. 6 1 6 6 6 aaa = = 0,26 0,27 — = _ 1,39 = = 0,28

Babolat ee een ncC 4,35 a 5,00 1,03 0,82 0,76 = 1,28 1,19 1,30 — 0,58 0,57 2,78 = = 1,94

RODENTIA

C.hottentotus. . . 2 2 8 = = = = 0,64 = 0,76 30 © 1,28 — — — 1,17 0,28

G. capensis. . . . . . ~ 811 21,74 40,00 10,00 4,62 5,22 7,63 130 7,05 8,33 10,39 5,68 5,85 2,87 5,56 8,65 8,11 6,65

Avsubspinosus. 5 6 so — — _— 0,39 27 — 0,58 1,15 1,39 1,92 _ 1,39

A, mamaquensis . . 6 6 = = — — 0,96 = —

DUCT SS a Gf ceeeceece | Poe — == — — 0,27 — = = = = = 0,28

M. minutoides sete ar ee 535) — —_— — 1,41 0,55 0,76 1,30 1,92 — 1,30 1,14 0,58 — — 0,96 — 0,83

P.verreauxi . . . . . . 4,05 = — 3,33 3,59 2,47 3,05 130 1,28 = 2,60 1,14 0,58 0,57 1,39 1,92 2,70 1,94

R.pumilio. . . . . . . 35,Al 8,70 20,00 1,67 2,57 1,65 0,76 130 2,56 2,38 2,60 — 1,17 2,30 2,78 3,85 2,70 3,05

M.albicaudatus . . . . . 5,41 4,35 — 5,00 3,98 3,30 1,53 130 3,85 5,95 3,90 3,41 3,51 2,30 1,39 — 2,70 2,22

T.afra. . . . . - «Sal 8,70 — of 3,72 1,65 3,82 130 4,49 5,95 11,69 12,50 12,28 6,32 8,33 6,73 8,11 6,65

D.melanotis . . . . . . (1,35 _ — = 1,80 0,82 1,53 — 064 2,38 1,30 1,14 0,58 0,57 — 0,96 — 0,28

D.mesomelaa. . . . . . 0,26 0,82 0,76 1,30 = =s = 1,14 0,58 0,57 1,39 0,96 = 0,28

i2Dendrormusit ware fy cue) 535) = 1,92 2,38 1,30 — 0,58 0,57

S.krebsi . . . . . . . 13,51 8,70 — 11,67 20,41 19,51 19,08 2857 17,31 «11,90 7,79» 4,55 3,51— 13,22, 4,17 3,85 2,70 5,26

O.laminaus . . . . . 0,90 0,82 0,76 2,60 — 2,38 2,60 2,27 1,17 2,30 2,78 0,96 2,70 1,39

O. saundersae. . . . . . 9,46 4,35 — 11,67 12,07 12,91 12,21 11,69 17,31 16,67 15,58 25,00 17,54 19,54 18,06 17,31 24,32 19,94

O.irroratus . . . . . . 6,76 13,04 20,00 11,67 5,52, 5,49 GI 519 8,97 8,33. 12,99 15,91 12,28 »:13,79--22,22:19,23 27,03 15,24

sub-total . . . . . . 62,16 69,57 60,00 60,00 61,87 55,77 58,78 5844 68,59 66,67 74,03 73,87 61,99 66,09 69,44 «68,27 81,08 65,65

NEW omen Cer ie 74 23 5 60 719 364131 71 156 84 71 88 171 174 2 104 37 361

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

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

Im 5 loam

10 eboulis secs

(1/12 loam, eboulis secs

0 14

1-14 MSA levels

Fig. 5. Stratigraphy of Die Kelders 1 (after Tankard & Schweitzer 1974).

201

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS

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202 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. duthiae ee. — — 0,28 0,26

A. hottentotus- . © s . 0,95 0,93 — —

IVE VQTIUSE © Far oar 8 2,86 0,93 1,87 IRS)

Se Vara so isl — — 0,52 0,13

C. cyanea 5. 0oe — — 0,03 —

CG. flavescens 11,43 5,61 3,92 3,62

Sub-total” 20s) 3% 15,24 7,48 6,63 5,56

CHIROPTERA

IRCUVOSUS <3 4 5. = 0,95 — 0,03 —

De COPCTISIS 50 53, 0,95 — — 0,13

PE plesicus. =» *. = \; — — — 0,13

M.schreibersi . . .. == = — 0,13

sub-total . . . . 1,90 — 0,03 0,39

RODENTIA

C.hottentotus . . . 2,86 2,80 1,63 3,10

GACGPCUSIS: =k. ie 4,76 4,67 1,53 1,68

Dincomtus 2) °. s : 2,86 — 0,07 0,26

M. minutoides a Me — — 0,03 —

PS VENTCQUKT a ey 4,76 —- 0,21 0,52

Repumilion 6 Sev 2,80 0,59 0,91

M. albicaudatus. . . ~- — 9,40 8,15

T. afra Sid eh: — — 25,70 22

DD melanous .° 5) 2% 0,95 — 0,31 0,26

Os laminatus. =. 7,62 ie 0,35 1,16

O. saundersae . . . 21,90 13,08 50,12 50,19

O-arroratus. =. Ss 2 31-43 57,01 3,09 3,49

OR UnISUICATUS.. 7... — — 0,31 1,81

G.ocilaris;, ~ % =~. — ~ 0,93 — =

sub-total . . . . 82,86 92,52 93,34 94,05

Ns nae ee yh Sore 105 107 2883 773

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS

B/|

BSBB/|

H/G

BSBH/G

GSL

BSL

YSL

a micromammalian samples

YGGL

Grey Loam

Rubble Horizon

Black Loam Ill

Black Loam Il

Black Loam |

Pale Brown Loam

Basal Loam

Fig. 6. Stratigraphy of Nelson Bay Cave (after Butzer 1973).

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 7yto alba (barn owl) on the basis of shed feathers collected in the

shelters. Both T. 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 Uuenkraal

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 205

TABLE 7

Percentage representation of species in modern samples.

BPB-C BNK2 GLEN NGA-B

INSECTIVORA

LIS — 2,38 — —

iniae = wks £19 — 2,19 0,98

A.hottentotus . . . — 0,13 0,73 —

547) 3. ¢ 0,48 0,13 — 0,25

MORI Ct 19,21 9,14 7,83 24,02

REONOIES 2. 2. lke — 0,75 —- =

ermal kt 1,83 9,51 4,01 1,96

Mercyunese St; 4,60 L735 — Pao |

eenaVeSECHS . .. 0,40 B13 1,46 0,49

SKogGgyasSp. . . .« — — — 0,12

subtotal ss... 24-10 26,91 16,21 30,02

CHIROPTERA

WNANNCOGIEG =... wt 0,08 — = =

CABCNSIS . -: .--. 0,08 0,25 = 0,37

Se) 0,08 — 0,36 0,12

re nonenmowus . . . 0,16 — = —

MCGDCHNES .° . .CS 0,16 0,13 0,55 0,12

WASSCRYEIDEVSE ~. . Cc == = 0,18 —

Chiroptera indet. . . — — 0,18 —

SHe-Oldl | sk 0,56 0,38 1,28 0,61

RODENTIA

Genoientous << . . 1,51 oss i 0,61

GS GOACHMIN OE — 6,26 5,10 —

A. subspinosus . . . 1,83 0,75 — 521

A. namaquensis . . . 11,59 2,63 — 13,36

Bimcomus = . . == 0,13 = 0,25

M.minutoides . . . 8,81 11,39 9,84 4,29

PMOIGICNSIS.. . .. . 17,70 — ~ 3551

Te VECVCGURE 5 se 1,83 4,88 0,73 8,46

ain (SC — 1,00 0,73 —

TEOPUMAIONI TO. 4,44 7,26 3,28 2,33

M. albicaudatus. . . 0,16 0,13 — 0,12

Cig 1! — 1526 — =

WCIGHOUS © 5 wt a 25 1,82 1535

Deamesomelas . . . £35 2,00 6,38 1,59

POCUOFOMUS. Sooo. — 4,63 = 1,35

Sa 0,16 4,01 — =

S. campestris i Le 1,43 —= — 0,61

Ovjammaius. . .° . 0,63 0,63 0,91 0,86

Oisanadersae 2... . 5,40 0,63 0,36 4,04

CD 11,67 15,89 53,37 19,00

Gemcwlaris . . . . 0,08 = — 0,37

sub-total 2° so. T1573 12,12 82,51 69,36

NS ee 1260 799 549 816

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 207

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 MNycteris 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 hoes 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 1s 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 Otomys 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 er al. 1973:

266). This finally leaves Tyto alba as the most likely creature to have accumu-

lated microfaunal remains in caves. In addition, 7. 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 (3”)

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 (%”) 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 2 mm 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 24

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

212 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 number

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. laminatus 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 ef 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 1s 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 215

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 ef a/. 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 DAT

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!~%) = upper tooth row (UTR)

alveolus

posterior palotal foramen

zygomatic arch

left mandible - lingual aspect

cheekteeth (M3) = lower tooth row (LTR)

ee Sema

incisor

body of mandible

Insectivora (Crocidura flavescens)

1-3

length of tooth row g |

5 lower alveolar length (LAL) ——

G) CICRD,

re)

right mandible - buccal aspect

| (coronoid spicule

O 1Omm coronoid process P P

| rocumbent

4 2 (p )

Soa a approx. condyle

length of mandible

plus incisor

height of ascending ramus

Chiroptera (Rhinolophus

We premaxilla

(A oe eer oman

Dre

TINS Zt

infraorbital canal

a Nera) right mandible - buccal aspect

Fig. 7. Diagrams of species representative of micromammalian orders with terminology used.

(See Meester 1963 and Rosevear 1969 for further explanations. )

clivosus)

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 small 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

hNO

ANNALS OF THE SOUTH AFRICAN MUSEUM

FACTOR |

5 _E.edwardi

C.flavescens

20 C.hottentotus

5S _S.campestris

0) es

5 en Se es ee soe De noe oe |

5 ,S-krebsi

10 M.albicaudatus

) oe eaten,

5 _0O.laminatus

Q.irroratus

A.namaquensis

5 _Rverreauxi

S ._R.pumilio

abcdefghijk Imnopq rs tuvwx

aso =o a © o& ©

DGL Wy {EI te q SOR 2

BLD3 jy VOL r OLPS

BL k BPI s BOL

BLA | SBiR2 t BOLI-3

BRL m BP3 u BOL4

CL n BP4 v BOLS

GWA o OLP w OCH

EP p OEP x LOH

FACTOR 3

O.unisulcatus

pees i Gee

FACTOR 2

5 C.duthiae

40 Mvyarius

5 S.varilla

5S D.melanotis

0) Wy aeoe

40 O,saundersae

A.subspinosus

5 _M.minutoides

s 2 e@eer cy

abcdefghijkIimnopqrs tuvwx

Fig. 8. Variation in percentage representation of species loading highly

in the analysis of species from Boomplaas A (. = less than 0,5%).

MICROMAMMALS AS

FACTOR |

30) OLP

40. OLP3

BOL

abcdefghijkIimnopqrst

C.duthiae

E.edwardi

M.varius

S.varilla

C.cyanea

C.flavescens

C.hottentotus

A.subspinosus

A.namaquesis

M.minutoides

--— FQ 4M A@ooTd0a

=A Sb fey asl Gy St Sf = ox

FACTOR 2

40 GWA

BPS

(0)

| _bar,

abcdefghij kIimnopqrst

PALAEOENVIRONMENTAL INDICATORS

Pverreauxi

R.pumilio

M.albicaudatus

D.melanotis

S.krebsi

S.campestris

O.laminatus

O.saundersae

O. irroratus

O.unisulcatus

FACTOR 3

30. DGL

30. BRL

abcdefghijk!Imnopqrst

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%).

D2? 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 223

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.

DOA 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. 7yto 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

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS DDS

composition of prey-remains of other birds of prey or carnivores which. hunt in

or near the territory of T. 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 T. 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 J. 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 7. 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

it.

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 T. 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 1s 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

iemancibie:: 1) M. varus + 3°C. cyanea +12? = 15

memandipic: 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 (1965b: 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:

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 231

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-

Doe 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) (commas eeshe

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 1965b). 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 233

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 DSS

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 M, 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 P, to M3, being the distance from the posterior surface of

M; to the anterior surface of P,, measured along the line of the jaw.

TOw>

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. Ms 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 2517.

D. foramen mentale to alveolar margin, being the distance between the

foramen mentale and the superior border of the body of the mandible

relia ear

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 of

M,, M,_>, 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,_>, 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 = 10s by Simpson er 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 1s 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 = BEN where S = number of species, N = number of individuals. The

index of evenness is e = —4— 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

OSs CUS):

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

239

Taxonomic list of species encountered, together with their English common names.

Order Family

INSECTIVORA — Chrysochloridae

Macroscelididae

Soricidae

CHIROPTERA Nycteridae

Rhinolophidae

Vespertilionidae

RODENTIA Bathyergidae

Muridae

Cricetidae

Muscardinidae

Genus, species

English common name

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

Myotis 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 pricier 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 ef 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 er 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 et 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; 19066: 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 ef 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 is 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 19056: 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 190S5a:

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

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 it 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. irroratus

O. saundersae

G. ocularis

P. verreauxi

D. melanotis

D. mesomelas

C. c. cyanea (500 mm)

A. namaquensis

S. campestris (500 mm)

TABLE 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

Hillside

E. edwardi

M. varius*

C. flavescens

C. c. cyanea

A. namaquensis

P. verreauxi

O. laminatus

O. saundersae

O. irroratus*

Plains

C. asiatica

A. hottentotus

G. capensis

R. pumilio*

M. albicaudatus

T. afra

D. melanotis

S. krebsi

Riverside

M. varius*

D. incomtus

R. pumilio*

D. mesomelas

O. irroratus*

Sundry

S. varilla

A. subspinosus

M. minutoides

R. rattus

* Score divided arbitrarily between two categories.

TABLE 13

Proportions of species in different habitats near Byneskranskop.

sparse vegetation

dense vegetation .

fynbos

sparse scrub .

fynbos

?restioid element

dense ?proteoid element

cultivated, grass .

TESS. lg Lae a ee

pans, cultivated, grass

grass, Acacia .

PEASS: Phe

cultivated, grass .

dense grass

grass

grass, fallen trees

reeds, grass

grass, Acacia .

near trees .

grass, reeds

Number

249

Percentage

0,13

4,78

5) P2

1,86

2,79

5,18

0,66

8,37

27,76

2592,

0,13

6,64

3535

0,13

8,23

Doe

4,25

28,69

4,91

0,13

3,85

8,50

19,52

— Se

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 Tyto 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 251

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 vA No Wa

Valley floor

C. duthiae PSE UNORES Pec, ica teea es. SLD 1,20 8 1,00

M. varius* Memscnverctation =." 5.4 92 .: « 12t 9,66 98 1207

Mauascmoms open vepctation . . . . . . . 19 1-52 5 0,63

D. incomtus dense waterside vegetation. . . . — — 2 0,25

P. natalensis PHASS CUlbiVated ..6 2 ee le we 6223 17,80 45 5,63

R. pumilio MEANS Wereiation:? 2) “eee. 3 4 56 4,47 19 2,38

MTIICHATIES PCASS ws Se st 2 0,16 1 0:13

D. melanotis MEHSEEetASS Bot eto et. Gee 4 AD 3,19 11 1,38

D. mesomelas grass +trees/bushes . . . . . 17 1,36 13 1,63

S. krebsi PERS. os en 2 0,16 — —

S. campestris open woodland + grass . .. . 8 1,44 5 0,63

O. irroratus* MeHSENVePCiAtOM 2 o:0- 5. 22 ig. 7A 5,91 78 9,76

Hillside

E. edwardi @pen scmi-amd scrub... 5 sy 6 0,48 2 0,25

M. varius* Meascyerctalion... .< ..... «,. «. 124 9,66 98 wart

C. c. cyanea PieIVAOOSsicg © 6 ee ee, wm ee SB 4,63 18 225

C. flavescens Memseavereiatien: =... Ses ss 5 0,40 4 0,50

PeMinosey | GSNSeSChUD . . ... . . . «* 23 1,84 43 5,38

Beumnnnueays Open SCtUD . . =.=... . . ». ~~. 146 11,65 109 13,64

P. verreauxi MMOS Tacnse-) 2 See ose | DB 1,84 69 8,64

O. laminatus EEE SS SA Se ee eo 8 0,64 A. 0,88

O. saundersae PRASS UNIDOS 4 Ek io 2 Tw OS 5,43 33 4,13

O. irroratus* Memseweremuon.. =. . =.) <«. «. 13 5,83 Th 9,64

G. ocularis epemiscnupt mr. Beco Ok 1 0,08 3 0,38

532 42,46 463 57,95

Sundry

S. varilla RPMS ee ed Ca =o ae eS 1,84 16 2,00

6 35 4,38

M. minutoides eee tte em ree ee cg RS. Ola 8,8

LN) SS oa ee ene Sie emer nt oP25 799

* Score divided arbitrarily between two categories.

252 ANNALS OF THE SOUTH AFRICAN MUSEUM

cases, however, the valley floor would appear to be heavily over-represented.

This suggests either that T. 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 253

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

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

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 —O,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 D355

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 1 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

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

YOL 1,01588 BL 0,97979 OLP2 1,02529

BOLS 0,91898 DGL 0,96491 OLPI 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 257

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 1

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.

258 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 17

Species loading highly on factors in the Varimax rotated factor matrix (A) and Oblique

rotated factor pattern matrix (B) for Byneskranskop 1.

Factor 1 Factor 2 Factor 3

M. varius 0,85083 C. flavescens 0,70743 O. laminatus 0,92387

S. varilla 0,88919 G. capensis 0,90159 O. saundersae 0,72222

D. melanotis 0,83358 R. pumilio 0,74821 O. irroratus 0,73970

S. krebsi 0,96512

M. minutoides 0,72959

S. krebsi 1,00507 G. capensis 0,95846 O. laminatus 0,87055

S. varilla 0,91596

M. varius 0,84172

D. melanotis 0,78344

81,6% of total variance accounted for. Factor correlations low or very low.

FACTOR |

M.varius

O FACTOR 3

5 _QO.laminotus

FACTOR 2 (0)

5 M.minutoides 20 C.flavescens O.saundersae

5 _D.melanotis 10

30 S.krebsi

O.irroratus

5 R.pumilio

0) fs)

125678 9IIIIIIIII i25678 9111 11ill 1256789 III II Itt

ABO1 2345679 ABOI 2345679 AB O12345679

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 1

“o

Te)

fo)

20 II

Ke)

fo)

20_ 14

20. 15

FACTOR 2 FACTOR 2 (cont.)

FACTOR | (cont.)

10 Te) | 2

0 O a

abcdefghijkIimno abcdef ghijkimno abcdefghi jkIimno

a C.asiatica e G.capensis i M.albicaudatus m Q.laminatus

b Mvarius f M.minutoides j T.afra n Q.saundersae

c Svarilla g Pverreauxi k D.melanotis o O.irroratus

d C.flavescens 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 7%).

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

) 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

15 0,70501 9B 0,84664

16 0,88258 10 0,71036

17 0,86701

19 0,88958

13 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 ay 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 I 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 |

SD .A.subspinosus FACTOR 2

10 _Pverreauxi 40_ M.varius

0 ! er a 30.

T.afra 20

10 FACTOR 3

) S ,C.flavescens

O e©eee?ee@

S.varilla

ne) lO M.albicaudatus

O.irroratus

30 O O ise rte Ses

20 S _M.minutoides 20 QO.saundersae

10 e 10

5S D.melanotis

O O O

eh Be Se As) L 3.5. ¢ Stl 13 l 33) meee lS

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

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

A, subspinosus 0,89375 D. melanotis 0,85728 O. saundersae 0,81984

P. verreauxi 0,89097 M. minutoides 0,92113

O. irroratus 0,96112 C. flavescens 0,98743 M. minutoides 0,94282

T. afra 0,93805 M. albicaudatus 0,78883 S. varilla 0,92614

P. verreauxi 0,91248 O. saundersae 0,74763 D. melanotis 0,84101

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

3 0,73229 1 0,86177

4 0,89936 DD 0,80720

5 0,86842 12 0,79641

6 0,91322 13 0,81997

W 0,91296 14 0,74322

8 0,91306

9 0,86275

10 0,87094

11 0,88203

6 1,04775 1 0,99601

8 1,04580 13 0,87854

Al 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

—OOWAIAMARWHN !,

0,78890 12 0,79377

0,75778 13 0,85395

0,89531 14 0,90062

0,84395

0,91891

0,89630

0,91986

0,85388

0,83454

0,90444

jemeh meh

ev)

NOWNUOANHE DO.

1,03659 14 0,84118

1,02994 is 0,81722

0,98597 |e 0,78936

0,97617

0,96771

0,88156

0,85075

0,83634

0,76808

—

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 ANNALS OF THE SOUTH AFRICAN MUSEUM

FACTOR |

“%o

FACTOR 2

SI) %o

20 FACTOR | (cont.) 30. =|

fo)

10 | ie 20

@) 20] 10

30 20 | (0)

20 10 | 40 2

10 fe) 1 = 30

O 4 20

30 10

20 0

10 12

(9) . A 20

(0)

10 20

O 10

20 20

10 10

O ® e O

abcde fghijk!Imno abc def ghi jk |Imno

a C.asiatica e G.capensis i R.pumilio m D.mesomelas

b M.varius f A.subspinosus j M.albicaudatus n OQO.saundersae

ce S.varilla g M.minutoides k T.afra o O.irroratus

d C.flavescens h P.verreauxi | 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

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

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 ;

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

verreauxl 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

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

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 BNK1 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 Croci-

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

BNKI1: 1 0,92329 DK1 M.S.A: 3 0,93394

y 0,86238 4 0,98497

5 0,90716 5 0,98596

6 0,93519 6 0,97390

7 0,89096 if 0,98565

8 0,86791 8 0,98093

9A 0,87869 9 0,97217

9B 0,78084 10 0,97645

10 0,95231 ilk 0,96473

11 0,97358 12 0,83609

We 0,90922 13 0,71645

13 0,86676

14 0,91204

15 0,90712

16 0,82248

17 0,79429

19 0,86448

BNKI: 6 0,98901 DK1 M.S.A: 7 102513

11 0,98452 8 LO1752

10 0,96754 4 MO 2Z2

9A 0,96334 5 1,00761

8 0,94502 6 0,99812

5 0,94372 10 0,98780

a 0,92715 9 0,97808

1 0,92101 11 0,96408

105) 0,88448 3 0,87517

14 0,85260 12 0,73803

12 0,84869

9B 0,83782

2 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 1 Factor 2 Factor 3 Factor 4

BNKI1: 1 0,91154 DK1 M.S.A: 3 0,88514

Zz 0,87867 4 0,97039

Bi 0,86818 5) 0,96722

6 0,90341 6 0,95426

Ui 0,83742 7 0,97045

8 0,80173 8 0,97866

9A 0,81366 9 0,96167

10 0,92246 10 0,95671

‘el 0,96692 11 0,95212

12 0,92068 —- 12 0,78523

13 0,91092

14 0,95965

15 0,87502

16 0,85971

by 0,82840

19 0,89241

iB:

BNKI: 11 1,00342 DKI1 M.S.A: 8 1,03686

14 0,99307 qi 1,01440

2 0,96243 4 1,01118

1 0,95995 J 0,99676

Ze 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

Ls) 0,84938

7 0,81348

ii 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 ZT

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

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

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

BPA: BLD3A 0,80190 BNKI1: 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

Crile 0,88865 12 0,73449

BNK 1-19 0,71169 15 0,81116

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 273

M.varius

O. saundersae 4

2 3 4 5 6 7 8 9 10 | l2 13

x [OOO yrs BP

Fig. 14. Correlation of Boomplaas A levels BLD3A to CL1 with the Byneskranskop |

sequence. (Solid line = Boomplaas A; dashed line = Byneskranskop {3)

pg: 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 DIS

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.

ee ee

Factor 1 Factor 2

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

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

eee ee

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 verreauxt 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

BPA: YOL 0,86001 BPA: OLP3 0,71777 DKI1 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

OLPI 0,73770 T 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

DK1 M.S.A:13 0,85308

NBC: YSL 0,87318

YGL 0,88311

BPA: OLP3 0,98289 NBC: YSL 0,71206 DKIM.S.A:2 0,91355 KRM1A:15 0,88213

BPI 0,97444 32 0,87497

OLP2 0,96506 BPA: BOL4 0,86883

OLP1 0,88768 BOLI 0,80520

DKI1M.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 Day

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 is

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 K S

BPA: DGL . . 10 ~ 6,9-8,7 7,74 0,36 0,09 106 2,4-3,5 2,86 0,23

BLD3 . . 70 7,2-8,5 7,80 0,31 0,07 131 2,5-3,5 2,90 0,21

BE sn 4 OP EQ ESl 7,88 0,38 0,11 69 2,5—3,4 2.95 0,21

BEA. . <<. 36 -7;4-9:0 8.12 -0:3%. 0:13 54 2,5—3,4 3,02 0,19

BRE. . .° 72. 7429.2 8,25 0,38 0,09 96 2,6-3,8 3,10 0,23

Clo... ae S50) 1290 8,34 0,39. 0,10 56 PGi) 3,16 0,23

GWA... 4_~—s 8, 2-92 8,43 22 3,0-3,5 3:25, O;17

LP ee oe Gr ES=89 8,41 0,35 0,19 69 2,9-3,7 BA - 0,22

YOL.. «. «. .4 °° 7,6-9.0 8,60 8 2,9-3,6 3,25 0,29

BP4. . . 8 _ 8,0-9,0 8,48 0,44 0,37 29 2,7-3,5 3:05 0,17

OLP 2s) 46-6) 2810292 8,52 0,45 0,47 12 2,/—3,3 3,01 0,19

OLP1 6 16> 7:9=972: 8,63 0,46 0,48 5 3,0-3,4 3,22 +~0,18

OEP2)" 373. 26) 77-955 8,44 0,44 0,18 40 2,7-3,6 3,19 0,19

OLP3 ee BiSwerS 2 —8rs 8,47 0,23 0,14 42 2,8-3,6 3.11 . 0,23

BOL * «7 dl -72=9'0 8,17 0,50 0,33 31 2,6—3,5 3,03 ~=0,21

OCH oy Be Ze POSS. 9. 8,38 0,30 0,19 19 2,9-3,6 3,13 0,18

BRB 88 Sc 56x a i3e8.3 7,70 0,40 0,41 A 2,5—3,2 2,80 0,23

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 95% N Range xX S

Unit DGL

DG 3. ae lS 7,6-8,5 8,01 0,26 0,15 14 2,7-3,4 2,92 0,20

DWGE2." .% see ol 6,9-8,7 7,66 0,39 0,14 44 2,6-3,5 2,88 0,20

BEDE 3 2. (re 9 7,2-8,3 7,66 0,33 0,25 16 2,0-3,2 293 0,30

BED. 40 oes aa 8 7,3-8,5 7,75 0,40 0,33 12 2,4-3,3 2,94 0,25

BEDQZA sec. 7 7,3-7,8 1263, O16 O85 20 2,6-3,1 Zot O15

Unit BLD3

BIDS fe Bie 39 7,3-8,5 a1). 031. 0:10 I 2,4-3,4 2.85 0,20

BEDSAL, Goss 8 ol 7,2-8,5 7,84 0,32 0,12 56 2,5—-3,5 2,96 0,23

Unit BRL

BRL Ed de gas walk 7,5-8,8 8,23 0,38 0,19 22 2,6-3,3 3,07 0,19

BRE2 ee Pe) eS lg 7,6-8,9 8,18 0,34 0,18 25 2 1=3e7 3,04 0,20

BRL4/4A . ... 6 7,7-8,4 Seis) O27 1029 14 2,7~3,3 3,01 0,19

BRS) 2 ce Hel 7,1-8,7 Sal2) O38) 2025 11 2,8-3,4 3,03 0,18

BREG? se 3205's .20 7,4-9,2 8.41 0,42 0,19 24 2,6-3,8 3.22, 0:28

Unit CL

BRO estan case 6 et LO 7,9-9,0 8,46 0,39 0,19 20 2,8-3,6 3-25, 0,21

Cl yg “acan® (eek 7,9-8,7 8,18 0,30 0,18 16 2,7-3,6 3,16 0,20

CL h ey Waka dee 6 9 8,1-8,9 8,50 0,31 0,24 8 2,8-3,7 3,20 ~=— 0,31

CE3BGy esc. he 9 7,7-8,8 S513) 0:43) 10533 12 2,8-3,1 2,98 0,11

Measurements in mm. 95% = 95% confidence interval for the mean.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 279

D: length of lower tooth row E: length of mandible plus incisor

95% N __ Range x s 95% N_ Range 4 sg 95%

0,07 6 11,4-12,6 12,08 0,40 0,42 9 16,8-19,2 18,32 0,85 0,28

0,06 S tis —-i3et 12,26; 0553, .0,45 6 17,4-19,3 17,97 0,72 0,75

O0G> 13 1158-13,1 12,27 0,38 0,23 25 16,5-20,0 18,40 0,77 0,32

OG. 6 11,9-12,4 12,00 0,31 0,33 9 17,1-19,1 18,12 0,66 0,51

0,05 19 12,0-13,6 12,71 0,43 0,21 20 17,9-20,5 19,19 0,73 0,34

0,08 11 13,0-13,9 13,30 0,27 0,18 11 19,0-20,9 20,06 0,72 0,48

O1s <2 1 (13,2) 3 19,0-20,4 19,70

2 12,2-13,6 12,90 3 18,3-20,6 19,40

0,26 1 (18,8)

1 (13,1) 2 19,9-20,4 20,15

i 8.7)

O09 6 13)1-142 13,47 0,50 0,52 6 19,5-21,3 20,45 0,67 0,70

Oi 8 12:6-13.9> 13,35 0,47 040 8 19,0-21,2 20,29 0,88 0,74

Oe? 10.7-13,2 12,95 37 18,4-196 19:10

ee, 3 12,3=13,0° 12,67 3 1922197 19:53

Gee 3 15-120 11,73 4 17,4-18,3 17,83

D: length of lower tooth row E: length of mandible plus incisor

95% N_ Range x s 95% -=N Ramee el Ga Boe

ft 233) 4 17,3-19,0 18,33

O10, 3. 11,4-12,1_ 11,87 3 16,8-19,2 17,93

2 V2-0-12,6° 12,30 2 18,6-19,2 18,90

0,14

oie. W1eS=1351 «12,22 0,57 0,60 3 17,5-18,0 17,67

Oi 2: 12,0-12,8. 12,40 3 17,4-19,3 18,27

O30" 3” 1234-129 12,70 4 18,6-19,5 19,08

Oe 6) 1221395 1268 044° 0,18 6 18,2-19,7 18,90 0,53" 056

Osos) DS, 9 12,0-1350- 12,42 045° 0,49" 3° 17,9-20,2 18,77

OhG) 2° 1257-13:0 12,85 2 192-1955; 719535

On2> 3) 12,9=13;6 13,17 5 18,8-20,5 19,82 0,68 0,84

19,0-20,9 19,94 0,83 1,03

19,0-20,7 19,85

19,8-20,9 20,30

(20,3)

1322-1355. 13,25

13,0-13,6 13,24 0,26 0,32

13,1-13,9 13,40

(13,4)

i=)

WwW

i=)

—

= W nbd

Re WNN

278

TABLE 30

ANNALS OF THE SOUTH AFRICAN MUSEUM

Size variation in mandibles of Crocidura flavescens from Boomplaas A and B.

B: depth of bod

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS

’

279

D: length of lower tooth row

E: length of mandible plus incisor

A: height of ascending ramus Y mandibk Ly, > 95%

N __ Range s 95% N Range RX me 5 95% N_ Range s 95% N_ Range Ke s

12,6 12,08 0,40 0,42 9 16,8-19,2 18,32 0,85 0,28

BPA: DGL ar a ee eet ee peer BE O10 sia 1228 053045 6 174198 1797 0,72 0,75

BLD3 Go es 20 Rea ae 8 ey 20 op On 13. 11,8-13,1 12,27 0,38 0,23 25 16,5-20,0 1840 0.77 0,32

BL ,2-9, i i F 2,5-3, } 0,17 0, 9-124 12,00 0,31 033 9 17,1-19,1 1812 0,66 051

BLA 36 7,4-9,0 8,12 937 ee os 25-34 3,02 0,26 0,10 6 ine 1271 043 021 20 179-205 1919 073 0134

BRE Pies aaa f ey 10 oes 70-139 13,30 0,27 0,18 11 19,0-20,9 2006 0.72 0.48

(Gil 50 7,7-9,0 8,34 0,39—0,10 ae ae 3,16 021 0,08 11 13,0-13, , ?

GWA 4 8,2-9,2 8,43 ,0-3,5 3,25 0,21 0,18 5 9

LP 16 7,9-8,9 841 0,35 019 69 29-37 394 025 013 1 (13,2) 3 oe 10

YOL 4 7,6-9,0 8,60 8 29-36 3,25 2 12,2-13,6 12,90 1) sai ,

BP4 8 80-90 848 0,44 0,37 29° -2,7-3,5 3,05 0,25 0,26 2 199-204 20.15

OLP 6 8,0-9,2 8,52 0,45 0,47 12 2,7-3,3 3,01 1 (13,1) é uae 20,

OLPI 6 7,9-9,2 8,63 0,46 0,48 5 30-34 319 :

OLP2 26 7,7-9,5 844 0,44 018 40 27-36 4719 O19 0109) 6, TS ETE2 147 01501052) 6 18s 2 a Oa

OLP3 13 8288 847 0,23 014 42 2836 3411 0,28 0,14 8 12,6-13,9 13,35 0,47 0,40 8 peal Eu a Haat

BOL 11 7,2-9,0 817 0,50 0,33 31 -2,6-3,5 3.03 0,25 0,21 2 12,7-13,2 12,95 atae 19,7 19.53

OCH 12 7,9-8,9 838 0,30 0,19 19 2,9-3,6 3,13 0,23 0,18 3 12,3-13,0 12,67 3 19,2- yee

BPB. . 6 73-83 7,70 0,40 0,41 7 2;5-3:2, 280) 0,19 0,24 3 11,5-12,0 11,73 4 17,4-18, ,

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 D: length of lower tooth row E: length of mans plus 95%,

N Range = =-X& s 95% N_ Range s 95% N Range & s 95% N_ Range s 6

Unit DGL

DGLI 15 76-85 801 0,26 0,15 14 2,7-34 292 0,20 1 (12,3) 4 17,3-19,0 ne

DGL2 31 6,9-8,7 7,66 0,39 0,14 44 26-35 288 020 018 0,10 3 114-121 11,87 3 16,8-19,2 17,

BLD . 9 72-83 7,66 0,33 0,25 16 20-32 273 030 : :

see 8 73-85 7,75 040 0,33 12 2433 294 025 2 12,0-12,6 12,30 2 18,6-19,2 18,90

BLD2A . 7 7,3-7,8 7,63 0,16 0,15 20 26-31 281 O15 O11 0,14

Unit BLD3

BLD3 . 39 73-85 7,77 0,31 010 7524-34 2,85 0,20 17,5-18,0 17,67

yeaa , , , 3, d y O19 0,07 6 11,5-13,1 12,22 0,57 0,60 3 17,5-18,

BDA 31 72-85 7,84 032 012 56 -2:5-35 2.96 023 021 O11 2 120-128 12,40 SU SUS ET

Unit BRL , . .

BRI2 18 75-88 8.23 0,38 0,19 22 «-2,6-3,3 3,070.19 017 0,10 3 12,4-12,9 12,70 ee UG) 03

BREGMA 17 76-89 818 034 O18 25 2,7-3,7 3,04 0,20 02 O11 6 122-135 1268 0,44 0,18 6 18,2-19,7 es , :

BRL6 | W 77-87 812 038 025 «1 esd 3,03 a O18 016 2 127-130 12.85 OO A Oy 0

Unit CL ® 7492 $41 042 O19 m4 2638 322 02 0,22 012 3 129-136 13,17 5 188-20,5 19,82 0,68 0,

nit 5 y > 's

BRL7 19 1 0,83 1,03

cli . 13 oe? «846 039 0,19 20 2,8:3,6 3,25 oF 021 016 2 1324133 1325 5 19,0207 9 334

CL3BG | 9 82 850 031 024 8 2a37 3,20 ait 025° 030 3 131-139 13,40 3 198-209 20,3

TI88 813 0,43 033 12 28-31 298 0 O14 O15 1 (34) 1 (20,3)

Measurements in mm. 2 :

95% = 95% confidence interval for the mean.

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 x S 95% N Range mA S

BNKGSS. 222. a 2 6,9-8,2 7,54 °0,36 - 0,14 31 2,5-3,2 2,83 0,20 }

LO ee Bre 16 7,2-8,3 Tis OAT 70:22 32 255-32 2,81 0,19%%

DRISEGQAI = 5 es es RESO 787 8 27-3286 One

M.S.A.4 .. . 6 6,9-7,6 7.33 O24 0325 i 2,5—2,8 2,74 0,10)

OR: 10 7,3-8,4 7,73 0,30 0,21 15 2,6-3,1 2,78. 0,129

10 5 7,38, | 7,67. 0135" 50343 8 2,6-3,2 2,81 0,184

KRMIA:6. ... 2 (8,5) 7 3,1-3,5 3,39 «=—s«O“, iS

15 6 8,0-8,9 8.37 «0339 > Oral 17 2,9-3,6 321° O25

DL aaa <a 4 7,2-8,4 7,80 14 2,7-3,4 3,04 0,19)

30 alee 3 7,38, 1 7,70 8 2,7-3,4 3,04 0,234

NBCAYSEa 3 13 8,1-8,7 S41 20519" Om 47 2,5-3,7 3.21 O22]

YGE eae 17 7,4-9,1 8,21 0:46 .0,23 37 2,8-3,5 3,15 0,204

BINK Oa) ot fe a eee 6,6-7,8 129) 0332 BONS 38 2A see 2,61 0,26 |

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

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 281

h of M,-; D: length of lower tooth row E: length of mandible plus incisor

ge xX s 95% N_ Range xX SF Oa, N_ Range 4 gs 95%

ects 0,19 0,08 23 10,9-12,2 11,70 0,39 0,17 31 16,4-18,7 17,70 0,63 0,23

6.2 5,87 0,19 0,08 10 11,3-12,7 11,89 0,47 0,34 15 16,5-19,2 1811 0,79 0,44

6,0 5,86 0,16 0,15 5 10,6-12,1 11,62 0,61 0,76 3 180-184 18,27

15,7 5,55 3 16,8-17,4 17,07

(6,0 5,85 1 5) 3 17,3-17,6 17,47

i) 1 (12,0) 2 17,9-18,3 18,10

6,6 6,40

(6,4 6,30 0,17 0,13 4 12,1-12,8 12,43 6 17,9-20,6 19,28 0,88 0,93

16,4 6,23

16,4 6,14 0,17 0,21 3 12,3-12,5 12,40 3 18,4-18,9 18,67

16,8 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

}-6,7 6,40 0,29 0,15 3 13,3-14,0 13,60 7 18,7-20,8 19,89 0,66 0,61

+-6,1 5,65 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 BRL5, and the second is in the uppermost levels, BLD2A to DGLI,

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

280 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 32

Size variation in mandibles of Crocidura flavescens from coastal sites

A: height of ascending ramus Bdepthiof body of andi

N Range RX s 95% N Range Xx 5

6,9-8,2 7,54 0,36 0,14 31

5 2,5-3,2 283 Qa

ap 16 72-83 7,75 O41 0,22 32 25-32 ai gis

K1:L.S.A.12. 3 77-80 7,87 PORTS 286 ong

DE Ds 6 69-76 7,33 0,24 025 11 25.28 974 gilt

6. 10 73-84 7,73 0,30 021 15 2631 7g old

‘a 5 73-81 7,67 0,35 043 8 26-32 281 gig

KRMIA: 6 . 2 (8,5) De E06

i 6 8089 837 039 O41 17 2936 31 95,

D 4 12-84 — 7,80 14-27-34 304 gig

5 3. 73-81 7.70 8 2734 dod on

SERS. 13 81-87 841 O19 O11 47 25-37 3.01 oy

YGL. 7 74-91 821 0,46 0,23 37 28-35 315 on

BNK2 2566-78 7,29 0,32 0,13 3821-32 2.61 gag

(9 5,87 0,19

ICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 281

: f lower tooth row E: length of mandible Plus incisor

95% ni ae > 4 s 95% N_ Range >< s| 95%

12,2 11,70 0,39 0,17 31 16,4-18,7 17,70 0,63 0,2

0,19 oe e Hee 11,89 0,47 0,34 15 16,5-19,2 18,11 0,79 0,44

0.15 5 10,6-12,1 11,62 0,61 0,76 18,0-18,4 18,27

5 3 16,8-17,4 ii.or

8 3. 17,3-17,6 17,

He i fr 2 179-183 18,10

en 0.17 0,13 4 12,1-12,8 12,43 6 17,9-20,6 19,28 0,88 0,93

i344 6,30 0,

Nerd 0,17 0,21 3 12,3-12,5 12,40 3 184-189 18,6

8,6-21,0 20,26 0,95 1,18

12,7-13,8 13,40 0,48 0,77 5 18,6-21, 1s

on 3 oe 13.3-14,0 13,60 7 18,7-20,8 19,89 0,66 0,

565 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

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 Crocidur

above. Tables 30-32 give the resultant d

Figures 16-21 present the same data in

a flavescens as was described

ata for different populations and

Ure 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

vullibe assumed that a larger mandible indicates greater body mass. In general

ee ee C. flavescens is shown to be larger during the Upper

theninesnseacs ieee Holocene. The only exception is level BOL where

an those for the earliest part of the Holocene

ples for level BOL may clarify this situation and it will be

i A

Mportant to discover whether the lower BOL levels follow the same trend or

whether specimens from these levels will be larger. In the ne ee

depth of mandible means are highest during the period pe ae see

to about 14.000 B.P., that is the last glacial bead e <a ets

height of the ascending ramus the mean is SSS at e oe Ovi

period. In all three parameters an earlier high occurs in leve a icra of

when means are highest for length of lower tooth row (Fig. Ree Bain

mandible plus incisor (Fig. 19). The data for level OEEZ nae neat In

based on a good sample, but those from level OLPI require i ot ieee eee

View of the correlation between large size and glacial pape. with Berg-

fo assume tentatively that the species is behaving In Be soit

mann’s Rule. This being the case, it would appear ee aed by a milder

during the Upper Pleistocene of two main cold periods an Meme the

period, and that the Holocene was era warmer than

recedi iod represented at Boomplaas A. : : ards

3 aa Hae 14 000 B.P. onwards there Is a Risen ch hie

smaller means in the various parameters up to and De ae by the

sample from Boomplaas B. That this trend ts far Pines are perhaps

means for the upper levels at Boomplaas A (Figs. ae et ostglacial levels,

‘wo main periods of fluctuation. The first is in the oye BLD2A to DGLI,

CL3BG to BRLS, and the second is in the uppermost eer ramus that

and Boomplaas B. In the first it is the height of the ne te coefficients of

fluctuates most noticeably. This parameter displays mo es than changes in

Variation (Table 33), which suggests that real fluctuation ei he are perhaps to

Population composition are being monitored. Such flucte the later period of

be expected during a period of climatic change. ae und in the depth of

fluctuation, on the other hand, the greatest amplitude is fo

282 ANNALS OF THE SOUTH AFRICAN MUSEUM

M.varius

C.flavescens

6,0

5,8

abcde fghj |mnopqs a b cde f abcdefghijkImnopqrst

1234512 123451234

a DGL hele op OlER2

b BLD3 LRG p OLP3 al DGLI b2 BLD3A e4 BRLS

c BE NAIL q BOL a2 DGL2 ec BL e5 BRL6

d BLA k BPI r BOL4/5 a3 BLD d BLA fl BRL7

e BRL | BP4 s OCH a4 BLD2 el BRL WZ CL

fF GIL m OLP t LOH a5 BLD2A e2 BRL2 es (eer2

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 oo 4 C

YGL |} —_______s 7

KRMIA-6 +—_}— 2

15 }—_—_—- i) 9

22 +—+— 3

32 5

BNK2 [$< SF

DKILSA- 12 | 67

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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

NBC-YSL | —¥_ i

YGL |__| 7

KRMIA - 6

15 | |g

32 —== 3

BNK2 F ney

DKILSA-12 H—}H 3

BNKI-5 eee)

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mm 57 16 165 169 173 / 17,7718! 185 18,9. 193 197 2ON mzo5neo9) 25

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 population 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 BRLS 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 [3

BLD |

BLD2 _ a ——___:

BLD2A |< 7

BLD3 BLD3 SSS 9

BLD3A | —____$_ ie | 6

BLD3AM ——oaEEEE 17

BLD3AM2 De ee

BL ee

BLA TS

BRL BRL |_-—_____ ig

BRL2 | —__ 7

BRL4-4A - - GSR 5

BRL5 =

BRL6 — —_—_— ae (20

BRL7 CL | _ a?

CLI | 3

C2 | a 9

CL3BG 9

mm SOP ee Se ero. tel 79. BS BS. BS Sr! BO. Ot > 293

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

288

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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 G D E

_. SENG?) i 4,51 6,61 3,44 3,54 3,90

MES we SG 3.33 2,61 3:27 2,87

BeeemmggIcnmSiS. =< ww kl kl 3,24 3,58 8,67

T. afra ee Se nS 3,08 3,17

Mmemmeeemiolws. 5 . wl wll 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.

bi di. P(1-sided)

BPA: BLD3/BLD2A

i Sr 3:15 44 0,005-0,0005

B PD ek we 0,83 93 n.s.

Cc ee 171i 38 0,05

BPA: BRL7/BRL2

c 2 2.35 24 0,025

BPA: BRL6/BRL5

A I ee 1,90 29 0,025

B ee ta 2,06 33 0,025

Cc PP ee 0,63 20 ES:

* 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

BPB/BNK2_ BPA: BL/BNK1:5 BPA: CL1/BNK1:19 BPA: LP/NBC:YGL BPA: GWA/NBC:YSL

4,51 S55) 2,44 0,24

> , >

A*® 5 >

B 7,28 4,24 12,46 2,86 1,25

C 4,07 4,50 8,86 2,34 1,55

D 2,00 4,87 R35

E 3,90 3,95 9,61

x 4,57 4,41 9,57 DAS 1,01

BPB/BPA: CL1 BNK2/BNK1: 19 BPB/BPA: BL BNK2/BNK1: 5

A 6,23 6,31 2,34 3,43

B 12,86 7,66 5,36 8,43

Cc ,6 3,89 De 2,30

D 12,87 3,89 4,60 1,74

E 11,33 5,54 3,20 — 3,05

x 10,39 5,36 3,64 221

M. varius

BPB/BNK2 BPA:BL/BNK1: 5

A 4,67 7,32

B 6,63 3,10

C 4,19 1,69

x 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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MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS

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‘SOUS SNOLIRA WO] sniuDA xadosodpw ul “Wd Jo yIsus] “qd iojowesed pure ‘siosrour snjd sjqipueuw Jo ysua] ‘gq Jo}oWRIed UI UONeLeA “EZ “SITY

t 4

ai Ole! ec] «Sie! El «U'el:«OCO)«C'z) «(fel «bel eel Ol S “Stl =p ail ol gos OM ee Oo HS 2g og ww

el Ee ee ace eee er non IE ae re eS

2 -—_}+_—" 00 = HOT

b) a HOO

2+ 8 aim S-p 108

ee ee }+—_— oy) 108

ee a €d10

a b ———— 2d10

a ae oc SE Id 10

OE ($$ d710

a 82 +} bad

aE 62 ida

9 a ———$ C=! OA

1 $$ od1

e+ _—4 0S —_— aa) 474

€ -_—________}+____ 2 a — vO

S| ee SS ae 62 [> — a 19

Q , —___{_ a ee —)— 92 + aT]

i] S —m-, vie

a ee 9) le aT

I 9| a ¢a19

6. a —§ 0! a! 490 -vaa

89)Q 2¢ -—_ gON

Es en 6¢ }+—_ a+ ada

6S) SSS EE ¢¢+—_ i

See + a 8

S¢ +} @___ SS} ——_ 2 See b -VSWING

ee +o }+-—_Sa S-1IyN@

ee a Zl-vSTINd

89 ~—eea a oe ZING

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 Ms, 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.

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.

Aethomys namaquensis and Tatera afra

TABLE 37

B: length of M,_;

Size variation in teeth and depth of mandible in Aethomys namaquensis from the Cango valley.

A: length of M?-%

C: depth of body of mandible

| N Range X Ss 957, ON ‘Rance Xx 5 ~95%, N Ranse “xX S 9507

Pei 23 «6—5,1-5.8 5,49 0,19 0,08 27. 5,1-5,7 5,38 0,20 0,08 43 3,1-4,5° 3,77 0,34 0,10

BEDS = 26 35,2-5.9 5,56 018 C07 34 5,0-60 5,45 0,17 006 34 3,0-43 3,79 0,34 0,12

BL 1, 4—-0,0 15,64 O17 0,08 31 5,1-5,9° 5,53 0,17 .0,06 - 31 3,2-4,8 3,90 0,32 0:12

BLA fees 5 on 97 O17 O10. Tt .5,3-5.7.. SAT 018 0,12 8 3,64,6 4,11 0,29 0,24

BRL 136 5,1-6,0 -5,60 0,16 0,03 88 5,1-6,0 5,54 0,18 0,04 83 3,0-4,5 3,88 0,34 0,07

CL B20) 5,09 O20 “O07 23 °~5:3-5,8 5,57 0316 0,07 28 -3,4-4,6 3,94.028 0.11

EP. wee -5.9, 5,74 0,17 021 1X5: 7) 5 3,0-3,8 3,40 0,40 0,50

BP4 Guns5—),9 5,75 0,16 0,17 J a5l—5,8p° 9,92 -0:350°°O,38 — 10) <3,;3-45'. 3.75 0,27 0,19

OLP1 a 5:6-6,1 5,90-.0,20. 0,25 9 5,4-5,8 5,60 0,13 0,10 12 2,8-4,1 3,63 0,41 0,26

OLP2 Bor 5-0,4 3,96 0,21 0,07 33 5,4-6,0 5,72 0,16 0,06 35 3,0-4,7 3,82 0,39 - 0:13

OLP3 34 5,6-6,4 5,94 0,20 0,07 28 5,460 5,64 0,17 0,07 30 2,8-4,4 3,70 0,44 0,16

BOL . 4 5,8-6,2 6,03 6. 5;5-6;1 5:80.,0;26° 0:27 10 3,7-4:3 +403 021 O.15

Peet 51-59 5,51 0.20 0,10 18 4,7-5,8 5,32 0,30 0,15 18 3,3-4,2 . 3,82 0,23 0,12

Measurements in mm. 95% = 95 % confidence interval for the mean.

TABLE 38

Size variation in teeth of Tatera afra.

A: length of M,_, B: length of M!-?

N_ Range x S 95% N_ Range x S 95%

eee Se SS GF «| 5.0-5,2 §=65,09 0,09 § 0,07 3 4,7-4,8 4,77

19 Be oO 920. 4 9,28: « 022° 0:27 4° -§:1=5.6' ~ 5,30

BMK2 .. 18 4,7-5,3- 4,99 0,16 0,08 7 (4,6-5,2 . 496, -019 . O18

DKIMSA: 1 BP (9:2) 6 ~4;8-5,2, 5,00. 613 O13

3 18 4,8-5,2 5,04 0.15 0,08 19° 48-53 -4,95 -0,15 0,07

5 D2 jAB 9-4 5.13) . <ON6r: (O07, 382° 477-5,3 34,93 —-O.15; “OL05

zy 297 4:8-5-54- 3613.00.12 * 0.05 34° 46-53 491 ..0,17 0,06

9 Oe: “S:0-54S 5222 O15 = O11 14°" 45-5.) 24,88 20,20 ‘0:12

ae 0 .4:8-54--> 4.09" O16 . 0,12 1S) 94.729. 14592 0:12”. 0:06

13S 2 1 (4,9) 3: 1 -4:9>5'3' = «5,10

YGL Bp 14. Sle 5.2-6:3-". 3:76, 0,22, 0108 AS 49-59 546 0,19 - 0,05

Measurements in mm.

95% = 95% confidence interval for the mean.

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 C

BPA-DGL a ee 43

BLD3 _————————— en

BL {]_ a

BLA TT I—-— 9

BRL {jg

CL (PO 28

LP Ma.

BP4 = nO

OLPI [nS

OLP2 (SSS Ey rs

OLP3 EE 50

BOL 15

A B

NGB mS 7 er = 2

BPA-DGL | —_ a 23 Sa 7

BLD3 -—__—s—)}— 26 |/___—— 3) ____-- 34

BL [S19 a 8

BLA | 14 |__| |

BRL pa RE - 136 (> $a 88

ol =i S30 _— 23

LP a 5 ] 1

BP4 | -E 6 TT =

OLPI ee 5 aE 9

OLP2 | —_———__—1 36 33

OLP3 SSS a —+ 28

BOL +—}—44 cr 6

5| 5:3 5,5 57 5,9 61 63 mm4y7 .49° 5,17” 5, seSipmeonmoroemol

Fig. 24. Variation in parameter A, length of M'?, parameter B, length of M,3, and parameter

C, depth of mandible, in Aethomys namaquensis from the Cango Valley.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 297

qd» DGL fe) BOL C.hottentotus

bee DS, .p. OCH A

aunt: (2 al DGLI

die BEA: a2 DGL2 mm

e BRL a3 BLD2

eC: a4 BLD2A

g GWA el BRL

Ae, e2 BRL2

ie VOL e3 BRL4

j BP4 e4 BRL5

k OLP e5 BRL6

Meoerl fi . SRL7

meeOLP2 f2 CLI

OERS f3° CLSBG

A.namaquensis

6,0

5,8

5,6

5,4

5,7

5,5

5)8)

3,9

3,7

3,5

3,3

abcdefhj imno abcdefghijkImnop

a bcde f

| 234 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

DID (IP

NY)

Oo)

OLP3 BPB NGA-B

25 123456 123456 123456 123456

BRL LP BP4 OLPI

: a. i. z,.

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 7. 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 M3 and posterior border of symphysis are

apparently little affected by age or sex (Table 33). There is much more

300

TABLE 39

Size variation in mandibles of Cryptomys hottentotus from Boomplaas.

BPA: DGL

BPB-C

Measurements in mm.

TABLE 40

ANNALS OF THE SOUTH AFRICAN MUSEUM

A: alveolar length

Range

4,5-6,1

4,9-5,8

4,7-5,6

4,7-5,8

4,9-6,1

4,8-6,2

5,0-5,8

4,9-6,2

5,3-6,0

5,1-6,3

5,4-6,4

5,2-6,2

5,3--6,4

5,2-6,1

5,3-6,3

5,2-6,3

4,8-5,4

93/5

0,08

0,09

0,08

B: posterior border Mz-sym'}

Range

7,3-9,1

7,3-9,3

7,5-8,8

7,5-8,7

7,3-9,1

7,3-9,3

7,5-8,7

7,3-8,8

7,9-8,9

7,9-9,3

8,0-9,4

7,5-9,7

7,8-9,2

8,3-9,4

7,8-9,4

7,8-9,2

7,8-8,8

8,56

8,49

8,29

95% = 95% confidence interval for the mean.

Size variation in mandibles of Cryptomys hottentotus from Boomplaas A upper levels.

Unit DGL

DGLI

DGL2

BED2: -

BLD2A

Unit BRL

BRL

BRL2

BRL4

BRLS5S

BRL6

Unit CL

BRL7

CLI :

CL3BG

Measurements in mm.

A: Alveolar length.

Range

4,8-5,7

4,5-5,9

4,8-6,1

4,9-5,7

5,0-5,8

5,2-6,1

5,0-6,0

4,9-6,1

4,9-5,8

4,8-6,0

5,1-6,2

4,9-5,6

3325

D322

5,36

5,21

5,34

Spa)

5,45

5,42

3539

5,49

d552.

5,30

0,23

0,29

0,26

0,28

0,24

0,27

0,26

0,19

95.74

B: posterior border M,-symph

Range

7,9-8,8

7,5-9,0

7,3-9,1

7,3-8,8

7,3-8,5

7,5—9,0

7,5-9,0

7,3-9,1

7,4-9,0

7,5-9,3

7,8-9,2

7,3-8,6

8,24

8,19

8,03

8,25

8,26

8,03

95% = 95% confidence interval for the mean.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 301

attachment ascending ramus D: foramen sae ee margin E: depth of mandible

Range Ss

i ascending ramus D: foramen mentale—alveolar margin E: depth of mandible

ange X 5 95% N Range X s 95% N Range X 5s 95%

PROS tne

44,5 4,04 0,39 0,28 10 1,6-2,5 1,93 0,29 0,21 4 4,0-5,6 4,90

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

‘54,8 4,28 0,38 0,23 3° 4524 31,97 0:25 0,15 5 43-60 5,10 0,72 0,89

\6-4,6 4,04 0,29 0,15 it £524 71:88" 0:25 O43 2 4,6-5,1 4,85

/5-4,9 4,10 0,39 0,25 12 1,7-2,6 2,04 0,23 0,14 1 (6,4)

16-4,8 4,13 0,33 0,16 19 1,7-2,4 2,07 0,24 0,11 4 4,2-5,3 4,85

12-5,0 4,15 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

a9 22,035 0,13 30° (,6-2;5 -2.09: 0:21 0,08 5 44°52 458 041: D5

55,1 4,21 0,39 0,15 30 1,7-2,7 2,12 0,23 0,09 2 “St=5.2- S15

5-5,0 4,29 0,33 0,12 307 127-26 =2,11 O21 008 5 45-63 5,12 0:71 (0:88

64,9 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

5,2 4,19 0,43 0,16 30 1527 3207 0:25 _ 0,09 18 4,2-6,7 5,26 0,76 0,38

MALS AS PALAEOENVIRONMENTAL INDICATO Ss 3

300 ANNALS OF THE SOUTH AFRICAN MUSEUM MICROMAM CTO 01

TABLE 39

f ‘ation i ibles of Cryptomys hottentotus from Boomplaas.

Size variation in mandible: Sea Bposten on Geni hment ascending ramus D: foramen mentale-alveolar margin E: depth of mandible

: alveolar u order M,. im oy o/

Ni Renee A 95% Nis iRane xm ata 5 95% N Range X s 95% N Range X& s 95%

60 4561 525 031 008 6 7391 ei Te 035 0,10 60 1,5-2,5 1,93 0,23 0,06 21 40-60 4,89 0,70 0,32

BPA: DGL 5-6, 7 , 0,12 30 1,5-2,9 1,97 0,30 0,11 10 3,6-5,8 4.89 0,70 0,55

D3 BONA, 5:8) 732415 10:25) 710,09) 830) 73-9531 S00 416 0,33 214 1,92 0.25 0,09 7 44-61 5,07 066 0,61

BL. SON ATs) 3119022559 0,08) ae 30) 7,58 8 aae7 93 aia 044 O18 45 iso's 197-002) ook 1 (5:5) es

BLA 30 4,7-5,8 5,37 0,29 0,11 30 7,5-8,7 8,08 ail 0,34 0,13 1627 208 0.23 0.04 20 42-64 5,04 0.50 0.23

BRL 212 SU 252 0:26) 10,05) 121) 973-2 ages Al. 0336-0006 TMD Dil O01 00s 3142-67 515 0,69 0,25

CL 90 48-62 5,43 0,26 005 90 73-93 gig 424 0,35 0.07 A 17-26 209 0.22 0.08 (31503 ae

GWA 30 50-58 5,49 0,27 010 30 75-87 gig 430 0,42 0,16 Se re aero oe

LP. 30 49-62 5,55 0,32 012 30 73-88 821 441 0,30 0,11 PE OE IN) i

YOL 10 53-60 5,74 019 014 10 79-89 933 434 0,23 0,16 MO Eee eas. Coston 5,60 0,39 0,49

BP4 30 51-63 5,67 0,29 011 30 79-93 gia 4,50 0,31 0,11 3! ae EE OT IL Pe ae ae

OLP 14 54-64 587 0,32 019 14 80-94 858 451 0,34 0,20 4 11628 219 030 Ol 5,50 0,52 0,35

OLP! 30 5,2-62 5.78 0,29 0,11 30 7,5-9,7 8.69 4,52 0,43 0,16 30 1,6-2, . , tt Thy aioe beba pee ee

OLP2 30 53-64 5,82 0,26 0,10 30 78-92 860 4,64 0,40 0,15 Boe eee, Ces A, EEE NAG

OLP3 30 52-61 5,80 0,24 0,09 30 83-94 893 468 0,31 0,11 30 1,6-3,1 2,32 0, a Ty Hee Ey. OES

BOL 30 53-63 567 028 010 30 78-94 856 TE Meee Cenc here 8 50-67 5.19 0,60 0:50

OCH 30 5,263 5,79 031 012 30 78-92 gag 466 0,41 0,15 30 1,9-3,0 2,36 0,24 0,09 Bal bie) Wiad

BPB-C 16 48-54 5,21 O18 010 16 78-88 8,29 406 0,34 0,18 16 1,7-2,4 2,03 0,18 0,10 16" 4,126.8" (5,12, (0,651 10,35

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,-symphysiggph attachment ascending ramus D: foramen mentale-alveolar margin Econ of mandible Fey

N_ Range x s 95% N_ Range s ex s. 95% N Range X& s 95% N_ Range 6

Unit DGL

DGLI 10 48-5,7 5,25 0,28 0,20 10 7,9-8,8 8,24 O48 FHS 404 0,39 0,28 10 1,6-2,5 1,93. 0,29 0,21 oo ES, ig OS?

DGL2 20 4,5-5,9 5,22 0,30 0,14 =. 20 7,5-9,0 8,19 0,38 HHS 423 034 0.16 20 1,6-2,3 1,93 0,17 0,08 Dey Cy OP LOD

BLD2 . 13 48-61 5,36 0,39 0,23. «13. 7,3-9,1 8,03 0,50 M48 428 0.38 0,23 Pes el Ne) Bega ces

BLD2A 17° 49-57 5,21 0,26 0,13 17 7,3-8,8 8,01 041 3646 404 0,29 015 17 1,5-2,4 1,88 0,25 0,13 2 4,6-5,1 4)

Unit BRL

BRL 12 50-58 5,34 0,23. 0,15 12 7,3-8,5 8,03 0,40 $349 410 0,39 0,25 12 1,7-2,6 2,04 0,23 0,14 1 (6,4)

BRL2 19 52-61 5,55 0,29 014 19 75-90 835 039 [M3 413 033 O16 19 1,7-2,4 2,07 0,24 0,11 As ee toanaoad

BRL4 30 50-60 5,45 0,26 0,10 30 7,5-9,0 832 042 $3250 415 035 O13 30 1,6-2,5 2,07 0,24 0,09 Bact ea sa plOrstenorst

BRLS 30 49-61 5,42 0,28 «0,10 = 30 7,3-9,1. 8,24 047 P3653 422 035 0.13 30 1,6-2,5 2,09 0,21 0,08 5 He ero e

BUS 30 49-58 5,39 0,24 0,09 30 7,490 823 04 Bist 421 039 os 30 1,7-2.7 2,12 0,23. 0,09 2 51-52 5,

Unit CL 88

7 71 0,

cL 30 2850 $49 0,27 O10 30755-93825 OF FSO 429 033 0,12 30 1,7-2,6 2,11 0,21 0,08 5 tees doa od 042

3049-56 5,30 0,19 0,07 30 7,3-8,6 8,03 38 2 419 0.43 O16 30 1,7-2,7 2,07 0,25 0,09 Te eel

Measurements in mm.

95% = 95% confidence interval for the mean.

ANNALS OF THE SOUTH AFRICAN MUSEUM

302

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SSanaiCums 6 “C's VS 6p. Eps = oS*e ww ¢'9 Oy we -GiGF= YrG cre, _ “cic I's Ge Lr

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MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS

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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 is,

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 M3 to symphysis. In this case the modern BPB-—C

sample continues the trend but in the other parameters this is not the case.

305

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS

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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.

3 def. P

Bi Ee 50 cr a ne ZI 58 0,01

ie BRL G0 a ae DAD 29 0,05

prem i we OS 1,93 58 0,05

cere ee sw eh we a L220 29 0,05—-0,02

CLL CLS Gy a 2,54 58 0,02-0,01

EU we 2,58 58 0,02-0,01

ESE CIL, 75 i re 3,03 58 0,010,001

C -

bE OI Oe BL ai30 58 0,001

ele DOr. gk ew ese 2,66 58 0,01

OIE a km 8 2,04 38 0,05

OOM es te te sl CU) 2,09 58 0,05

i Ole Se 2,02 58 0,05

UL OCIS, ot eS 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 Ss c d € H

BPASDGE. So & ak & 848 Ppa 0,15 36h 0,73 2,26

BEDSie 0) ee Mae oe 1055 23 0,14 3,16 0,74 ZS

| 5 heal a Ne orm ea ee 280 A 0,12 3555 0,80 2,44

BAe ota he 604 23 0,16 3,44 0,73 DOM |

BRE ge 2s 5, tee le ae 1761 7) 0,14 Be 0,73 235

CE ar a) Pe 1449 21 0,16 219 0,72 2,20

GWaAs ge, 2 Sar 1805 19 0,28 2,40 0,57 1,69

| i ee el rd ae 2 a 1839 20 0,25 Mess) 0,60 1,79

LPC eis eae ee 140 15 0,27 2,83 0,65 175

VOGe a) ae Se 315 18 0,20 2,96 0,67 1,95

BP ag) ee, Me ie 353 18 0,24 2 90 0 66 1,90

BP2MA 5) seen BS 110 15 0,16 2,98 0,78 DAD

16] £28 eh eee eRe 93 16 0,20 3.30 0,73 2,03

ESP Ayp eer Sk ee awe 2 1708 pope 0,21 2ESZ 0,63 1,96

OLR ee ee 353 18 0,20 2,90 0,69 2,00

OLP1 ND negra ieee occ 1189 22 0,24 2397 0,62 1,91

OUP te eS 29-9500 23 0,25 2,40 0,61 ut 90

OLP3 (oe eee SAO. 23 0,23 2,64 0,62 1,94

BOE acetic sole. “2 1802 23 0,22 2,93 0,61 1,91

BOIS 0) 5 ee os 110 le, 0,17 3,40 0,76 2,14

BORA aT chen We 106 10 0,22 1,93 0,77 1,78

BOERS i) eM a Ge 96 2 0,21 2,41 0,73 1,81

OCH Saessatin Same ta 529 9543 0,20 3,19 0,66 2,00

EOE fe a ee 120 19 0,20 3,76 0,69 2,02

BE Bret tk. Wrnree ros cP T47 Wop 0,11 3,17 0,80 2,47

SE Geek Fehon er wa Pa 504 18 0,13 2713 0,81 2,33

INGA Ke. Pgs eT ae 352 21 0,14 3,19 0,76 as).

ING BW ante ooh ee a, Wc Glos 261 23 0,13 3,95 0,77 2,42

KERMA; 1065 ora 9 cad Carle AG gh) lO 0,14 2,34 ty OZ amar? 12

Say cl eee 103 12 0,18 2.30 0,81 2,01

VG: pet Ute ALS: Ae a Wis 43 8 0,24 1,86 0,81 1,69

D1 a te ra 42 8 0,26 1,87 0,77 1,61

5 A eee Re Pi 107 10 0,36 1,93 0,64 1,47

NBCBBSEY 7. oe uae ae 59 8 0,35 [Zz 0,68 1,42

WiSiiss. tts wn eee 2882 18 0,33 Dis 0,52 1,51

GES POL Sn eee 770 16 0,32 2,26 0,58 1,62

GIPEING So Oyo al wee dee 542 15 0,32 DDapi 0,64 rs

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 1 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 |

= e

e 0,9

0,7 0,7

0,5 0,5

abcdefghijkImnopgqrs tuvwx

lS 8 7 OAs

am DGE d BLA g GWA ols m BP3 p OLPI s BOL vy BOLS

DeBEDS e BRE i) (le k BPI n BP4 qe OlEP2 t BOLI-3 w OCH

c BE i GIL | {LPC iP BR2 Q OLIP r OLP3 u BOL4 x LOH

Bige sill

Aspects of community structure in samples from Boomplaas A, Byneskranskop 1 and

Die Kelders 1. (H = general diversity; c = dominance; d = species richness;

e = evenness.)

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 4 |

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.

a1 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

Variation in aspects of species diversity at Byneskranskop and Die Kelders 1.

BNKI: la .

9A .

9B.

£2

17,

BNK2

DK1: LSA 12.

ToaanZd

Me ith ih

MSA 1.

umbers of individuals

umbers of species

ominance

species richness

evenness

general diversity

TABLE 43

S13

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.

Holocene Unit 1

1 630+50 Herder scrub on hillsides, sparse

?semi-arid in places,

1955+65

Wilton

6 400+75

9 100+135 Albany

10 425+125

elsewhere ?dense microphyllous;

fairly open grass, ?trees on valley

floor, extensive dense riverside reeds

Upper

Pleistocene

12 425+ 130

or grass

14 200+240

fluctuating

rapid

temperature

rise

fluctuating

rapid

temperature

rise

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

moderate

mild

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

cold moderat?

MSA

SSE

relatively warm

UNIT 5

general increase in scrub

on hillsides, grass on valley floor

and dense riverside vegetation:

BOLS glacial maximum as in Unit 3

mild

harsh

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 BOLS5 indicates full glacial conditions, on the

evidence of the factor analysis, with the samples from levels BOL4 and BOL1

suggesting progressive amelioration thereafter. At the time of level BOL5 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 BOL5 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.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 347

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 BPl 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 11000 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 1 600 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

BRL5S, 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.

1 880+50 1 ioe

+ intermediate 1 mil

3 220+45 as unit 3 but more scrub and sia

3 400+55 2-4 less restioid vegetation on hills mild

UNIT 2

3 900 +55 =) extensive ?more open grass on flats; warmer

rather less restioid or ‘grassy’ vegetation

6 on hills and more scrub; less dense

; vegetation near river and lower slopes

Wilton 7 (?relatively dry) moderate

D 8

6 370+90 9A

6 100+140 9B ae

6 540+55 10

changing

vegetation

UNIT 3

11 mild

extensive dense vegetation near river

cS 12 and lower hillslopes; extensive ?more 12 mildest

closed grass on flats with pans and

13 some low scrub; restioid or ‘grassy’ 13 rather

?and proteoid element on hillsides general harsher

9 760+85 : 14 (peak restioid element in level 13) increase in

Pre-Wilton 15 (?relatively wet) ; temperature

= ze moderate

Au 18 harsh

12 730+185 A 19 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 6500 B.P. to

approximately 3500 B.P. is covered by samples from levels 10 up to 5

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 1 800 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 1 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 321

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-

tation and approximate dates (after Tankard & Schweitzer 1974).

Approx.

dates B.P. Vegetation Climate General Sea-level

UNIT 1

35 000 | extensive grass, ? lower

dense vegetation on

flats; scrub on hills

UNIT 2

3 intermediate ? higher

reduced grass, ? general

dense vegetation on increase in

flats; moderate temperature

restioid or “grassy”

vegetation on hills generally

mild conditions

perhaps

deteriorating

gradually

UNIT 3

12 slight change; ? lower

extensive grass,

dense vegetation on

flats; restioid or

80 000 ‘grassy vegetation

on hills

327 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 323

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 B25

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 BNKI1

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 327

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 '*O. In compensation the oceans will be relatively high in '°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 III, 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 '8O 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 +5-12 m 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 faunal 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 '*O 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 ef 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 eft 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 '%O 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 331

(Mangerud et al. 1979) has correlated the Eemian (Last) Interglacial with

oxygen-isotope stage 5e 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 er 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. (‘SO stages 5d

and 5b) interrupted a long, relatively warm period which lasted until about

73 000 B.P. ('8O stages 5c and Sa). 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 Br@rup interstadials of northern Europe can

apparently be correlated with '*O stages 5c and 5a 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 et al. 1971: 396) with the Bglling 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 11 800 B.P. and 10900 to 10000 B.P.

respectively. This represents the beginning of ‘SO 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 is,

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 10000 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 1000 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

336 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 !80 stages and major divisions of the last glacial

cycle.

70 last glacial cycle BPA BNKI1 DKIMSA

1 present interglacial

—13 000 B.P.-

2 upper pleniglacial unit 3

32000 EP

3 middle pleniglacial

——64 000 B.P

4 lower pleniglacial unit 5

—75 000 B.P.

Sa early glacial

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 5a, 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 BOLI. 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 et 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 Allergd 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,

338 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 (1972b,

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

9¢ I? ell c6l VCC Iv 8S 901

— _ 80°L os‘ZI 16'r prt Cre —

gfe sol. 16S 60'r 6S'L e9'yl ZS'ST SELE

6] “IS. 619 IS EhIC GEC. 9857 80'CE

— — —_ — Ut vor

3 — +60

weet Ole RO SC«‘iSCt(‘iRT:*CO:«CO YT «= CLO'ZE SSL

-- ee Sey HOT ONT. OG = OR - eL'e

8L‘7 ee Tp v0'l 7 = <

a = 697 v0'l 68°0 5 — —

a = [es = = = re “aes

rg = 88°0 os fi a7 ete —

9¢°¢ ESGT= “OLET S19 “86:6 pr'z LVS age

ane = pS'€ Iz‘s 6L'T pr'z aa: v6'0

— Ks <7 9ST 6L'I Ov7r = OL ET pox

— ies = ae cv‘0 Bi or ease

~~ = 69°C €£'8 LS‘€ “4 < ae:

a Bars 7% 7S‘0 680 “ — Sa

— Fa LL‘T 80'°7 — = 7L'T ca

aa but v0'T = = a —

oa = an zS‘0 680 Tas LT 160

a <0 = zs‘0 ae Sa = a

a =. = hak 680 37 sas —_

8L‘Z = 61°9 8£°6 ory 88°P SPE tL’

— = 880 80°7 = an a =

o> a 2 Ey gi mate on 160

dooys SIsouUOp

(yoqqui eva) yonqoys Acid

yoqsars /yOqusa}s

Josutrdsdiyy

yoqsurids

ysooqoyIey preyseq

ysooqoIeY pol/JsooqopyIM youyq

uvod/yonqg onyq

yonqpse.d ureyuNoUW

yonqposa1

puryo

npny 19}vo1d

oyeyng uPoLyy

SOY-}IEM

sid-ysnq

edsenb/erqoz ureyuNoU

SOIDOOUIYT YOuiq

SOId0ds JOUIIXA »

en ne EEIEEEEEEEEEEEE

nnn nnn LEE EERE

Od]

WINIpPOW od.1e]

WNnTpour |/euws

j[vtus— *jopul oeplAog

Sd1AD SIAC

SnjOaddvd vajad

souvjaul/siajsadiupa snaaaiydoy

SNSDAJOIAO SNSVAJOAAC

‘ds spoaopiyup

SDIAOP|,OAIU SHOSIOUDCG

“yopur rurydepsoy Vv

4SNIstdd SNSDALO]VSAW,

puipps snydvjasnq

snydpjaajp [nous sajapyaouu0)

snuinba|,snavydoona] snsvajod diy

pjnfnaoajnf pounpoy

wWinulpunaDd DIUNPAY

XAAO SNSIDAJOAND

sosaaisdaajs Snydvjasv [

daf{va snaaouds!

4 SMMDYUD SIAOAO] AT

‘JOpUl SePING

snaidolyjav sndaoyoroovyd

SnddO0d SNADOYIOUDI OT

VIALOVGOLLAV

4D8sonb|vaqaz snnby

4 sisuadpa snnbsy

SIUAOIIG SOADIIG ¢

VIALOVGOSSI add

Fe ee ee ae

HOT dtI0O TOA TD Tua vid caida Od

“Tod -dél “VM9D ~Ta

EN ee ee

‘(QL6[ Ul Ul ByEp UO poseq) V sevjduioog ye s[Ajoeporjse pue s[Ajoepossisod jo WOT}EJUOSIId9I 99"JU9I10d

Qp ATV L

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 E. 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 Taurotragus 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 1

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

Leo P PO aP ye — 0,84 133

puaeecrmidacimdct. . . « . . «= « me «6 0,68 1,68 Sela

POMEAEEIMECHIICU SIS. 9 (0), ee — _— 0,44

ARTIODACTYLA

RIMPMIMICUMCENS FOFCUS...- 2.2 2 \ fe Rk 4,79 D2 33

Suidae indet. ube BM SOR ee ie Go Fos. | ue 4,79 2,94 4,00

Hippopotamus Eeeeieuis es yee a ie tak 0,68 1,26 0,44

Se RES BEEP) ES Sane ee 6,85 7,56 4,89

SUIS ES SEUSS a — 0,84 222

PRE OT OPE nr — 0,84 2,67

WEECPVOVIUIG = 5 ll 0,68 — 1,33

Hippotragus spp. . PE age Soa est eee ge 0,68 Des) 4,89

Connochaetes sp. [Alcelaphus Sp. Eee cag ted hag eta es 2,74 2,10 14,11

Damaliscus dorcas . . Joe Cary ee et eae — — 222,

MECEPNIORCOUVGSUS 5 ee le ke we 2,05 1,68 1533

INU CENOECEIIDCSTIS. 5 6 «8k a Os 2,74 0,84 oe

DeIMNNIMELQHOWS Vi ae ww we 6,16 8,82 12338

JESU BES US SOO est 17312 DDT 8,00

Prramommrem eee eet Ss LR Me 0,68 0,84 0,44

Ovis aries. UE A ee ae oa a ae T55 — os

Bovidae indet. all Sly hoes a ee ae 19,86 23,95 10,67

SMIalmMeadn. “2 ws we Ok 10,27 2,94 4,44

FIBRES TTS UCN Gel a eee A ee a 4,11 5,46 20,89

PCr ke ir wer LEA aS 1,50 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 19746: 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

MRM oc.) ay Ge EY la kt eo 2,78 1,16 2,78

200 20. PL ea er — 0,29 —

ARTIODACTYLA

Hippopotamus amphibius 1-39 1,74 2,78

ReMOEMOTINS. 3) gw 1,39 0,87 =

Syncerus caffer .. TT, SR ee. tee, Re a 1,16 —

Tragelaphus strepsiceros ee eee ic 2 ine ES 39 0,58 —

PINNED Ce ee! ye lee Pe ee ey eS 8,33 4,93 13,89

BED ORS SOOT E A rrr a 339 1,45 —

meer IVOTIWHG we a — 0,29 —

iippeirarusieucopnaeus*s . . ... 6 1 ew ele — 1,74 —

2 EUS EALE ES SO, 6S ea eee 2,78 1,45 2,78

Damaliscus sp. . Sak Ro ek eer eer ee ne — 0,29 —

Antidorcas cf. pial ee eS A ee 2,78 1,45 —

CREIEGSONOLLOW GGUS... 5 a ee Deg less 3,48 2,78

SO RIAGHOUS 0 26.) ee we 16,67 33,04 16,67

POE BD IDRIS A er ae a 6,94 4,06 ——

emimacmect-—smdll 2 2. te ee 2222 20,87 27,78

Sia Me@ au: a Oe het 6,38 5,56

Kingemmegiums “bo .5 « 5 5 1,39 4,93 5,56

Cue et ee SN eS 5,56 2,90 5,56

very large SD oe a eh ee ea eT a 6,96 13,89

IN =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

S.varilla

. 5 6

x 10 000 yrs BP

Fig. 32. First alternative correlation of the Boom-

plaas A and Die Kelders 1 M.S.A. sequences. BPA

levels OLP to BOLE3 and the DKI 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

FACTOR |

“fo

100

FACTOR 4

10)

FACTOR 2

40 40

30 30

——, ,

~ ¢

S02

‘

\ U

Oy)

om ,

‘

e fe 4

“KN 2Ne /

¢ ‘\ Z ‘

¢ NG

3 4 5) 6 3 4 5 6

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 BOL1 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 “359 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 BOL1 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 (19765), 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

352 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

354 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 355

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 Bil

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 '*O stage 5e 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 lies 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 (19725) 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. ak.

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 lkelihood

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.

REFERENCES

Acocks, J. P. H. 1975. Veld types of South Africa. Mem. bot. Surv. S. Afr. 40: 1-128.

ANDREWS, P., Groves, C. P. & Horne, J. F. M. 1975. Ecology of the lower Tana River flood

plain (Kenya). J. E. Afr. nat. Hist. Soc. & natl Mus. 151: 1-31.

ANSELL, W. F. H. 1960. Mammals of Northern Rhodesia. Lusaka: Government Printer.

ANSELL, W. F. H. & ANSELL, P. D. H. 1973. Mammals of the north-eastern montane areas of

Zambia. Puku 7: 21-69.

Avery, D. M. 1977. Past and present distribution of some rodent and insectivore species in the

southern Cape Province, South Africa: new information. Ann. S. Afr. Mus. 74: 201-209.

Avery, G. 1974. Open station shell midden sites and associated features from the Pearly Beach

area, south-western Cape. S. Afr. archaeol. Bull. 29: 104-114.

Avery, G. 1976. A systematic investigation of open station shell midden sites along the

' south-western Cape coast. Unpublished M.A. thesis, University of Cape Town.

Baba, J. L. & Dees, L. 1975. Accuracy of dates beyond the “C dating limit using the aspartic

acid racemization technique. Nature, Lond. 255: 218-219.

Be, A. W. H. & Duptessy, J. C. 1976. Subtropical convergence fluctuations and Quaternary

climates in the middle latitudes of the Indian Ocean. Science 194: 419-421.

BEHRENSMEYER, A. K. 1975. The taphonomy and paleoecology of Plio-Pleistocene vertebrate

assemblages east of Lake Rudolf, Kenya. Bull. Mus. comp. Zool. 146: 473-578.

366 ANNALS OF THE SOUTH AFRICAN MUSEUM

BIGALKE, R. C. 1978. Mammals. Jn: WerGeR, M. J. A., ed. Biogeography and ecology of

southern Africa. Monogr. biol. 31: 981-1048.

Binrorp, L. R. & BERTRAM, J. B. 1977. Bone frequencies—and attritional processes. In:

BinForD, L. R. ed. For theory building in archaeology: 77-153. New York: Academic

Press.

BODENHEIMER, F. S. 1949. Problems of vole populations in the Middle East. Report on the

population dynamics of the Levant vole (Microtus guentheri D. et A.). Jerusalem: Research

Council of Israel.

Bonp, W., FeRGusON, M. & Forsytu, G. 1980. Small mammals and habitat structure along

altitudinal gradients in the southern Cape mountains. S. Afr. J. Zool. 15: 34-43.

BRAIN, C. K. 1967. Hottentot food remains and their bearing on the interpretation of fossil

bone assemblages. Sci. Pap. Namib Des. Res. Sta. 32: 1-11.

Brain, C. K. 1969. New evidence for climatic change during Middle and Late Stone Age times

in Rhodesia. S. Afr. archaeol. Bull. 24: 127-143.

BraINn, C. K. 1974. The use of microfaunal remains as habitat indicators in the Namib. S. Afr.

archaeol. Soc. Goodwin Ser. 2: 55-60.

Brain, C. K. & Brain, V. 1977. Microfaunal remains from Mirabib: some evidence of

palaeoecological changes in the Namib. Madoqua 10: 285-293.

Brain, C. K. & MEEsTER, J. 1964. Past climatic changes as biological isolating mechanisms in

southern Africa. In: Davis, D. H. S., ed. Ecological studies in southern Africa: 332-340.

The Hague: W. Junk.

Brooks, P. M. 1974. The ecology of the four-striped field mouse, Rhabdomys pumilio

(Sparrman, 1784), with particular reference to a population on the Van Riebeeck Nature

Reserve, Pretoria. Unpublished D.Sc. thesis, University of Pretoria.

Butzer, K. W. 1971. Environmental archaeology. 2nd ed. London: Methuen.

Butzer, K. W. 1973. Geology of Nelson Bay Cave, Robberg, South Africa. S. Afr. archaeol.

Bull. 28: 97-110.

Butzer, K. W. 1974. Reflections on the stability of Holocene environmental zonation in South

Africa. S. Afr. archaeol. Soc. Goodwin Ser. 2: 37-38.

Butzer, K. W. 1978. Sediment stratigraphy of the Middle Stone Age sequence at Klasies River

Mouth, Tzitzikamma coast, South Africa. S$. Afr. archaeol. Bull. 33: 141-151.

Butzer, K. W. & HELGREN, D. M. 1972. Late Cenozoic evolution of the Cape coast between

Knysna and Cape St Francis, South Africa. Quat. Res. 2: 143-169.

Butzer, K. W. Fock, G. J., STUCKENRATH, R & ZiLcH, A. 1973. Palaeohydrology of late

Pleistocene lake, Alexandersfontein, Kimberley, South Africa. Nature, Lond. 243:

328-330.

ButTzer, K. W., STUCKENRATH, R., BRUZEWICZ, A. J. & HELGREN, D. M. 1978. Late Cenozoic

paleoclimates of the Gaap Escarpment, Kalahari margin, South Africa. Quat. Res. 10:

310-339.

CarRTMILL, M. 1967. The Early Pleistocene mammalian microfaunas of sub-Saharan Africa and

their ecological significance. Quaternaria 9: 169-197.

CaTTELL, R. F. 1965a. Factor analysis: an introduction to essentials (I) the purpose and

underlying models. Biometrics 21: 190-215.

CATTELL, R. F. 1965b. Factor analysis: an introduction to essentials (II) the role of factor

analysis in research. Biometrics 21: 405-435.

CHALINE, J. 1972. Les rongeurs du Pléistocéne moyen et supérieur de France. Cahiers de

Paléontologie. Paris: Editions du C.N.R.S.

Cuitty, D. 1960. Population processes in the vole and their relevance to general theory. Can. J.

Zool. 38: 99-113.

CLARK, J., BEERBOWER, J. R. & KieERTZKE, K. K. 1967. Oligocene sedimentation, stratigraphy,

paleoecology and paleoclimatology in the Big Badlands of South Dakota. Fieldiana: geol.

Mem. 5: 1-158.

CiarK, J. D. 1959. The prehistory of southern Africa. Harmondsworth: Penguin.

CiLarK, J. D. 1963. Ecology and culture in the African Pleistocene. S. Afr. J. Sci. 59: 353-366.

Ciason, A. T. 1972. Some remarks on the use and interpretation of archaeozoological data.

Helinium 12: 139-153.

Coe, M. D. & FLANNERY, K. V. 1964. Microenvironments and Mesoamerican prehistory.

Science 143: 650-654.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 367

CoETZEE, C. G. 1963. The prey of owls in the Kruger National Park as indicated by owl pellets

collected during 1960-61. Koedoe 6: 115-125.

CoETZEE, C. G. 1969. The distribution of mammals in the Namib desert and adjoining inland

escarpment. Scient. Pap. Namib Des. Res. Sta. 40: 23-36.

CoETZEE, C. G. 1971. Genus Steatomys. In: MEESTER, J. & SETZER, H. W., eds. The mammals

of Africa: an identification manual. Washington: Smithsonian Institution Press.

CoETZEE, C. G. 1972. The identification of southern African small mammal remains in owl

pellets. Cimbebasia (A) 2: 53-64.

CoETZEE, C. G. 1975. The biology, behaviour and ecology of Mastomys natalensis in southern

Africa. Bull. Wld Hlth Org. 52: 637-644.

CoETZEE, J. A. 1967. Pollen analytical studies in East and southern Africa. Palaeoecol. Afr. 3:

1-146.

Coon, C. S. 1962. The origin of races. London: Jonathan Cape.

CorBeT, G. B. & HAnkKs, J. 1968. A revision of the elephant-shrews, family Macroscelididae.

Bull. Br. Mus. nat. Hist. (Zool.) 16: 47-111.

CowciLL, G. L. 1968. Archaeological applications of factor, cluster and proximity analysis.

Amer. Antiquity 33: 367-375.

CRAIGHEAD, J. J. & CRAIGHEAD, F. C. 1956. Hawks, owls and wildlife. Harrisburg: Stackpole.

DANSGAARD, W., JOHNSEN, S. J., CLAUSEN, H. B. & LaNGway, C. C. 1971. Climatic record

revealed by the Camp Century ice core. Jn: TUREKIAN, K. K. ed. The late Cenozoic glacial

ages: 37-56. New Haven: Yale University Press.

Davies, O. 1971. Pleistocene shorelines in the southern and south-eastern Cape Province

(Part 1). Ann. Natal Mus. 21: 183-223.

Davies, O. 1972. Pleistocene shorelines in the southern and south-eastern Cape Province

(Part 2). Ann. Natal Mus. 21: 225-279.

Davis, D. H. S. 1958. Notes on small mammals in the Kalahari Gemsbok National Park, with

special reference to those preyed upon by barn owls. Koedoe 1: 184-188.

Davis, D. H. S. 1959. The barn owl’s contribution to ecology and palaeoecology. Proc. Ist

Pan-Afr. Ornith. Congr. Ostrich suppl. 3: 144-153.

Davis, D. H. S. 1962. Distribution patterns of southern African Muridae, with notes on some

of their fossil antecedents. Ann. Cape prov. Mus. 2: 56-76.

Davis, D. H. S. 1965. Classification problems of African Muridae. Zool. afr. 1: 121-145.

Davis, D. H. S. 1966. The small rodents of the Transvaal: some taxonomic, biogeographic,

economic and health problems. Fauna Flora, Pretoria 17: 4-12.

Davis, D. H. S. 1974. The distribution of some small southern African mammals (Mammalia:

Insectivora, Rodentia). Ann. Transv. Mus. 29: 135-184.

Davis, R. M. 1973. The ecology and life history of the vlei rat, Otomys irroratus (Brants,

1827), on the Van Riebeeck Nature Reserve, Pretoria. Unpublished D.Sc. thesis, Univer-

sity of Pretoria.

Deacon, H. J. 1969. Melkhoutboom Cave, Alexandria District, Cape Province: a report on the

1967 excavation. Ann. Cape prov. Mus. (nat. Hist.) 6: 141-169.

Deacon, H. J. 1972. A review of the post-Pleistocene in South Africa. S. Afr. archaeol. Soc.

Goodwin Ser. 1: 26-45.

Deacon, H. J. 1976. Where hunters gathered. S. Afr. archaeol. Soc. monogr. Ser. 1: 1-232.

Deacon, H. J. 1979. Excavations at Boomplaas Cave—a sequence through the Upper Pleisto-

cene and Holocene in South Africa. Wid Archaeol. 10: 241-257.

Deacon, H. J. & Brooker, M. 1976. The Holocene and Upper Pleistocene sequence in the

southern Cape. In: Proc. sth Afr. Soc. Quat. Res., Ann. S. Afr. Mus. 71: 203-214.

Deacon, H. J., DEAcon, J. & BROOKER, M. 1976. Four painted stones from Boomplaas Cave,

Oudtshoorn District. $. Afr. archaeol. Bull. 31: 141-145.

Deacon, H. J., DEACON, J., BROOKER, M. & Witson, M. L. 1978. The evidence for herding at

Boomplaas Cave in the southern Cape, South Africa. S. Afr. archaeol. Bull, 33: 39-65.

Deacon, J. 1972. Wilton: an assessment after fifty years. S. Afr. archaeol. Bull. 27: 10-48.

Deacon, J. 1978. Changing patterns in the late Pleistocene/early Holocene prehistory of

southern Africa as seen from the Nelson Bay Cave stone artifact sequence. Quat. Res. 10:

84-111.

DEAN, W. R. J. 1973. Age distribution of Praomys natalensis in Tyto alba pellets. Zool. afr. 8:

140.

368 ANNALS OF THE SOUTH AFRICAN MUSEUM

Dean, W. R. J. 1977. The ecology of owls at Barberspan, Transvaal. Proc. Symp. Afr.

predatory Birds: 25-45.

DE GraarFF, G. 1960. A preliminary investigation of the mammalian microfauna in Pleistocene

deposits of caves in the Transvaal System. Palaeont. afr. 7: 59-118.

De GRAAFF, G. 1974. Notes on the occurrence of rodents in South African national parks.

Koedoe 17: 173-183.

De Graaff, G. 1981. The rodents of South Africa. Durban: Butterworth.

De GraarF, G & NEL, J. A. J. 1970. Notes on the smaller mammals of the eastern Cape

national parks. Koedoe 13: 147-149.

DELANEY, M. J. 1972. The ecology of small rodents in tropical Africa. Mamm. Rev. 2: 1-42.

DELANEY, M. J. & NEAL, B. R. 1966. A revue of the Muridae (Order Rodentia) of Uganda.

Bull. Br. Mus. nat. Hist. (Zool.) 13: 295-355.

DE MENESES CABRAL, J. C. 1966. Some new data on Angolan Muridae. Zool. afr. 2: 193-203.

DINGLE, R. V. & RoGers, J. 1972. Pleistocene palaeogeography of the Agulhas Bank Trans. R.

Soc. S. Afr. 40: 155-165.

Doran, J. E. & Hopson, F. R. 1975. Mathematics and computers in archaeology. Edinburgh:

Edinburgh University Press.

Dorst, J. & DANDELoT, P. 1970. A field guide to the larger mammals of Africa. London:

Collins.

EMILIANI, C. 1955. Pleistocene temperatures. J. Geol. 63: 538-578.

EMILIANI, C. 1966. Paleotemperature analysis of Caribbean cores P6304-8, P6304-9 and a

generalised temperature curve for the last 425 000 years. J. Geol. 74: 109-126.

EMILIANI, C. 1972. Quaternary paleotemperatures and the duration of high-temperature

intervals. Science 178: 398-401.

Estes, R. D. 1971. Social organisation of the African Bovidae. Jn: Geist, V. & WALTHER, F.,

eds. The behaviour of ungulates and its relation to management. IUCN Publs 24: 166-205.

Ewer, R. F. 1967. The fossil hyaenids of Africa—a reappraisal. In: BisHop, W. W. & CLARK,

J. D., eds. Background to evolution in Africa: 109-123. Chicago: University of Chicago

Press.

FaGAN, B. M. 1960. The Glentyre shelter and Oakhurst re-examined. S$. Afr. archaeol. Bull.

15: 80-94.

FAIRBRIDGE, R. W. 1961. Eustatic changes in sea-level. Jn: AHRENS, L. H., PREss, F.,

RANKAMA, K. & Runcorn, S. K., eds. Physics and chemistry of the earth 4: 99-185.

London: Pergamon Press.

FAIRBRIDGE, R. W. 1971. Quaternary shoreline problems at INQUA 1969. Quaternaria 15:

1-17.

FeRRAR, A. A. & WALKER, B. H. 1974. An analysis of herbivore/habitat relationships in Kyle

National Park, Rhodesia. J. sth Afr. Wildl. Mgmt Ass. 4: 137-147.

FLANNERY, K. V. 1967. Vertebrate fauna and hunting patterns. Jn: Byers, D. S. ed. The

prehistory of the Tehuacan Valley. 1. Environment and subsistence: 132-177. Austin &

London: University of Texas Press.

FLANNERY, K. V. 1968. Archeological systems theory and early Mesoamerica. In: MEGGARS,

B. J., ed. Anthropological archeology in the Americas: 67-88. Washington: Anthropologi-

cal Society of Washington.

FLIntT, R. F. 1976. Physical evidence of Quaternary climatic change. Quat. Res. 6: 519-528.

GuvE, D. E. 1967. Prey taken by the barn owl in England and Wales. Bird Study 14: 169-183.

Goopwin, A. J. H. & VAN RieEr Lowe, C. 1929. The Stone Age cultures of South Africa. Ann.

S. Afr. Mus. 27: 1-289.

Grayson, D. K. 1978. Minimum numbers and sample size in vertebrate faunal analysis. Amer.

Antiquity 43: 53-65.

GRINDLEY, J., SIEGFRIED, W. R. & VERNON, C. J. 1973. Diet of the barn owl in the Cape

Province. Ostrich 44: 266-267.

GUuILCHER, A. 1969. Pleistocene and Holocene sea level changes. Earth-Sci. Rev. 5: 69-97.

GuiLpay, J. E. 1971. The Pleistocene history of the Appalachian mammal fauna. Jn: HOLr,

P. C., ed. The distributional history of the biota of the southern Appalachians. III.

Vertebrates. Virginia Polytech. Inst. State Univ. Res. Div. Monogr. 4: 233-262.

Hanney, P. 1962. An aid to the identification of skulls of some small Nyasaland mammals.

Rep. Bull. Nyasaland Mus. 1962: 34-38.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS 369

Hanney, P. 1963. Observations upon the food of the barn owl (Tyto alba) in southern

Nyasaland, with a method of ascertaining population dynamics of rodent prey. Ann. Mag.

nat. Hist. (13) 6: 305-313.

Hanney, P. 1965. The Muridae of Malawi (Africa: Nyasaland). J. Zool. 146: 577-633.

HARPENDING, H. & Davis, H. 1977. Some implications for hunter-gatherer ecology derived

from the spatial structure of resources. Wid Archaeol. 8: 275-286.

Hays, J. D., Lozano, J. A., SHACKLETON, N. J. & IRvING, G. 1976. Reconstruction of the

Atlantic and western Indian Ocean sectors of the 18 000 B.P. Antarctic Ocean. Jn: CLINE,

R. M. & Hays, J. D., eds. Investigation of late Quaternary paleooceanography and

paleoclimatology. Geol. Soc. Amer. Mem. 145: 337-372.

HENDEY, Q. B. 1974. The late Cenozoic Carnivora of the south-western Cape Province. Ann.

S. Afr. Mus. 63: 1-369.

Hicuam, C. F. W. 1967. Faunal sampling and economic prehistory. Z. Sdugetierk. 33: 297-305.

Hirt, J. N. 1972. The methodological debate in contemporary archaeology: a model. Jn:

CriarK, D. L., ed. Models in archaeology: 61-107. London: Methuen.

Hirst, S. M. 1975. Ungulate—-habitat relationships in a South African woodland/savanna

ecosystem. Wildl. Monogr. 44: 1-60.

HOFMANN, R. R. & STEWART, D. R. M. 1972. Grazer or browser: a classification based on the

stomach-structure and feeding habits of East African ruminants. Mammalia 36: 226-240.

Honer, M. R. 1963. Observations on the barn owl (Tyto alba guttata) in the Netherlands in

relation to its ecology and population fluctuations. Ardea 51: 158-195.

HusBparpb, C. A. 1972. Observations on the life histories and behaviour of some small rodents

from Tanzania. Zool. afr. 7: 419-449.

HuFFAKER, C. B. & MESSENGER, P. S. 1964. The concept and significance of natural control. Jn:

DeEBacu, P., ed. Biological control of insects and weeds: 74-117. London: Chapman &

Hall.

Impric, J. & Kipp, N. G. 1971. A new micropaleontological method for quantitative paleo-

climatology: application to a late Pleistocene Caribbean core. Jn: TUREKIAN, K. K., ed.

The late Cenozoic glacial ages: 71-183. New Haven: Yale University Press.

ImpriE, J.,. VAN Donck, J. & Kipp, N. G. 1973. Paleoclimatic investigation of a late Pleistocene

Caribbean deep-sea core: comparison of isotopic and faunal methods. Quat. Res. 3: 10-38.

JARMAN, P. J. 1974. The social organisation of antelope in relation to their ecology. Behaviour

48: 215-267.

Jarvis, J. U. M. & Davin, J. n.d. Population ecology of the striped field mouse Rhabdomys

pumilio on the Cape Flats, including their dependence on invasive Acacias. In: Fynbos

research project: first annual research meeting (unedited abstracts). (Duplicated. )

JOHNSEN, S. J., DANSGAARD, W., CLAUSEN, H. B. & LANGway, C. C. 1972. Oxygen isotope

profiles through the Antarctic and Greenland ice sheets. Nature, Lond. 235: 429-434.

Kim, J. 1975. Factor analysis. In: Nie, N. H., Hutt, C. H., JENxkins, J. G., STEINBRENNER, K.

& Brent, D. H., eds. SPSS: statistical package for the social sciences: 468-514. New York:

McGraw-Hill.

Kine, L. C. 1963. South African scenery. Edinburgh: Oliver & Boyd.

Kincpon, J. 1974. East African mammals IIA & ITB. London & New York: Academic Press.

KLEIN, R. G. 1972a. Preliminary report on the July through September 1970 excavations at

Nelson Bay Cave, Plettenberg Bay (Cape Province, South Africa). Palaeoecol. Afr. 6:

177-208.

KieIn, R. G. 19726. The late Quaternary mammalian fauna of Nelson Bay Cave (Cape

Province, South Africa): its implications for megafaunal extinctions and for environmental

and cultural change. Quat. Res. 2: 135-142.

KLEIN, R. G. 1974a. Environment and subsistence of prehistoric man in the southern Cape

Province, South Africa. Wild Archaeol. 5: 249-284.

_K etn, R. G. 1974b. On the taxonomic status, distribution and ecology of the blue antelope,

Hippotragus leucophaeus (Pallas, 1766). Ann. S. Afr. Mus. 65: 99-143.

KLEIN, R. G. 1975. Middle Stone Age man—animal relationships in southern Africa: evidence

from Die Kelders and Klasies River Mouth. Science 190: 265-267.

KLEIN, R. G. 1976. The mammalian fauna of the Klasies River Mouth sites, southern Cape

Province, South Africa. S. Afr. archaeol. Bull. 31: 75-98.

KLEIN, R. G. 1977. The ecology of early man in southern Africa. Science 197: 115-126.

370 ANNALS OF THE SOUTH AFRICAN MUSEUM

KieIn, R. G. 1978. A preliminary report on the larger mammals from the Boomplaas Stone

Age site, Cango Valley, Oudtshoorn District, South Africa. S. Afr. archaeol. Bull. 33:

66-75.

KLEIN, R. G. 1979. Stone Age exploitation of animals in southern Africa. Amer. Sci. 67:

151-160.

KieIn, R. G. 1981. Later Stone Age subsistence at Byeneskranskop Cave, South Africa. In:

HarpinG, R. S. O. & TELEKI, G., eds. Omnivorous primates: gathering and hunting in

human evolution: 166-190. New York: Columbia University Press.

Krome, H. 1962. The influence of climate and weather on the mean density level, the

fluctuations and the regulation of animal populations. Arch. Néerl. Zool. 15: 68-109.

Knox, E. 1976. Upper alveolar patterns of some Muridae in Queensland and Papua New

Guinea. Mem. Queensland Mus. 17: 457-460.

Kowatsk!, K. 1971. The biostratigraphy and paleoecology of Late Cenozoic mammals of

Europe and Asia. Jn: TUREKIAN, K. K., ed. The late Cenozoic glacial ages: 465-477. New

Haven: Yale University Press.

Kress, C. J. 1972. Ecology. New York: Harper & Row

KruGEr, F. J. 1977. Ecological reserves in the Cape fynbos: towards a strategy for conserva-

tion. S. Afr. J. Sci. 73: 81-85.

KruGer, F. J. 1979. South African heathlands. Jn: Specut, R. L.,. ed. Ecosystems of the world.

9a. Heathlands and related shrublands. Descriptive studies: 19-80. Amsterdam: Elsevier.

KuKLaA, G. J. 1975. Loess stratigraphy of central Europe. Jn: Burzer, K. W. & Isaac, G. LL,

eds. After the australopithecines: 99-188. The Hague & Paris: Mouton.

Kuk1a, G. J. 1977. Pleistocene land-sea correlations. 1. Europe. Earth-Sci. Rev. 13: 307-374.

Lamprey, H. F. 1963. Ecological separation of the large mammal species in the Tarangire

Game Reserve, Tanganyika. E. Afr. Wildl. J. 1: 63-92.

LIVINGSTONE, D. A. 1975. Late Quaternary climatic change in Africa. A. Rev. Ecol. Syst. 6:.

249-280. ;

Luz, B. 1973. Stratigraphic and paleoclimatic analysis of Late Pleistocene tropical southeast

Pacific cores (with appendix by N. J. Shackleton). Quat. Res. 3: 56-72.

Luz, B. 1977. Late Pleistocene paleoclimates of the south Pacific based on statistical analysis of

planktonic foraminifers. Palaeogeogr. Palaeoclimatol. Palaeoecol. 22: 61-78.

Lyncu, C. D. 1975. The distribution of mammals in the Orange Free State, South Africa.

Navors. nas. Mus. Bloemfontein 3: 109-139.

MANGERUD, J., S@NSTEGAARD, E. & SesRuP, H.-P. 1979. Correlation of the Eemian (inter-

glacial) Stage and the deep-sea oxygen isotope stratigraphy. Nature, Lond. 277: 189-192.

Martin, A. R. H. 1962. Evidence relating the Quaternary history of the Wilderness lakes.

Trans. Proc. geol. Soc. S. Afr. 65: 19-42.

Martin, A. R. H. 1968. Pollen analysis of Groenvlei lake sediments, Knysna (South Africa).

Rev. Palaeobotan. Palynol. 7: 107-144.

Martin, A. R. H. 1969. Final report on the palynology of Groenvlei, Knysna, South Africa.

Palaeoecol. Afr. 4: 190-191.

Mazel, A. D. & ParRKINGTON, J. E. 1978. Sandy Bay revisited: variability among Late Stone

Age tools. S. Afr. J. Sci. 74: 381-382.

McLacuHian, G. R. & LiversipGe, R. 1970. Roberts birds of South Africa. Johannesburg:

Trustees of the John Voelcker Bird Book Fund.

MEESTER, J. 1955. A mammal survey of the S.A. Lombard Nature Reserve (1954). Fauna

Flora, Pretoria 6: 45-57.

Meester, J. 1958. Variation in the shrew genus Myosorex in southern Africa. J. Mammal. 39:

325-339.

MeEEsTER, J. 1961. A taxonomic revision of southern African Crocidura (Mammalia: Insecti-

vora). Ann. Mag. nat. Hist. 4: 561-571.

Meester, J. 1962. The distribution of Crocidura Wrangler in southern Africa. Ann. Cape prov.

Mus. 2: 77-84.

MEEsTER, J. 1963. A systematic revision of the shrew genus Crocidura in southern Africa.

Transv. Mus. Mem. 13: 1-127. —

MEESTER, J. & Setzer, H. W., eds. 1971. The mammals of Africa: an identification manual.

Washington: Smithsonian Institution Press.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS ah

MISONNE, X. 1969. African and Indo-Australian Muridae: evolutionary trends. Ann. Mus. r.

Afr. cent. (in-8°) (Sci. zool.) 172: 1-219.

MISONNE, X. 1971. Rodentia. Jn: MEESTER, J. & SETZER, H. W., eds. The mammals of Africa:

an identification manual. Washington: Smithsonian Institution Press.

MorreTr, R. O. & Deacon, H. J. 1977. The flora and vegetation in the surrounds of

Boomplaas Cave, Cango Valley. S. Afr. archaeol. Bull. 32: 127-145.

MonrTAGGIONI, L. 1976. Holocene submergence on Réunion Island (Indian Ocean). In: Proc.

sth Afr. Soc. Quat. Res., Ann. S. Afr. Mus. 71: 69-75.

Morner, N.-A. 1978. Palaeogeoidal changes and palaeoecological changes in Africa with

respect to real and apparent palaeoclimatic changes. Palaeoecol. Afr. 10: 1-12.

Neal, B. R. 1970. The habitat distribution and activity of a rodent population in western

Uganda, with particular reference to the effects of burning. Rev. Zool. Bot. afr. 81: 29-50.

NEAL, B. R. & Cock, A. G. 1969. An analysis of the selection of small African mammals by

two break-back traps. J. Zool. 158: 335-340.

Nex, J. A. J. & Pretorius, Jj. J. L. 1971. A note on the smaller mammals of the Mountain

Zebra National Park. Koedoe 14: 99-110.

NEL, J. A. J. & RAuTensacu, I. L. 1975. Habitat use and community structure of rodents in

the southern Kalahari. Mammalia 39: 9-29.

Nir, N. H., Hutz, C. H., Jenkins, J. G., STEINBRENNER, K. & Bent, D. H. 1975. SPSS:

statistical package for the social sciences. New York: McGraw-Hill.

Ovum, E. P. 1971. Fundamentals of ecology. Philadelphia: W. B. Saunders.

PARKINGTON, J. E. 1972. Seasonal mobility in the Later Stone Age. Afr. Stud. 31: 223-243.

PARKINGTON, J. E. 1976. Coastal settlement between the mouths of the Berg and Olifants

Rivers, Cape Province. S. Afr. archaeol. Bull. 31: 127-140.

PATERSON, W. S. B., KOERNER, R. M., FISHER, D., JOHNSEN, S. J., CLAUSEN, H. B.,

DANSGAARD, W., BUCHER, P. & OrSCHGER, H. 1977. An oxygen isotope climatic record

from the Devon Island ice cap, Arctic Canada. Nature, Lond. 266: 508-511.

PIENAAR, U. DE V. 1964. The small mammals of the Kruger National Park: a systematic list and

zoogeography. Koedoe 7: 1-25.

RAUTENBACH, I. L. 1971. Notes on the small mammals of the Kalahari Gemsbok National Park.

Koedoe 14: 137-144.

RAUTENBACH, I. L. 1976. A survey of the mammals occurring in the Golden Gate Highlands

National Park. Koedoe 19: 133-144.

RAUTENBACH, I. L. & NEL, J. A. J. 1975. Further records of smaller mammals of the Kalahari

Gemsbok National Park. Koedoe 18: 195-198.

Roserts, A. 1951. The mammals of South Africa. Johannesburg: Trustees of ‘The mammals of

South Africa’ Book Fund.

RosENZWEIG, M. L. & Winaxvur, J. 1969. Population ecology of desert communities: habitats

and environmental complexity. Ecology 50: 558-572.

RosEvEAR, D. R. 1969. The rodents of West Africa. London: Trustees of the British Museum

(Natural History).

Roussouw, P. J., Meyer, E. I., MULDER, M. P. & STOcKEN, C. G. 1964. Die geologie van die

Swartberge, die Kangovallei en die omgewing van Prins Albert, K.P. Pretoria: Dept. of

Mines, Geological Survey.

RuMMEL, R. J. 1967. Understanding factor analysis. J. Conflict Resolution 11: 444-480.

SaInT Girons, M.-C. 1977. Morphologie végétale et répartition des Mammiféres. Bijdr. Dierk.

47: 120-130.

Sampson, C. G. 1974. The Stone Age archaeology of southern Africa. New York & London:

Academic Press.

SANCETTA, C. ImprRie, J. & Kipp N. G. 1973. Climatic record of the past 130 000 years in north

Atlantic deep-sea core V23-—82: correlation with the terrestrial record. Quat. Res. 3:

110-116.

SCHALKE, H. J. W. G. 1973. The Upper Quaternary of the Cape Flats area (Cape Province,

South Africa) Scripta geol. 15: 1-57.

ScHaAPERA, I. 1930. The Khoisan peoples of South Africa. Bushmen and Hottentots. London:

George Routledge & Sons.

ScHRIRE, C. 1962. Oakhurst: a re-examination and vindication. S. Afr. archaeol. Bull. 17:

181-195.

372 ANNALS OF THE SOUTH AFRICAN MUSEUM

ScHuLz, K. H. 1953. The control of veld rodents responsible for damage to pine forests in the

eastern Transvaal. Public Health (Johannesburg) 17: 402-465.

SCHULZE, B. R. 1965. Climate of South Africa. Part 8: general survey. Pretoria: Government

Printer.

SCHWEITZER, F. R. 1970. A preliminary report of excavations at a cave at Die Kelders. S. Afr.

archaeol. Bull 25: 136-138.

SCHWEITZER, F. R. & Scott, K. 1973. Early occurrence of domestic sheep in sub-Saharan

Africa. Nature, Lond. 241: 547.

SCHWEITZER, F. R. & Wixson, M. L. 1978. A preliminary report on excavations at Byneskrans-

kop, Bredasdorp District, Cape. S. Afr. archaeol. Bull. 33: 134-140.

ScLATER, W. L. 1900-1. The mammals of South Africa. London: R. H. Porter.

Scott, L. & VoGEL, J. C. 1978. Pollen analysis of the thermal spring deposit at Wonderkrater

(Transvaal, South Africa). Palaeoecol. Afr. 10: 155-162.

SERVICE, E. R. 1966. The hunters. Englewood Cliffs: Prentice-Hall.

SHACKLETON, N. J. 1973. Oxygen isotope analysis as a means of determining season of

occupation of prehistoric midden sites. Archaeometry 15: 133-141.

SHACKLETON, N. J. 1975. The stratigraphic record of deep-sea cores and its implications for the

assessment of glacials, interglacials, stadials and interstadials in the mid-Pleistocene. Jn:

BuTzer, K. W. & Isaac, G. LL., eds. After the australopithecines: 1-24. The Hague &

Paris: Mouton.

SHACKLETON, N. J. & MATTHEws, R. K. 1977. Oxygen isotope stratigraphy of late Pleistocene

coral terraces in Barbados. Nature, Lond. 268: 618-620.

SHACKLETON, N. J. & OppykeE, N. D. 1973. Oxygen isotope and paleomagnetic stratigraphy of

equatorial core V28-238: oxygen isotope temperatures and ice volumes on a 10° and 10°

scale. Quat. Res. 3: 39-55.

SHACKLETON, N. J. & OppykeE, N. D. 1976. Oxygen-isotope and paleomagnetic stratigraphy of

Pacific core V28—239 late Pliocene to latest Pleistocene. Jn: CLINE, R. M. & Hays, J. D.,

eds. Investigation of late Quaternary paleooceanography and paleoclimatology. Geol. Soc.

Amer. Mem. 145: 449-464.

SHEPPE, W. A. 1973. Notes on Zambian rodents and shrews. Puku 7: 167-190.

SHEPPE, W. A. & OspBorn_e, T. 1971. Patterns of use of a flood plain by Zambian mammals.

Ecol. monogr. 41: 179-205.

SHORTRIDGE, G. C. 1934. The mammals of South West Africa. London: William Heinemann.

SHORTRIDGE, G. C. 1942. Field notes on the first and second expeditions of the Cape museums’

mammal survey of the Cape Province: descriptions of some new subgenera and subspecies.

Ann. S. Afr. Mus. 36: 27-100.

Simpson, G. G., RoE, A. & LEwontin, R. C. 1960. Quantitative zoology. New York: Harcourt

Brace.

SMITHERS, R. H. N. 1971. The mammals of Botswana. Natl Mus. Rhodesia Mem. 4: 1-340.

Smuts, J. 1832. Dissertatio zoologica, ennumerationem mammalium capensium. Leiden: J. C.

Cyfveer.

SOUTHERN, H. N. 1954. Tawny owls and their prey. /bis 96: 348-410.

SPARKS, J. & Soper, T. 1972. Owls. Their natural and unnatural history. Newton Abbot: David

& Charles.

STEYN, P. & TREDGOLD, D. 1977. Observations on the Cape eagle owl. Bokmakierie 29: 31-42.

Stuart, C. T. 1976. Diet of the black backed jackal Canis mesomelas in the Namib Desert,

South West Africa. Zool. afr. 11: 193-205.

STuarT, C. T. 1977. Analysis of Felis libyca and Genetta genetta scats from the central Namib

Desert, South West Africa. Zool. afr. 12: 239-241.

Stuart, C. T. n.d. Preliminary report on a rodent and insectivore survey in the Jonkershoek

Valley, Stellenbosch. Unpublished report, Cape Provincial Administration.

SWANEPOEL, P. 1975. Small mammals of the Addo Elephant National Park. Koedoe 18:

103-137.

SycH, L. 1965. Fossil Leporidae from the Pliocene and Pleistocene of Poland. Acta zool.

Cracov. 10: 1.

TaLMA, A. S., VoGEL, J. C. & PartipGe, T. C. 1974. Isotopic contents of some Transvaal

speleothems and their palaeoclimatic significance. S. Afr. J. Sci. 70: 135-140.

TANKARD, A. J. 1976a. The stratigraphy of a coastal cave and its palaeoclimatic significance.

Palaeoecol. Afr. 9: 151-159.

MICROMAMMALS AS PALAEOENVIRONMENTAL INDICATORS Bes.

TANKARD, A. J. 1976b. Cenozoic sea-level changes: a discussion. Jn: Proc. sth Afr. Soc. Quat.

Res., Ann S. Afr. Mus. 71: 1-17.

TANKARD, A. J. & SCHWEITZER, F. R. 1974. The geology of Die Kelders Cave and environs: a

palaeoenvironmental study. S. Afr. J. Sci. 70: 365-369.

TANKARD, A. J. & SCHWEITZER, F. R. 1976. Textural analysis of cave sediments: Die Kelders,

Cape Province, South Africa. Jn: Davipson, D. A. & SHACKLEY, M. L., eds. Geoarchae-

ology: 289-316. London: Duckworth.

Taytor, H. C. 1961. Ecological account of a remnant of coastal forest near Stanford, Cape

Province. J. S. Afr. Bot. 27: 153-165.

TayLor, H. C. 1978. Capensis. In: Wercer, M. J. A., ed. Biogeography and ecology of

southern Africa. Monogr. Biol. 31: 171-229.

TcCHERNOV, E. 1968. Succession of rodent faunas during the Upper Pleistocene of Israel.

Mammalia depicta. Hamburg: Verlag Paul Parey.

TcHERNOV, E. 1975. Rodent faunas and environmental change in the Pleistocene of Israel. Jn:

PrakasH, I. & Guosu, P. K., eds. Rodents in desert environments: 331-362. The Hague:

W. Junk.

Tuomas, O. & SCHWANN, H. 1904. The Rudd exploration of South Africa. I-British Namaqua-

land. Proc. zool. Soc. Lond. 1904: 171-183.

Tuomas, O. & SCHWANN, H. 1905a. The Rudd exploration of South Africa. I—Wakkerstroom,

S.E. Transvaal. Proc. zool. Soc. Lond. 1905: 129-138.

THomas, O. & SCHWANN, H. 1905b. The Rudd exploration of South Africa. HJ—Zululand.

Proc. zool. Soc. Lond. 1905: 254-276.

Tuomas, O. & ScHWANN, H. 1906a. The Rudd exploration of South Africa. [V—Knysna. Proc.

zool. Soc. Lond. 1906: 159-168.

Tuomas, O. & SCHWANN, H. 1906b. The Rudd a of South Africa. V-N.E. Transvaal.

Proc. zool. Soc. Lond. 1906: 575-591.

TriGGER, B. G. 1971. Archaeology and ecology. W/d Archarol: 2: 321-336.

TRUSWELL, J. F. 1971. The geological evolution of South Africa. Cape Town: Purnell.

VAN DER HAMMEN, T., WusmstTrA, T. A. & ZaGwun, W. H. 1971. The floral record of the late

Cenozoic of Europe. Jn: TUREKIAN, K. K., ed. The late Cenozoic glacial ages: 390-424.

New Haven: Yale University Press.

VAN ZINDEREN BAKKER, E. M. 1957. A pollen analytical investigation of the Florisbad deposits

(South Africa). In: Clark, J. D., ed. Third Pan-African Congress on Prehistory, Living-

stone 1955: 56-67. London: Chatto & Windus.

VAN ZINDEREN BAKKER, E. M. 1976. The evolution of Late-Quaternary palaeoclimates of

southern Africa. Palaeoecol. Afr. 9: 160-202.

VAN ZINDEREN BAKKER, E. M. & Butzer, K. W. 1973. Quaternary environmental changes in

southern Africa. Soil Sci. 116: 236-248.

VERNON, C. J. 1972. An analysis of owl pellets collected in southern Africa. Ostrich 43:

109-124.

VESEY-FITZGERALD, D. F. 1964. Mammals of the Rukwa Valley. Tanganyika Notes & Records

62: 61-72.

VESEY-FITZGERALD, D. F. 1966. The habits and habitats of small rodents in the Congo River

catchment region of Zambia and Tanzania. Zool. afr. 2: 111-122.

VINCENT, E. 1972. Climatic change at the Pleistocene—Holocene boundary in the southwestern

Indian Ocean. Palaeoecol. Afr. 6: 45-54.

WELLINGTON, J. H. 1955. Southern Africa—a geographical study. I. Physical geography.

Cambridge: Cambridge University Press.

WenrcER, M. J. A. 1978. The Karoo—Namib Region. Jn: WerGER, M. J. A., ed. Biogeography

and ecology of southern Africa. Monogr. biol. 31: 231-299.

White, F. 1978. The Afromontane Region. Jn: WerGeR, M. J. A., ed. Biogeography and

ecology of southern Africa. Monogr. biol. 31: 463-513.

WinGate, L. R. & MEEstTER, J. 1977. A field test of six types of live-trap for African rodents.

Zool. afr. 12: 215-223.

WOILLARD, G. M. 1978a. Grande Pile peat bog: a continuous pollen record for the last 140 000

years. Quat. Res. 9: 1-21.

WOILLARD, G. M. 19785. Végétation et climat des derniers 140 000 ans dans la tourbiére de la

Grande Pile (N.E. France). Palaeoecol. afr. 10: 125—134.

374 ANNALS OF THE SOUTH AFRICAN MUSEUM

Wyner, J. J. & SINGER, R. 1972. Middle Stone Age occupational settlements on the Tsitsikama

coast, eastern Cape Province, South Africa. In: Ucko, P. J., TRINGHAM, R. & DIMBLEBY,

G. W., eds. Man, settlement and urbanism: 207-210. London: Duckworth.

YELLEN, J. E. 1977. Long-term hunter-gatherer adaptation to desert environments: a biogeo-

graphical perspective. Wid Archaeol. 8: 262-274.

YELLEN, J. E. & LEE, R. B. 1976. The Dobe-/Du/da environment. Jn: LEE, R. B., & DE VoreE,

I., eds. Kalahari hunter—gatherers: 27-46. Cambridge & London: Harvard University Press.